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Quaestiones
Entomologicae
A periodica! record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 23
NUMBER 1
WINTER 1987
Publication of Quaestiones Entomologicae was started in 1965 as part of a
memorial project for Professor E. H. Strickland, the founder of the
Department of Entomology at The University of Alberta in Edmonton in 1922.
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length. However, shorter papers about insects in the Prairie Provinces of
Canada are acceptable. Page charges are normally levied, the rate determined
by printer’s charges. For information about current page charges, consult the
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Published quarterly by:
Department of Entomology
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Issued March 1987
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 23 1987
CONTENTS
Nimmo-The adult Arctopsychidae and Hydropsychidae (Trichoptera) of Canada and adjacent
United States 1
Book Review-Campos-Ortega, J.A. and V. Hartenstein. 1985. The Embryonic Development of
Drosophila melanogaster 191
Book Review-Tauber, M.J., C.A. Tauber and S. Masaki. 1986. Seasonal Adaptations of
Insects 194
Sperling-Evolution of the Papilio machaon species group in western Canada (Lepidoptera:
Papilionidae) 1 98
Clark-Revision of the Anthonomus Subgenus Anthomorphus Weise (Coleoptera:
Curculionidae) 317
Book Review-Mitchell, Andrew W. 1986. The Enchanted Canopy: A Journey to the Last
Unexplored Frontier, The Roofs of the World’s Rainforests 365
Book Notice-Belles, X. 1985. Systematica, filogenia y biogeografia de la subfamilia Gibbiinae
(Coleoptera: Ptinidae) 368
Book Review-D’Alberta, B. 1987. Sphingidae Mundi 369
Book Notice-Cox, J.M. 1987. Pseudococcidae (Insecta: Hemiptera) 371
Craig-A Taxonomic Account of the Black Flies (Diptera: Simuliidae) of the Society Islands -
Tahiti, Moorea and Raiatea 372
Anderson-Systematics, phylogeny and biogeography of New World weevils traditionally of the
tribe Cleonini (Coleoptera: Curculionidae; Cleoninae) 431
Book Notices 711
Editor’s Acknowledgements and Comments 713
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 23 Number! 1987
CONTENTS
Nimmo-The adult Arctopsychidae and Hydropsychidae (Trichoptera) of Canada
and adjacent United States 1
Book Review-Campos-Ortega, J.A. and V. Hartenstein. 1985. The Embryonic
Development of Drosophila melanogaster 191
Book Review-Tauber, M.J., C.A. Tauber and S. Masaki. 1986. Seasonal
Adaptations of Insects 194
THE ADULT ARCTOPSYCHIDAE AND HYDROPSYCHIDAE (TRICHOPTERA) OF
CANADA AND ADJACENT UNITED STATES
Andrew P. Nimmo
Department of Entomology
University of Alberta
Edmonton, Alberta, T6G 2E3 Quaestiones Entomologicae
CANADA 23:1-189 1987
ABSTRACT
Of the six species of Arctopsychidae here reported from Canada and adjacent States of the
United States, three belong to each of Arctopsyche McLachlan and Parapsyche Betten. Of 72
species of Hydropsychidae, 24 belong to Cheumatopsyche Wallengren, 42 to Hydropsyche
Pictet, three to Macrostemum Kolenati, and one each to Potamyia Banks, Diplectrona
Westwood, and Aphropsyche Ross.
Keys are provided (for males, and females where possible ) to genera and species. For each
species the habitus is described in some detail, with diagnostic statements for the genitalia.
Also included are brief statements about way of life and known distribution. Distributions are
mapped, and genitalia are illustrated.
RESUME
Soixante et dix-huit especes d’ Arctopsychidae et d’ Hydropsychidae (Trichoptera) sont mentionnees pour le Canada
et les etats frontaliers des Etats-Unis, representant les genres suivants: Archtopsyche MeLachlan (3), Parapsyche Betten
(Archtopsychidae) (3), Cheumatopsyche Wallengren (24), Hydropsyche Pictet (42), Macrostemum Kolenati (3), Potomyia
Banks (1), Diplectrona Westwood (I) et Aphropsyche Ross (1).
Des clefs d’ identification augenre et a I’espece (pour les males, et les femelles lorsque possible) sont presentees par
1’ auteur. Un habitus ainsi qu’ une description diagnostique des pieces genitales sont donnes pour chacune des especes.
Une description resume ce qu’ il y a de connu sur I’histoire naturelle et la repartition geographique de ces especes. Les
aires de distributions et les pieces genitales sont largement ilustrees.
Table of Contents
Introduction 2
Techniques 3
Geographical Distribution 3
The Family Arctopsychidae Martynov .......4
Genus Arctopsyche McLachlan .....5
Genus Parapsyche Betten 14
The Family Hydropsychidae Curtis 22
Genus Cheumatopsyche Wallengren ...23
Genus Hydropsyche Pictet 76
2
Nimmo
Genus Potamyia Banks . 164
Genus Aphropsyche Ross 167
Genus Diplectrona Westwood 170
Genus Macrostemum Kolenati 173
Acknowledgements 179
References 180
Index 187
INTRODUCTION
This paper brings together information about all species of Arctopsychidae and
Hydropsychidae presently known from Canada, or with the potential to be found in Canada.
Consequently, all species of the two families recorded from contiguous States of the United
States, but not yet from Canada, are also included here.
The Hydropsychidae is one of the larger families of Trichoptera, world-wide, with 72 species
recorded here. These species are distributed among three subfamilies and six genera. The
Arctopsychidae, on the other hand, is a small family confined to the Holarctic region and its
fringes in south Asia. There are six species recorded here.
Characterisations of supra-specific taxa are abridged from Schmid (1980).
The key to genera is translated from Schmid (1980). For a key to genera of larvae, Wiggins
(1977) should be consulted. Keys to species are original, with the exception of that to
Cheumatopsyche which is partially adapted from Gordon (1974). These keys make fullest
possible use of the illustrations, and have been kept as simple as possible: use of more than two
characters per couplet has been avoided where possible; for most couplets, only one character is
used.
In the text for each species a general habitus description is presented in some detail. This is
based on the male, with mention of the female only when she is significantly different.
Regarding genitalia, a diagnostic statement only is presented, which makes reference to those
characters which will ensure correct recognition of a species in conjunction with the
illustrations. The male and female (where known) genitalia of each species are fully illustrated.
Notes about way of life and known distribution (with maps) complete the presentation for each
species.
Within subfamilies the genera are presented in alphabetical order. Within the genera the
species are also presented in alphabetical order, except where the genus has been subdivided to
species groups and subgroups, when the alphabetical ordering is used only within the lowest
grouping. This latter arrangement is used in Cheumatopsyche and Hydropsyche. In
Cheumatopsyche I use the infra-generic group names of Gordon (1974). In Hydropsyche I use
a numbering system for the groups, and a letter designation for the subgroups.
At the beginning of each species treatment (as also for the treatment of each genus,
subfamily, and family) a brief synopsis of the synonymy attaching to that taxon is presented,
with citations of the more important papers relevant to the history of the taxon. For a complete
listing of literature for each taxon, up to 1961 inclusive, Fischer (1963; 1972) should be
consulted.
Statements about colour are based on alcohol-preserved material. With such material,
fading may occur over time, and wing and body hairs may be lost. Wing colouration, therefore,
is based on membrane colour. This is the normal situation with fluid-preserved material, which
Arctopsychidae and Hydropsychidae (Trichoptera)
3
is usual in bulk collecting especially.
Species recognition in adults is based almost entirely on male and female genitalia. In the
males I follow Snodgrass (1957:35) in considering the aedeagus to be the entire evertible
assemblage located between the claspers (inferior appendages). I also prefer to use the term
clasper, rather than inferior appendage , as it has the merit of brevity and of describing the
apparent function of the appendage in question. The abdominal segments are referred to by use
of roman numerals, counting from the abdomen-thorax junction.
From the illustrations of genitalia I omit setae or hairs except in instances where they may
be of use in identification of a given species; otherwise they simply clutter the drawing and
obscure other features. The genitalia of many species are, however, well invested with setae or
hairs.
One final point to note, regarding descriptions or characterisations, is the use of the singular
and plural. Several components of the genitalia are paired but, in certain views only one
member of a pair is visible. When one member only is visible the singular is used. When both
are visible, the plural is generally used. Reference to the wings and legs is normally on the basis
of one member of each pair.
TECHNIQUES
Refer to Nimmo (1971) for details about collection, preservation, and preparation of
material for examination.
GEOGRAPHICAL DISTRIBUTION
A distribution map, or maps, is presented for each species dealt with here. For those species
presently known from Canada a detailed map of the Canadian distribution is given. An inset
map of North America presents a by state/province overview of the total known distribution in
North America. For those species not yet recorded from Canada only the overall North
American map is presented.
The State records for the United States portion of the North American maps are derived
from the literature. Most of the detailed Canadian records are new to the literature and are
derived from examination of museum or newly collected material. Such detailed records as
were obtained from the literature are considered to be trustworthy and no distinction is made
on the maps.
Apart from the transcontinental species the fauna may be divided into those relatively few
species known from the Rocky Mountain foothills and west, and the great bulk of species which
occur east of these foothills. A very few of these last are apparently restricted to the Great
Plains. The remainder are centred on eastern North America with extensions to the north-west,
to the north-east, and to the south, or combinations of these. Some species appear to be
confined to the environs of the Appalachians, or to the southern fringes of the Great Lakes,
with or without extensions southward along the valley of the Mississippi River. Many species
are still too poorly known for useful speculation regarding their distribution patterns.
Other than records of a few species from northern Quebec, these two families appear to be
limited northwards by the tree line.
A single, transcontinental, species is holarctic in total distribution, being known from
Eurasia as far west as northern Europe.
Quaest. Ent., 1987, 23 (1)
4
Nimmo
NOTE ADDED IN PRESS
Schefter, Wiggins, & Unzicker (1986) recently synonymized names of two species dealt
with here: Hydropsyche jewetti Denning ( = H. cockerel li Banks); and Hydropsyche riola
Denning (=H. alhedra Ross). Similarly, Schefter & Unzicker (1984) synonymised
Hydropsyche bifida Banks with H. morosa Hagen.
These synonymizations may be correct; indeed I am inclined to accept that they are.
However, in course of preparation of this paper, I encountered no material which might support
the above synonymizations, and prefer, for present purposes, to leave the text as originally
written. Thus, I have inserted guiding notes, in the appropriate parts of the text, relating to this
particular note.
THE FAMILY ARCTOPSYCHIDAE MARTYNOV
Arctopsychidae Martynov, 1924:25; Schmid, 1968:4; Schmid, 1980:51.
Arctopsychinae (Hydropsychidae); Milne, 1940:13, 19; Flint, 1961:5; Ross, 1956:10; Wiggins, 1977:93.
Description. — Females distinctly larger, more robust than males. Ocelli absent. Maxillary palpi of five articles;
with basal two articles very short, sub-equal; article four slightly shorter than article three; article five long, flagellate.
Antennae thickened, especially in male; with short, globular scape. Spurs large; spur formula 2,4,4. Fore- and hind-wings
(Fig. 2) virtually identical in individuals and between sexes; nearly oval in outline but hind-wings with anal edge evenly
rounded. Venation similar in the two genera dealt with here: fl-V present in fore-wings, fl-III and fV present in
hind-wings. Fore-wings with discoidal, median, and thyridial cells closed; thyridial cell especially long; cross-veins C-Sc,
Sc-Rl, and R1-R2 + 3 present; Cu2 terminated on A rather than on anal edge of wing; postcostal cell very large.
Hind-wing with cross-veins C-Sc and Sc-Rl present; with four separate anal veins.
Genitalia. Male. (Fig. 7-9, 25-28). Segment IX well developed dorsally. Preanal appendages free or fused to segment
X. Intermediate appendages also free or fused to each other. Claspers (inferior appendages) large or small, bipartite.
Aedeagus large, located high in the abdomen composed of tubular phallotheca with invaginated small, membranous,
erectile endotheca.
Genitalia. Female (Fig. 11-12, 29-30). Segments X and XI short. Tergite VIII very large; lateral edges produced
quite far ventrad. Sternite VIII correspondingly reduced, with posterior edge terminated in two large lobes. Segment IX
absent. Segment X enclosed by lobes of Sternite VIII, short, simple, without clasper receptacles; postero-dorsal edge with
two small tubercles, each with slender brush of very long hairs. Segment X with large, postero-ventral, membranous vulval
scale. Ano-vaginal opening on posterior end of segment X.
The Arctopsychidae are very closely related to the Hydropsychidae, and some authors regard it as a subfamily of the
Hydropsychidae (Wiggins, 1977). Based on Schmid (1968), and taking account of Smith (1968), the Arctopsychidae
encompass two known genera and a minimum of 43 species worldwide. The family is oriental and holarctic in overall
distribution, with most species concentrated in a zone extending from the Himalaya to Japan. Eleven species (one of which
is holarctic) are currently recognised in North America, with five known to occur in Canada and a sixth possibly to be
found here. Both genera occur in Canada, with three and two (three?) species of each represented.
Key to genera of Arctopsychidae of Canada
la Eyes glabrous. Male genitalia protuberant, not recessed into segment VIII.
Tibia and tarsus of female middle leg flattened, enlarged
Arctopsyche McLachlan, p. 5
lb Eyes hairy. Male genitalia barely protuberant, recessed into segment VIII.
Tibia and tarsus of female middle leg not flattened, not
enlarged Parapsyche Betten, p. 14
Arctopsychidae and Hydropsychidae (Trichoptera)
5
Genus Arctopsyche McLachlan
Maps 1-3; Fig. 2, 7-24
Arctopsyche McLachlan, 1868:300; Betten, 1934:179, 180; Milne, 1936:65; Flint, 1961:6; Schmid, 1968:21, 29, 32;
Wiggins, 1977:98; Schmid, 1980:52.
Description. — Eyes glabrous. Third article of maxillary palpi twice as long as wide; not much longer than fourth.
Spur formula 2,4,4. Tibia and tarsus of female middle leg enlarged, flattened, fringed with hairs. Discoidal cell of fore-
and hind-wings (Fig. 2), and median cell of fore-wing small.
Genitalia. Male. (Fig. 7-10, 13-16, 19-22). Segment IX nearly as large as VIII, not recessed within VIII. Preanal
appendages free (Fig. 7, 9); long, narrow, rounded. Intermediate appendages (Fig. 7, 9) stout, long blades; single or paired;
simple or with dentitions. Segment X entirely membranous, either very short or as long slender tube (Fig. 19, 20)
depending on species group. Claspers (inferior appendages) (Fig. 7, 8) with two articles, reduced in size, complex; basal
article massive, with dorsal lobe or spine and two or three ventral spines; distal article small, inserted between spines of
basal article. Aedeagus (Fig. 10) large, stout, with recurved internal phallotremal sclerite.
Genitalia. Female. (Fig. 11-12, 17-18, 23-24). Postero-dorsal margin of segment X with inconspicuous flange (Fig.
11). Segment XI developed basad.
In Canada Arctopsyche is represented by two species; one of these (A. ladogensis) is
holarctic in distribution, being recorded in boreal regions from northwestern Europe to
Newfoundland. A third species (A. irrorata) is presently known only from the southeastern
United States but may eventually be recognised from eastern Canada. A. grandis is primarily a
western montane species, but has been recorded from northwestern Quebec.
Key to known or potential species of Arctopsyche McLachlan of Canada
la Males (Fig. 7-10)
1 b Females (Fig. 11-12)
2a (la) Segment X prominent, projected posterad of intermediate appendages
(Fig. 19, 20) A. ladogensis (Kolenati), p. 7
2b Segment X not evident (Fig. 7, 13)
3a (2b) Each member of intermediate appendage pair with single dorsal and
ventral processes (Fig. 13, 15) A. irrorata Banks, p. 6
3b Each member of intermediate appendage pair with only one process
A. grandis (Banks), p. 5
4a (lb) Vulval scale with complex outline in ventral aspect (Fig. 12)
A. grandis (Banks), p. 5
4b Vulval scale with relatively simple outline in ventral aspect (Fig. 18, 24);
evenly tapered posterad
5a (4b) Ventral surface of vulval scale with darker, w- shaped transverse line in
ventral aspect (Fig. 24). Thorax and head purplish brown-black
A. ladogensis (Kolenati), p. 7
5b Ventral surface of vulval scale whithout transverse darker line (Fig. 18).
Thorax and head greyish brown A. irrorata Banks, p. 6
2
4
3
5
Arctopsyche grandis Banks
Map 1; Fig. 7-12
Arctopsyche grandis Banks, 1900:258; Milne, 1936:66 (A. phryganoides as synonym); Ross, 1 938c: 1 4; Schmid, 1968:54;
Smith, 1968:109 (A. inermis as synonym); Wiggins, 1977:99.
Arctopsyche phryganoides Banks, 1918:21; Milne, 1936:66 (as synonym of A. grandis).
Arctopsyche inermis Banks, 1943:368; Smith, 1968:109 (as synonym of A. grandis)-, Schmid, 1968:54.
Quaest. Ent., 1987,23 (1)
6
Nimmo
Description. — Male fore-wing length 12.56 mm; pale grey-brown with uniform faint irroration except for
coalescence of pale areas along costal edge; female irroration more evident. Hind-wing faintly tinted golden brown;
grey-brown in female. Antennae brown-cream; basal 19 flagellar annuli of male each with simple black bands; 17-18 in
female. Vertex dark brown. Spurs brown; lateral member of middle leg pairs notably shorter than mesal companions.
Thorax dark brown, to pale yellow-brown laterally. Legs pale brownish yellow.
Genitalia. Male. (Fig. 7-9). (Specimen from Wildhorse camp, Ya Ha Tinda Ranch road, Alberta). Males
distinguished by apparent lack of segment X (Fig. 7, 9); by dorsal lobe of clasper basal article broad in lateral aspect,
angled posterad (Fig. 7); and by ventral lobe of clasper basal article terminated in several acuminate teeth in ventral
aspect (Fig. 8).
Genitalia. Female. (Fig. 11-12). (Specimen from Wildhorse camp, Ya Ha Tinda Ranch road, Alberta). Females
distinguished by complexity of vulval scale in ventral aspect; scale with semi-circular posterior edge (Fig. 12) terminated
laterally by lateral processes; anterad of processes scale markedly constricted and anterad of constriction scale abruptly
expanded to greater width.
Biology. — British Columbia and Alberta records give flight season extremes of June 6 to
August 27, and May 26 to August 12 respectively, with definite peak indicated for July as a
whole. Smith’s (1968) records from Idaho conform to much the same pattern. He adds that the
commonest life stages in winter were mature larvae, with pupation occurring from April to
May. He concludes that this species has a two-year life cycle. My records indicate that adults
emerge from the largest mountain and foothill rivers, from smaller, riffled foothill streams,
from turbulent mountain streams, and from all intermediate types of flowing water in mountain
and foothill country. Adults have also been collected from clear, riffled, cool streams in
low-altitude terrain of little relief. Larvae appear to be, ecologically, very diverse or tolerant.
Wallace (1975a) presents information in support of the predaceous nature of larvae of
Arctopsyche. Mecom (1972) contends that A. grandis larvae are carnivorous in summer
months, but primarily phytophagous, diatom feeders, or detritovores at other times.
Distribution. — Recorded from the Yukon and western Northwest Territories in Canada, to
California and New Mexico in the United States, with one record from northwestern Quebec
(Map 1) in Canada; excepting the Quebec record, this species appears to be confined to the
Cordillera west of the Great Plains.
Arctopsyche irrorata Banks
Map 2; Fig. 13-18
Arctopsyche irrorata Banks, 1 905b:2 1 7; Milne, 1936:66.
Description. — Male fore-wing length 14.43 mm; medium grey-brown; randomly irrorate throughout, except more
regular alternate hyaline and coloured patches along costal edge and posterad as far as Rs. Hind-wing tinted grey-brown
(very pale). Antennae pale orange-brown; each annulus with deep chocolate-brown band on dorsal and lateral surfaces, at
right-angles to axis of annulus; scape with rectangular patch of dark brown laterally. Vertex pale orange-brown to
red-brown. Spurs red-brown; lateral member of each pair shorter than mesal companions. Thorax deep red-brown dorsally,
pale straw to cream laterally. Legs pale red-brown. Female overall darker than male.
Genitalia. Male. (Fig. 13-16). (Specimen from Citico Ck, Monroe Co., Tennessee, USA). Males distinguished by
each member of intermediate appendage pair bifid (Fig. 13, 15); by apparent lack of segment X; by ventral lobe of clasper
single, not subdivided (Fig. 14), with distal hook turned dorsad in lateral aspect (Fig. 13); and by dorsal lobe of clasper
directed dorsad, irregular in outline.
Genitalia. Female. (Fig. 17-18). (Specimen from Citico Ck, Monroe Co., Tennessee, USA). Females distinguished by
vulval scale fairly regular in ventral outline (Fig. 18), straight-edged laterally, with central portion slightly raised, tapered
posterad, and with semi-circular membranous lobe on distal edge.
Biology. — Flint (1961) records small larvae in early summer, mature larvae in fall, and
adult flight season as late May and June. Overwintering is by mature larvae. He adds that
larvae live in fairly large streams up to 25 m wide, and 1 m deep; the water is clear, cold, with
gravel and boulder substrate. He also records abundance of larvae in shallow water falling
rapidly over bedrock. Both animal and plant material are consumed, with animal matter
predominating. Wallace (1975a) provides full details about retreat and net structure, and
Arctopsychidae and Hydropsychidae (Trichoptera)
7
seasonal feeding habits.
Distribution. — To date this species is recorded definitely only from Tennessee, and North
& South Carolina (Map 2). Ross (1944) records the species from ‘eastern States and eastern
Canada’, hence its inclusion here. But as, in the same sentence, he also includes A. ladogensis,
this ‘record’ may simply result from imprecise wording. More definite records could not be
found to substantiate Ross’ statement.
Arctopsyche ladogensis (Kolenati)
Map 3; Fig. 19-24
Aphelocheira ladogensis Kolenati, 1859:145, 147, 165, 183, 201.
Arctopsyche ladogensis ; McLachlan, 1868:301; Milne, 1936:66, 67; Schmid, 1968:56.
Arctopsyche ladogensis form, ramosa McLachlan, 1878:379; Milne, 1936:67.
Arctopsyche ladogensis form, obesa McLachlan, 1868:301; Milne, 1936:67.
Description. — Male fore-wing length 1 1.23 mm; light purplish brown, no pattern evident. Hind-wing palely tinted
brown. Antennae pale yellow-brown; basal 19 flagellar annuli with at least trace of darker encircling band; 14 in female.
Vertex deep red-brown. Spurs brown; lateral member of middle leg pairs markedly shorter than mesal companions. Thorax
very deep purplish brown-black, to paler, mottled red-brown laterally. Legs brown, to straw distally.
Genitalia. Male. (Fig. 19-22). (Specimen from House R., Hwy 63, S of Ft McMurray, Alberta). Males distinguished
by segment X prominent (Fig. 19, 20); by much smaller clasper with finger-like dorsal lobe accompanied by spine
immediately anterad (Fig. 19); and by distinct dorsal spine on distal edge of aedeagus (Fig. 22).
Genitalia. Female. (Fig. 23-24). (Specimen from House R., Hwy 63, S of Ft McMurray, Alberta). Females
distinguished by vulval scale simple, tapered, with slight mesal extension of posterior edge (Fig. 24); and by transverse line
of darker colour in form of a W in ventral aspect.
Biology. — Flint (1961) records larvae of this species from clear, cold streams of up to 25 m
width and 1 m depth, on gravel or boulder beds. Retreat and net are typical for the genus.
Mature larvae are recorded in August and May, and are apparently the overwintering stage.
Pupation occurs in late May or June. Flight records are for June (Flint, 1961). Records
available to me from Canada give a flight season from May 12 to August 18, with the bulk in
June and July. I have records, from northwestern Canada, of adults taken adjacent to rivers of
75 m width or more.
Distribution. — In North America this circumboreal species is known from western Alaska
to Newfoundland, the northeastern United States, and Michigan (Map 3). From the Map one
might conclude that the species is limited northward by tree line.
Quaest. Ent., 1987, 23 (1)
8
Nimmo
Map 1. Collection localities for Arctopsyche grandis (Banks) in Canada, with known distribution in North America by
state or province.
Map 2. Known distribution of Arctopsyche irrorata Banks in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
9
Map 3. Collection localities for Arctopsyche ladogensis (Kolenati) in Canada and Alaska, with known distribution in
North America by state or province.
Quaest. Ent., 1987,23 (1)
10
Nimmo
Fig. 1-6. Wing venation, males. 1, Apropsyche doringa (Milne). 2, Arctopsyche irrorata Banks. 3, Cheumatopsyche
pettiti (Banks). 4, Diplectrona modesta Banks. 5, Hydropsyche alhedra Ross. 6, Potamyia flava (Hagen), ‘a’ figures are
fore-wings; ‘b’ are hind-wings, d, discoidal cell; m, median cell; th, thyridial cell; pc, postcostal cell.
Arctopsychidae and Hydropsychidae (Trichoptera)
11
Fig. 7-12, Arctopsyche grandis (Banks): 7, genital capsule of male, lateral aspect; 8, claspers of male, ventral aspect; 9,
intermediate and preanal appendages of male, dorsal aspect; 10, aedeagus of male, lateral aspect; 11, genital segments of
female, lateral aspect; 12, genital segments of female, ventral aspect, inf, inferior appendage (clasper); int, intermediate
appendage; pr, preanal appendage; ce, cercus; vs, vulval scale.
Quaest. Ent., 1987,23 (1)
12
Nimmo
Fig. 13-18, Arctopsyche irrorata Banks: 13, genital capsule of male, lateral aspect; 14, claspers of male, ventral aspect; 15,
intermediate appendages of male, dorsal aspect; 16, aedeagus of male, lateral aspect; 17, genital segments of female,
lateral aspect; 18, genital segments of female, ventral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
13
Fig. 19-24, Arctopsyche ladogensis (Kolenati): 19, genital capsule of male, lateral aspect; 20, genital capsule of male,
dorsal aspect; 21, claspers of male, ventral aspect; 22, aedeagus of male, lateral aspect; 23, genital segments of female,
lateral aspect; 24, genital segments of female, ventral aspect.
Quaest. Ent., 1987,23 (1)
14
Nimmo
Genus Parapsyche Betten
Maps 4-6; Fig. 25-43
Parapsyche Betten, 1934:179, 181; Schmid, 1968:26,31,60; Wiggins, 1977:1 14; Schmid, 1980:53.
Description. — Eyes densely clothed with long hairs. Palpi little longer than in Arctopsyche. Third article of
maxillary palpi at least three times longer than wide; with noticeable bulge on mesal face. Spur formula 2,4,4. Tibia and
tarsus of female middle leg not flattened, not enlarged, not fringed with hairs. Hind leg femur very long, tarsi reduced,
especially in female. Venation identical to that of Arctopsyche except discoidal and median cells of fore- wing distinctly
longer.
Genitalia. Male. (Fig. 25-28, 31-35, 38-41). Segment IX rather small; partly recessed into segment VIII; short, with
dorsal part prominent due to lower position of intermediate appendages (Fig. 25). Preanal appendages closely blended with
base of intermediate appendages apparently little more than scars. Intermediate appendages large, horizontal, slightly
sclerotised blades; fused basally (Fig. 26) or throughout most of their length (Fig. 40); connected to aedeagus by two
clearly visible internal straps (Fig. 25, 26, 28). Claspers (inferior appendages) large, directed somewhat postero-dorsad
(Fig. 25); v/ith two articles. Basal article of clasper large; distal article on apex or middle of basal article; articles partly
fused (Fig. 25, 27). Aedeagus with tubular phallotheca, and membranous, erectile endotheca with external, paired
phalotremal sclerites curved basad (Fig. 28). Phallotheca surmounted by dorsal lobe, a prolongation of postero-dorsal
margin of basal phallocrypt.
Genitalia. Female. (Fig. 29-30, 36-37, 42-43). Very similar to Arctopsyche. Dorso-lateral posterior margin of
segment X not flanged (Fig. 29). Segment XI produced less prominently basad.
In Canada Parapsyche is represented by three species, two of which are western cordilleran,
and the third eastern, in distribution. The larvae of all three species are known.
Key to species of Parapsyche of Canada
la Males 2
lb Females 4
2a (la) Intermediate appendages, in dorsal aspect (Fig. 26, 33), fused only at base 3
2b Intermediate appendages fused throughout length (Fig. 40)
P. elsis Milne, p. 16
3a (2a) Dorsal part of segment IX smoothly continuous with main body of
segment, along posterior edge (Fig. 25). Western cordilleran species (Map
4) P.almota Ross, p. 14
3b Dorsal part of segment IX angled sharply posterad from main body of
segment (Fig. 31). Eastern species (Map 5) . ... P. apicalis (Banks), p. 15
4a (lb) Segment X slender in lateral aspect, small (Fig. 29, 36). Vulval scale
medium to large in ventral aspect (Fig. 30, 37) 5
4b Segment X massive, wide in lateral aspect (Fig. 42). Vulval scale small in
ventral aspect (Fig. 43) P. elsis Milne, p. 16
5a (4a) Vulval scale with triangular, membranous tip (Fig. 30). Eastern species
(Map 5) P. apicalis (Banks), p. 15
5b Vulval scale with rounded tip, scarcely membranous (Fig. 37). Western
cordilleran species (Map 4) P. almota Ross, p. 14
Parapsyche almota Ross
Map 4; Fig. 25-30
Parapsyche almota Ross, 1 938a: 1 19; Schmid, 1968:106-107, Fig. 84; Smith, 1968:105.
Arctopsyche oregonensis Ling, 1938:65; Ross, 1944:293.
Description. — Male fore-wing length 9.36 mm; grey-brown, heavily irrorate, with relatively large areas of hyaline
membrane. Hind-wing faintly tinted brown. Female fore-wing more faintly irrorate; hind-wing grey-brown. Antennae
Arctopsychidae and Hydropsychidae (Trichoptera)
15
brown; about 19 basal flagellar annuli with dark brown band around each. Vertex red-brown; posterior warts pale. Spurs
yellow; lateral member of middle leg pairs notably shorter than mesal companions. Thorax deep chocolate-brown to paler,
grey-brown laterally. Legs straw-coloured.
Genitalia. Male. (Fig. 25-28). (Specimen from small, turbulent creek, Hwy 12, 12.7 km NE of Lilloet, British
Columbia). Males distinguished by dorsal portion of segment IX smoothly continuous with main body of segment along
posterior edge, in lateral aspect (Fig. 25); by intermediate appendages curved dorsad; by distal article of clasper short,
conical in lateral aspect; and by intermediate appendages fused only at base (Fig. 26).
Genitalia. Female. (Fig. 29-30). (Specimen from small, turbulent creek, Hwy 12, 12.7 km NE of Lilloet, British
Columbia). Females distinguished by segment X narrow (Fig. 29); and by vulval scale with distal half membranous,
triangular (Fig. 30).
Biology. — Smith (1968) gives the Idaho flight season as April to October; my scanty
Canadian records are within this range. Smith also concludes that medium to mature larvae are
the overwintering stages; they are found on the small to medium rubble of small, clear creeks
and streams. Of my two records from British Columbia one locality was a steep hill stream
flowing over and around large boulders; the second stream was very shallow with fine gravel, at
the outlet of a swamp.
Distribution. — This species is presently recorded from the Cariboo-Chilcotin district of
west-central British Columbia, to Nevada and California (Map 4) and is probably confined to
the Cordillera west of the continental divide. In Canada it is known only from western British
Columbia in the southern coastal mountains.
Parapsyche apicalis (Banks)
Map 5; Fig. 31-37
Arctopsyche apicalis Banks, 1908:266.
Parapsyche apicalis\ Betten, 1934:181; Flint, 1961:8; Schmid, 1968:Fig. 108-110; Wiggins, 1977:115.
Description. — Male fore-wing length 8.58 mm; largely uncoloured membrane with scattered fragments of
grey-brown. Antennae pale purplish brown with darker ring near distal end of each annulus. Vertex purple-brown, warts
paler; red-brown in female. All spur pairs with lateral members shorter than mesal members. Thorax dark purple-brown
dorsally; paler laterally, with some red-brown areas. Female thorax red-brown dorsally, to yellow-brown laterally. Legs
yellow.
Genitalia. Male. (Fig. 31-35). (Specimen from Fox Point, Cumberland Co., Nova Scotia). Males distinguished by
dorsal portion of segment IX sharply angled posterad (Fig. 31), in lateral aspect; by intermediate appendages more or less
linear, sloped slightly ventrad; by distal article of clasper prominent, finger-like; and by intermediate appendages fused
only at base.
Genitalia. Female. (Fig. 36-37). (Specimen from Baden, Ontario). Females differentiated by segment X narrow (Fig.
36); and by vulval scale broad, rounded distally, with little distal membrane (Fig. 37).
Biology. — Flint (1961) records the larvae as inhabiting cold (below 10°C), spring-fed
brooklets only a metre or so wide and, occasionally, in rushing mountain streams several metres
wide. The retreat and net are typical for the family. Overwintering is by larvae of several
instars. Pupae occur throughout summer. Adults recorded from May 11 to October 1. Flint
considers it possible that there are several generations per year, but without definite broods.
The above seasonal information is derived from Massachussetts. Records from Canada give a
flight season of May 7 to October 12; some agree with Flint’s characterisation of the habitat as
cold, spring-fed brooks. The remainder, however, indicate that larvae also inhabit warmer,
larger streams.
Distribution. — In North America this species is confined to the eastern half of the
continent (Map 5), ranging from Newfoundland and Ontario to Wisconsin, Tennessee, and
North Carolina. In Canada the species is recorded from St John’s, Newfoundland, to Lake
Nipigon, Ontario, and south to the Niagara peninsula. An old record from Colorado almost
certainly represents a misidentification of P. almota before that species was recognized by Ross
- the two species are very similar as adults.
Quaest. Ent., 1987,23 (1)
16
Nimmo
Parapsyche elsis Milne
Map 6; Fig. 38-43
Parapsyche elsis Milne, 1936:66, 67; Schmid, 1968:Fig. 1 1 1-1 13; Smith, 1968:107.
Description. — Male fore-wing length 12.4 mm; pale red-brown (grey-brown in female), very heavily irrorate.
Hind-wing tinted pale grey-brown on distal third. Antennae brownish cream; each flagellar annulus encircled in distal
quarter by brown band. Vertex brownish yellow except antero-mesal warts brown. Spurs brown; lateral member of middle
and hind-leg pairs notably shorter than mesal companions. Thorax yellow-brown, to brownish cream laterally. Legs pale
yellow-brown.
Genitalia. Male. (Fig. 38-41). (Specimen from Fiddle R., Hwy 16, Jasper National Park, Alberta). Males
distinguished by dorsal portion of segment IX inclined posterad at about 45° to remainder of segment (Fig. 38); by
intermediate appendages fused throughout length (Fig. 40), more or less linear, horizontal; by clasper with articles almost
indistinguishably fused in lateral aspect (Fig. 38), massive, long; by phallotremal sclerites at tip of aedeagus (Fig. 41)
small, with fine spines directed basad.
Genitalia. Female. (Fig. 42-43). (Specimen from Fiddle R., Hwy 16, Jasper National Park, Alberta). Females
distinguished by massive, wide segment X in lateral aspect (Fig. 42); and by small vulval scale tapered sinuously distad,
distal half trapezoidal, membranous (Fig. 43).
Biology.— Smith (1968) suggests that this species has a two-year life cycle. Overwintering
is by young or nearly mature larvae; pupation evident in June and July. Larvae found in flowing
waters ranging from small, clear creeks, to largest rivers with boulder beds; from hill streams to
mountain torrents. Canadian records indicate flight season ranges from June 26 to September
8.
Distribution. — This species is (with one improbable exception) restricted to the western
Cordillera of North America (Map 6), ranging from the Yukon and the Mackenzie Mountains
of the Northwest Territories of Canada south to California and Utah. In Canada it is recorded
generally from the various ranges of the western Cordillera as far north as the Ogilvie
Mountains of the Yukon Territory. An isolated occurrence was recorded for North Carolina.
Milne and Milne (1938) had only the five type specimens available at that time, to one of which
this curious record is attributed. They make no comment on it. It seems to be so improbable
that one is inclined to conclude that it is the result of locality mis-labelling of specimens at some
earlier date.
Arctopsychidae and Hydropsychidae (Trichoptera)
17
Map 4. Collection localities for Parapsyche almota Ross in Canada, with known distribution in North America by state or
province.
Map 5. Collection localities for Parapsyche apicalis (Banks) in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987, 23 (1)
18
Nimmo
Map 6. Collection localities for Parapsyche elsis Milne in Canada and Alaska, with known distribution in North America
by state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
19
Fig. 25-30, Parapsyche almota Ross: 25, genital capsule of male, lateral aspect; 26, genital capsule of male, dorsal aspect;
27, right clasper of male, ventral aspect; 28, aedeagus of male, lateral aspect; 29, genital segments of female, lateral
aspect; 30, genital segments of female, ventral aspect, int, intermediate appendage; ce, cercus; vs, vulval scale.
Quaest. Ent., 1987,23 (1)
20
Nimmo
Fig. 31-37, Parapsyche apicalis (Banks): 31, genital capsule of male, lateral aspect; 32, right clasper of male, ventral
aspect; 33, genital capsule of male, dorsal aspect; 34, aedeagus of male, lateral aspect; 35, aedeagus of male, distal half,
ventral aspect; 36, genital segments of female, lateral aspect; 37, genital segments of female, ventral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera) 21
Fig. 38-43, Parapsyche elsis Milne: 38, genital capsule of male, lateral aspect; 39, right clasper of male, ventral aspect; 40,
genital capsule of male, dorsal aspect; 41, aedeagus of male, lateral aspect; 42, genital segments of female, lateral aspect;
43, genital segments of female, ventral aspect, pr, preanal appendage.
Quaest. Ent., 1987, 23 (1)
22
Nimmo
THE FAMILY HYDROPSYCHIDAE CURTIS
Hydropsychidae Curtis, 1835:P1. 544 (& text); Betten, 1934:116, 123, 176; Milne, 1936:63, 67; Ross, 1944:76; Wiggins,
1977:92; Schmid, 1980:55.
Hydropsychinae; (Subfamily of Phryganeidae) Burmeister, 1839:907.
Description. — Ocelli absent. Eyes very large in males of some species. Maxillary palpi with five articles in both
sexes; basal article variable in length; fifth article very long, flagellate. Tibia and tarsus of female middle legs enlarged,
flattened in many taxa. Spur formula generally 2,4,4. Sternite V in many taxa with lobe with aperture of internal gland.
Fore- and hind-wings different in shape. Fore-wing smoothly, slightly broadened distally, with truncate apex (Fig. 3a).
Hind-wing larger, more rounded (Fig. 3b). Venation complete, with five marginal forks (fl-fV) generally present in
fore-wing, and fl-fHI and fV present in hind-wing. Discoidal and median cells closed in fore-wing; thyridial cell very long.
In hind-wing Sc in contact with Rl; discoidal and median cells open or closed; four free anal veins extended to wing
margin.
Genitalia. Male. (Fig. 44-47, 469-472, 483-487). Quite similar in all genera. Segment IX in most taxa, short, well
developed dorsally (Fig. 44, 469, 483). Segment X bulky, located roof-like over aedeagus, with various specifically
characteristic lobes. Preanal appendages present; entirely integrated with segment X; reduced, simple tubercles, each with
tuft of short hairs. Intermediate appendages absent. Claspers (inferior appendages) directed postero-dorsad; of two simple,
slender articles; curved mesad (Fig. 45) and, as pair, pincer-like. Aedeagus large, with greatly expanded base (Fig. 46);
entirely, or almost entirely, composed only of the phallotheca. Endotheca absent or minute; simple or complex.
Genitalia. Female. (Fig. 48-49, 473-474, 488-489). As in males, females are equaly similar in all genera. Sternite
VIII more or less divided to two large lobes by postero-ventral Fissure. Segment X large, roughly triangular in lateral
aspect (Fig. 49); produced deeply ventrad. Dorsal extremity of segment X with two tufts of stout hairs (omitted from
illustrations here). Segment XI basically high, oblique rectangle, clothed in short hairs; with two small, tube-like, papillate
lobes with cerci between, latter in many taxa with vestiges of distal article. Vulval scale large, elongate, largely
membranous, basally articulated with ventral extremity of segment X.
Hydropsychid larvae are filter-feeders, building a sheltered retreat with attached net
projected into the current, which is typical. Macronematine larvae are more specialised in their
structures, and frequent warmer, less turbulent waters. Very few species are recorded west of
the Great Plains of North America, in the Cordillera. Larvae of some species live along
wave-swept lake shores.
While retaining a basic uniformity throughout, the Hydropsychidae are represented by a
multitude of species on every continent but Antarctica. In Canada are found six genera. The
species known, or likely to be recorded from Canada number 72.
Key to known or potential genera of Hydropsychidae of Canada
la Discoidal and median cells of fore- wing small. Discoidal cell of hind-wing
open. Fore-wing brown, barred with yellow
Macronematinae Ulmer p. 173
Macrostemum Kolenati, p. 173
lb Sternite V with long filament. Hind-wings large, rounded, with Sc and Rl
markedly bowed distally (Fig. 4b). Antennae slightly crenate
Diplectroninae Ulmer, p. 167 2
lc Sternite V at most with lobe or faint prominence. Hind-wing constricted
somewhat distally (Fig. 3b, 5b, 6b). Antennae slender,
cylindrical Hydropsychinae Curtis, p. 23 3
2a (lb) Hind-wing apex rounded; with Sc and Rl bowed deeply apically (Fig.
4b) Diplectrona modesta Banks, p. 170
2b Hind-wing apex more bluntly rounded (Fig. lb); with Sc and Rl not bowed
deeply apically Aphropsyche doringa Milne, p. 167
3a (lc) Stems of M and Cul of hind-wing parallel, very close together (Fig. 5b,
6b); fl present
4
Arctopsychidae and Hydropsychidae (Trichoptera)
23
3b Stems of hind-wing M and Cul not parallel, not close; fl absent (Fig.
3b) Cheumatopsyche Wallengren, p. 23
4a (3a) In fore-wing, cross-veins M3 + 4-Cul and Cul-Cu2 close (Fig. 6a); A in
contact with Cu2 prior to wing margin. Hind-wing median cell open (Fig.
6b) Potamyia flava (Hagen), p. 164
4b In fore-wing, cross-veins M3 + 4-Cul and Cul-Cu2 clearly separated (Fig.
5a); A terminated at wing margin. Hind-wing median cell closed (Fig.
5b) Hydropsyche Pictet, p. 76
The Subfamily Hydropsychinae Curtis
Hydropsychinae Curtis, 1835: pi. 544 (text); Betten, 1934:118, 177; Milne, 1936:67; Ross & Unzicker,
1977:298; Wiggins, 1977:93; Schmid, 1979:48; Schmid, 1980:48.
Description. — Antennae slender, slightly longer than fore-wings, especially in male. Maxillary palpi with first and
second articles equal; third and fourth articles short. Vertex with more than two warts. Tibia and tarsus of female middle
legs flattened, enlarged. Fore-wing slightly, evenly widened distad (Fig. 3a, 5a, 6a); distally truncate. Hind-wing (Fig. 3b,
5b, 6b) slightly larger than fore-wing; with evenly rounded anal edge, but distally constricted. Venation complete; fl-fV
present in fore-wing, fl-fHI and fV in hind-wing. Discoidal and median cells rather small; fl and fill petiolate. Hind-wing
discoidal cell closed; median cell open (Fig. 6b) or closed (Fig. 5b).
Genitalia. Male. (Fig. 44-47, 182-186, 463-466). Segment IX rather short; with postero-lateral edge more or less
projected posterad. Segment X either simple, or bilobed with distal lobes or processes. Claspers (inferior appendages)
slender, of two articles. Aedeagus simple or with complex apex.
Genitalia. Female. (Fig. 48-49, 187-188, 467-468). Segment X with only narrow ventro-lateral extremity which
serves to support vulval scale. Dorso-lateral face of segment X with more or less evident clasper receptacle, or apparently
absent.
Genus Cheumatopsyche Wallengren
Maps 7-30; Fig. 3, 44-181
Cheumatopsyche Wallengren, 1891:138, 142; Denning, 1943:138; Ross, 1944:108; Gordon, 1974:117; Wiggins, 1977:100;
Schmid, 1980:60.
Ulmeria Navas, 1918:15; Navas, 1933:98.
Hy dropsy chodes Ulmer, 1905:34; Kimmins, 1963:130.
Description. — Small, slender insects. Tarsal claws variously /deformed or normal. Cross-veins M3 + 4-Cul and
Cul-Cu2 of fore-wings adjacent (Fig. 3a). Hind-wing fl absent; median cell open; stems of M and Cul divergent, not very
close (Fig. 3b).
Genitalia. Male. (Fig. 44-47, etc.). Very similar to Hydropsyche species. Segment IX postero-lateral angle generally
less prominent, more ventrad (Fig. 44, 50). Segment X less bulky; terminated in two or four setose, specifically distinct
lateral lobes (Fig. 47, 53), separated by inconspicuous median bridge. Claspers (inferior appendages) slender, with distal
article of many species curved mesad, claw-like (Fig. 45). Aedeagus proximally very large; simple; distally terminated by
two large endothecal valves.
Genitalia. Female. (Fig. 48-49, etc.). Tergite VIII with posterior edge not notched. Sternite VIII divided
longitudinally to two halves or lobes. Segment X without postero-lateral margin produced posterad. Clasper receptacle or
not; the term “chimney” is used here to refer to the ‘apical two-thirds’ of clasper receptacle (Jordan, 1974).
Wiggins (1977), partly quoting from earlier authors, wrote us that Cheumatopsyche larvae
tend to be more dominant in warmer streams than Hydropsyche larvae, and to be more
pollution tolerant than most other species of caddisflies. Larvae of this genus have also been
found as deep as 20 cm in stream-bed gravels. Gut-content analysis indicates that feeding is
largely on algae and small animals, with little detrital component.
Cheumatopsyche is represented in all regions but the Neotropical and the Antarctic
continent. Gordon (1974) identifies 39 species known from the Nearctic region. In this work I
deal with 24 species known from Canada, or potentially to be found here.
Quaest. Ent., 1987,23 (1)
24
Nimmo
Key to known or potential species of Cheumatopsyche of Canada (Adapted from Gordon, 1974)
la Males
lb Females
2a (la) Distal article of clasper not produced to tapered apex in lateral aspect (Fig.
50); short, blunt sordida complex, p. 27
2b Distal article of clasper long, tapered (Fig. 135) . . . gracilis complex, p. 32
3a (2a) Aedeagus base very large (Fig. 46). Lobes of tergum X, acuminate hooks
directed dorsad (Fig. 44). Tergum X with long, sclerotised, distally
bulbous, median process emergent posteriorly from below tergum (Fig. 44,
47) C. minuscula (Banks), p. 27
3b Aedeagus base moderately large at most (Fig. 52). Lobes of tergum X wide
in lateral aspect (Fig. 50). Median process absent
C. sordida (Hagen), p. 28
4a (2b) Dorsum of tergite X domed (Fig. 99) C. wabasha Denning, p. 46
4b Dorsum of tergite X flat (Fig. 142)
5a (4b) Dorsal lobes of segment IX indistinctly defined, with single seta (Fig. 130).
Aedeagus elongate, almost linear C. vannotei Gordon, p. 60
5b Dorsal lobes of segment IX clearly defined, with numerous setae (Fig.
163). Aedeagus not elongate, not linear
6a (5b) Preanal appendages vertically long, linear in lateral aspect (Fig. 163) ....
C. mollala Ross, p. 65
6b Preanal appendages vertically short, circular in lateral aspect (Fig. 63)
7a (6b) Distal lobes of tergum X with apices elongate, reflexed (Fig. 59, 66, 72)
7b Distal lobes of tergum X with apices not reflexed (Fig. 148)
8a (7a) Distal article of clasper, in posterior aspect, short (Fig. 64); not fully
curved; distal portion straight or possibly slightly recurved. Lobes of
tergum X broad, rounded in lateral aspect (Fig. 63); anterad of turned-up
distal edges of tergum; lobes not visible in posterior aspect
C. pinaca Ross, p. 32
8b Distal article of clasper, in posterior aspect (Fig. 57, 70), longer; entire
article almost straight. Distal lobes of tergum X with apices visible in
posterior aspect (Fig. 59, 72); distal edge of tergum turned dorsad (Fig.
59), or not (Fig. 72)
9a (8b) Distal article of clasper, in lateral aspect (Fig. 56) hooked sharply dorsad
at tip. Distal edge of tergum X produced posterad of distal lobes, flared
dorsad C. speciosa (Banks), p. 32
9b Distal article with only slightly curved tip (Fig. 69). Distal edge of tergum
X not produced posterad of distal lobes, not flared dorsad
C. lasia Ross, p. 32
10a (7b) Distal article of clasper two-thirds length of basal article (Fig. 145, 146) . .
C. oxa Ross, p. 64
10b Distal article, at most, only half length of basal article (Fig. 107, 113, 129) . .
11a (10b) Distal lobes of tergum X circular or ovate dorsally, in posterior aspect (Fig.
110, 116, 131)
1 lb Distal lobes of tergum X not circular or ovate dorsally, in posterior aspect
2
25
3
4
5
6
7
8
10
9
11
12
Arctopsychidae and Hydropsychidae (Trichoptera)
25
(Fig. 141, 153, 179)
12a (11a) Distal lobes of tergum X widely separated from main body of tergum, in
lateral aspect C. pettiti (Banks), p. 48
12b Distal lobes with only short gap between them and main body of tergum X
(Fig. 107, 129)
13a (12b) Distal article of clasper, in posterior aspect, clearly recurved (Fig. 108) .
C. smithi Gordon, p. 48.
14
13
13b Distal article curved dorsad only (Fig. 130)
C. h. harwoodi Denning, p. 54
14a (lib) Basal article of clasper strongly curved mesad in posterior aspect (Fig.
177). Mesal face of distal article concave C. enonis Ross, p. 66
14b If clasper curved mesad than mesal face of distal article not concave (Fig. 127) . . 15
15a (14b) Distal lobes of tergum X directed antero-laterad (Fig. 125) in lateral
aspect, appressed against side of tergum C. wrighti Ross, p. 54
15b Distal lobes of tergum X not as above (Fig. 151) 16
16a (15b) Distal lobes of tergum X wide, rectangular in lateral aspect (Fig. 151) .
C. aphanta Ross, p. 64
16b Distal lobes of tergum X narrow, or not rectangular (Fig. 129) 17
17a (16b) Distal lobes of tergum X subquadrate, with lateral processes near venter
(Fig. 84) C. mickeli Denning, p. 38
17b Distal lobes of tergum X not as above 18
18a (17b) Distal lobes of tergum X oval or lanceolate (Fig. 78, 172) 19
18b Distal lobes of tergum X either square, clavate, or shouldered (Fig. 87, 93, 1 19) ... .
. 20
19a (18a) Distal lobes of tergum X without gap between them and main body of
tergum X (Fig. 169); dorsal tips clearly separated (Fig. 172)
C. burksi Ross, p. 65
19b Distal lobes of tergum X inclined posterad away from main body of tergum
X (Fig. 75); dorsal tips close to each other (Fig. 78)
C. pasella Ross, p. 38
20a (18b) Clasper markedly curved; distal article with broad base in posterior aspect
(Fig. 136, 158) 21
20b Clasper not markedly curved, or distal article narrow, sinuate (Fig. 88, 94,
105, 121) 22
21a (20a) Distal lobes of tergum X high, narrow, finger-like in lateral aspect (Fig.
157) C. halima Denning, p. 65
21b Distal lobes of segment X short, wide, with distinctly angled antero-dorsal
edge in lateral aspect (Fig. 135) C. gracilis (Banks), p. 60
22a (20b) Distal lobes of tergum X, in posterior aspect (Fig. 120), short, clearly
separated, with distinct lateral angle C. helma Ross, p. 54
22b Distal lobes of tergum X, in posterior aspect (Fig. 90, 95, 104), close, with
dorsal prolongations 23
23a (22b) Segment IX with distinct angular development of postero-lateral margin
C. ela Denning, p. 39
23b Postero-lateral margin of segment IX not angled (Fig. 87, 103) 24
24a (23b)Dorso-lateral lobes of segment IX well developed (Fig. 87)
Quaest. Ent., 1987, 23 (1)
26
Nimmo
C. campyla Ross, p. 38
24b Dorso-lateral lobes of segment IX barely evident (Fig. 103)
C. logani Gordon, p. 48
25a (lb) Clasper receptacle absent, or not visible, or minute in lateral aspect (Fig.
49, 55) sordida complex, p. 27
25b Clasper receptacle clearly visible in lateral aspect (Fig. 60, 68, 156)
gracilis complex, p. 32
26a (25a) Clasper receptacle, in lateral aspect (Fig. 49), a minute circle high up on
segment X C. minuscula (Banks), p. 27
26b Clasper receptacle, in lateral aspect (Fig. 55), very small, triangular
invagination high up on segment X on border between darker anterior bulk
of segment, and paler posterior area C. sordida (Hagen), p. 28
27a (25b) Clasper receptacle short, small, located high on segment X (Fig. 86, 174, 181)
27b Clasper receptacle longer, larger, in most specimens located at a level
ventrad of ventral lobe of segment XI (Fig. 97, 112, 118, etc.)
28a (27a) Inner end of clasper receptacle, in lateral aspect, apparently not open,
rounded (Fig. 174, 181)
28b Inner end of clasper receptacle, in lateral aspect, clearly open (Fig. 86) . . .
C. mickeli Denning, p. ...
29a (28a) Inner end of clasper receptacle, in lateral aspect, directed anterad (Fig.
181). Vulval scale without sclerotised band C. enonis Ross, p. 66
29b Inner end of clasper receptacle, in lateral aspect, directed dorsad (Fig.
174). Vulval scale with very narrow sclerotised strap, proximal end
abruptly flared C. burksi Ross, p. 65
30a (27b) Clasper receptacle outer edge without of marginal incision (Fig. 60, 68,
118, 140, 156, 162, 168)
30b Clasper receptacle outer edge with incision, either rounded or angular (Fig.
74, 80, 92, 97, 112, 124, 134, 150)
31a (30a|Fosterior margin of segment X overlapped by marginal flange developed
from outer edge of clasper receptacle
31b No such flange or overlap (Fig. 60, 68, 1 18, 140, 168)
32a (31a) Development of flange dorsad, along posterior edge of segment X; clasper
receptacle apparently with two chimneys (Fig. 156)
C. aphanta Ross, p. 64
32b No such development of flange (Fig. 162) C. halima Denning, p. 65
33a (31b) Anterior end of outer margin of clasper receptacle continued on lateral face
of segment X as thin, black line; length various with species (Fig. 60, 1 18, 140)
33b No such continuation of outer margin (Fig. 68, 168)
34a (33a) Clasper receptacle tubular (Fig. 60) C. speciosa (Banks), p. 32
34b Clasper receptacle papillate (Fig. 1 18, 140)
35a (34b) Clasper receptacle long, located dorsally (Fig. 140)
C. gracilis (Banks), p. 60
35b Clasper receptacle small, located at level ventrad of ventral lobe of segment
XI (Fig. 118) C. pettiti (Banks), p. 48
36a (33b) Posterior edge of each half of sternite VIII with small, rounded process
close to lateral corner (Fig. 68) C. pinaca Ross, p. 32
26
27
28
30
29
31
37
32
33
34
36
35
Arctopsychidae and Hydropsychidae (Trichoptera)
27
36b No such process present C. mollala Ross, p. 65
37a (30b) Incision of outer margin of clasper receptacle angled (Fig. 80, 92, 97, 134) 38
37b Incision of outer margin of clasper receptacle rounded; large (Fig. 112,
150), through small (Fig. 74), to minute (Fig. 124) 41
38a (37a) Inner opening of clasper receptacle, in lateral aspect, visible (Fig. 80, 92) 39
38b Inner opening of clasper receptacle not visible in lateral aspect, rounded
(Fig. 97, 134) 40
39a (38a) Incision of clasper receptacle margin directed anterad (Fig. 80)
C. pasella Ross, p. 38
39b Incision of clasper receptacle margin directed dorsad (Fig. 92)
C. campy la Ross, p. 38
40a (38b) Anterior end of clasper receptacle margin extended antero-ventrad across
lateral face of segment X as fine, sinuate, dark line (Fig. 134)
C.h. harwoodi Denning, p. 54
40b No such continuation of clasper receptacle margin (Fig. 97)
C. ela Denning, p. 39
41a (37b) Inner opening of clasper receptacle visible in lateral aspect (Fig. 74, 150) 42
41b Inner opening of clasper receptacle not visible in lateral aspect (Fig. 1 12, 124) 43
42a (41a) Inner opening of clasper receptacle as wide as chimney (Fig. 150)
C. oxa Ross, p. 64
42b Inner opening narrower than chimney (Fig. 74) C. lasia Ross, p. 32
43a (4 lb) Marginal incision of clasper receptacle margin narrow, minute; anterior
end of margin associated with fine, dark, sinuate line just anterad (Fig.
124) C. helma Ross, p. 54
43b Marginal incision of clasper receptacle wide, of medium size; no fine, dark
line associated with anterior end of margin (Fig. 112)
C. smithi Gordon, p. 48
Gordon (1974) divides this genus into sordida and gracilis complexes, which are included in
the above key. These complexes are each further subdivided to species groups. Other than
presenting the species included here in the order of complexes, species groups, and species used
by Gordon, no further details are given regarding them. It is considered that the minutiae
involved are beyond the scope of the present work.
THE SORDIDA COMPLEX
THE SORDIDA GROUP
Cheumatopsyche minuscula (Banks)
Map 7; Fig. 44-49
Hydropsyche minuscula Banks, 1907:130; Milne, 1936:73 (as synonym of C. sordida).
Hydropsychodes minuscula-, Carpenter, 1933:43; Betten, 1934:195.
Cheumatopsyche minuscula-, Ross, 1 938c: 1 5; Denning, 1943:142; Ross, 1944:1 10; Gordon, 1974:127.
Cheumatopsyche montrealensis Nimmo, 1966a:689; Gordon, 1974:127.
Description. — Male fore-wing length 7.72 mm; red-brown, without evident pattern. Hind-wing paler than
fore-wing, but distinctly tinted. Antennae greyish brown; each flagellar annulus with oblique, faint, darker band around
mid-point. Vertex dark red-brown; warts slightly paler, surrounded by very dark boundary. Spurs dark brown; lateral
member of middle and hind-leg pairs slightly smaller than mesal companions. Thorax deep red-brown dorsally, to paler
Quaest. Ent., 1987, 23 (1)
28
Nimmo
laterally. Legs straw-coloured, with tarsal articles darker.
Genitalia. Male. (Fig. 44-47). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec - Holotype of C.
montrealensis Nimmo). Males distinguished by distal lobes of tergum X small, hooked dorsad in lateral aspect (Fig. 44);
by blunt, rounded distal article of clasper in lateral aspect (Fig. 44); and by long, thin, distally bulbous process emergent
from beneath tergum X (Fig. 44, 47).
Genitalia. Female. (Fig. 48-49). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Females
distinguished by minute, circular or elliptical clasper receptacle high on lateral face of segment X, in lateral aspect (Fig.
49).
Biology. — Available records indicate that larvae of this species prefer fair-sized to
extremely large, turbulent, rubble-bottomed rivers. Flight dates range from June 6 to
September 9, peaking in late June and July.
Distribution. — To date the species has been recorded from Manitoba and the lower St
Lawrence River in the north, to Oklahoma and Georgia in the south (Map 7). In Canada it is
now known from eastern Manitoba to the Saguenay River of Quebec, but most records are
from southern Quebec and Ontario.
Cheumatopsyche sordida (Hagen)
Map 8; Fig. 50-55
Hydropsyche sordida Hagen, 1861:290; Milne, 1936:70, 72, 73.
Hydropsychodes sordida\ Ulmer, 1 905b: 1 00; Betten, 1934:196.
Cheumatopsyche sordida', Ross, 1 938c: 1 5; Denning, 1943:142; Ross, 1944:110; Gordon, 1974:126.
Description. — Male fore-wing length 6.86 mm; uniform rich dark brown. Hind-wing uniformly, palely tinted deep
brown. Antennae deep brown; basal five flagellar annuli with oblique, darker bands. Female with six banded annuli.
Vertex very dark brown. Spurs brown; lateral member of middle and hind-leg pairs notably shorter than mesal
companions, in males; not so in females. Thorax very dark brown, to red-brown laterally. Legs yellow-brown.
Genitalia. Male. (Fig. 50-53). (Specimen from He Ste Helene, St Lawrence R., Montreal, Quebec). Males
distinguished by short, blunt distal article of clasper, in lateral aspect (Fig. 50); by widely separated distal lobes of tergum
X in posterior aspect (Fig. 53), with dorso-lateral angles toothed; and by distal portion of aedeagus phalotheca deeply
keeled ventrally (Fig. 52).
Genitalia. Female. (Specimen from He Ste Helene, St Lawrence R., Montreal, Quebec). Females differentiated by
very small, triangular clasper receptacle, in lateral aspect (Fig. 55), located well dorsad on lateral face of segment X and
with outer margin as part of boundaryline between anterior darker part of segment, and posterior lighter part.
Biology. — Available records indicate that larvae of this species inhabit a wide variety of
flowing waters in Canada, ranging from small to very large rivers, and from slow-flowing
boreal waters to fast, turbulent rivers. Flight dates range from June 7 to August 14, with
possible peak about end of June, early July.
Distribution. — Recorded from Manitoba to New Brunswick in the north, to Texas and
Georgia in the south (Map 8). In Canada it is recorded from Lake Winnipeg to southern New
Brunswick, with most records being from the Ottawa River drainage.
Arctopsychidae and Hydropsychidae (Trichoptera)
29
Map 7. Collection localities for Cheumatopsyche minuscula (Banks) in Canada, with known distribution in North
America by state or province.
Map 8. Collection localities for Cheumatopsyche sordida (Hagen) in Canada, with known distribution in North America
by state or province.
Quaest. Ent., 1987,23 (1)
30
Nimmo
Fig. 44-49, Cheumatopsyche minuscula (Banks): 44, genital capsule of male, lateral aspect; 45, left clasper of male,
posterior aspect; 46, aedeagus of male, lateral aspect; 47, segment X of male, posterior aspect; 48, genital segments of
female, dorsal aspect; 49, genital segments of female, lateral aspect, inf, inferior appendage (clasper); pr, preanal
appendage; cr, clasper receptacle; vs, vulval scale.
Arctopsychidae and Hydropsychidae (Trichoptera)
31
Fig. 50-55, Cheumatopsyche sordida (Hagen): 50, genital capsule of male, lateral aspect; 51, left clasper of male,
posterior aspect; 52, aedeagus of male, lateral aspect; 53, segment X of male, posterior aspect; 54, genital segments of
female, dorsal aspect; 55, genital segments of female, lateral aspect, cr, clasper receptacle.
Quaest. Ent., 1987,23 (1)
32
Nimmo
THE GRACILIS COMPLEX
THE SPECIOSA GROUP
Cheumatopsyche speciosa (Banks)
Map 9; Fig. 56-62
Hydropsyche speciosa Banks, 1 904a:2 1 4; Milne, 1936:71, 73.
Hydropsychodes speciosa] Neave, 1929:190; Betten, 1934:197.
Cheumatopsyche speciosa-, Ross, 1938c; 15; Denning, 1943:154; Ross, 1944:114; Gordon, 1974:134.
Description. — Male fore-wing length 5.54 mm; grey-brown with scattered irroration and larger areas of hyaline
membrane which give transverse banded appearance. Antennae yellowish brown; basal five flagellar annuli each with
slightly darker, oblique band (not evident in female). Vertex reddish brown. Spurs yellow; members of each pair essentially
equal. Thorax deep reddish brown, to yellowish brown laterally. Legs straw-coloured.
Genitalia. Male. (Fig. 56-59). (Specimen from Whitemud Ck, Ellerslie, Alberta). Males distinguished by deep cleft
between dorsum of segment IX and tergum X, in lateral aspect (Fig. 56); by distal lobes of segment X adjacent to this
cleft, and posterior extremity of tergum X flared dorso-laterad as pair of flanges (Fig. 56, 59); and by long, slender distal
article of clasper, more or less linear in posterior aspect (Fig. 57), sinuate, hooked dorsad in lateral aspect (Fig. 56).
Genitalia. Female. (Fig. 60-62). (Specimen from Whitemud Ck, Ellerslie, Alberta). Females distinguished by
postero-lateral angle of each half of sternite VIII (Fig. 62) with small, triangular process; by inner opening of clasper
receptacle not visible in lateral aspect (Fig. 60), receptacle of more or less uniform width; and by outer margin of clasper
receptacle with anterior extremity higher than posterior extremity.
Biology. — This species is recorded from small, sluggish streams, and the largest, turbulent,
rubble-bottomed rivers. Flight dates range from July 6 to August 29, with a diffuse peak in late
June and July.
Distribution. — Recorded from Alberta to Labrador in the north, to Oklahoma and South
Carolina in the south (Map 9). Canadian records are thinly scattered from east of the Rocky
Mountain Foothills in Alberta, to northern Quebec and Labrador in the east, and south to the
United States border.
Cheumatopsyche pinaca Ross
Map 10; Fig. 63-68
Cheumatopsyche pinaca Ross, 1941:82; Ross, 1944:294; Gordon, 1974:133.
Description. — Male fore-wing length 5.75 mm; pale brown; faintly irrorate. Antennae orange-brown; basal five
flagellar annuli each with oblique, darker band. Vertex dark brown; warts paler. Spurs brownish yellow; lateral member of
middle leg pairs notably shorter than mesal companions. Thorax orange-brown. Legs straw-coloured.
Genitalia. Male. (Fig. 63-66). (Specimen from southern Appalachians, USA). Males distinguished by distal lobes of
tergum X, in lateral aspect, short, rounded (Fig. 63); by extreme distal edge of tergum X flared dorso-laterad as pair of
triangular flanges [distal lobes not visible in posterior aspect (Fig. 66)]; by distal article of clasper short, sharply but
smoothly tapered distad, hooked dorsad in lateral aspect (Fig. 63); and by distal end of aedeagus slightly keeled ventrally
(Fig. 65).
Genitalia. Female. (Fig. 67-68). (Specimen from southern Appalachians, USA). Females differentiated by lateral
corner of each half of sternite VIII with short, rounded process (Fig. 68); by sclerotised band of vulval scale very wide,
saddle-like across dorsum of scale; by clasper receptacle, in lateral aspect, papillate; and by anterior end of clasper
receptacle outer margin much higher than posterior end.
Biology. — Neves (1979) records flight period in Massachusetts as June to August.
Distribution. — This species, not yet recorded from Canada, is recorded from Maine to
Florida in the United States, west to Tennessee (Map 10).
Cheumatopsyche lasia Ross
Map 11; Fig. 69-74
Cheumatopsyche lasia Ross, 1938b:154; Denning, 1943:153; Ross, 1944:114; Gordon, 1974:132.
Arctopsychidae and Hydropsychidae (Trichoptera)
33
Description. — Male fore-wing length 4.3 mm; uniform golden brown. Hind-wing palely tinted golden brown.
Antennae yellow-brown; with at least basal six flagellar annuli each with oblique, darker band. Vertex deep red-brown;
warts rather paler. Spurs yellow; lateral member of mid-leg pairs notably shorter than mesal companions. Thorax very
deep red-brown, to paler grey-brown laterally. Legs yellow-brown.
Genitalia. Male. (Fig. 69-72). (Specimen from Red Deer R., Drumheller, Alberta). Males distinguished by segment
IX with distinct dorsal lobes (Fig. 69); by distal lobes of Tergum X projected well dorsad of main body of tergum X, in
lateral aspect; and by these distal lobes with distinct basal processes, best seen in posterior aspect (Fig. 72).
Genitalia. Female. (Fig. 73-74). (Specimen from Brazos R., Palo Alto Co., Texas, USA). Females distinguished by
clasper receptacle slightly expanded distally (Fig. 74); by inner opening visible in lateral aspect, narrower than chimney of
receptacle, directed postero-dorsad; by outer margin of receptacle incised, incision small, rounded, directed antero-dorsad;
by anterior end of receptacle outer margin higher at anterior end than posterior end; and by sclerotised strap of vulval scale
broad, saddle-like across dorsal area of scale.
Biology. — Ross (1944) records the Illinois flight period as May to August, peaking in July
and August. Available records from western Canada are for mid-July. It appears that adults
emerge from the slower, less turbulent creeks and large rivers.
Distribution. — Recorded from Mexico north to Alberta and Saskatchewan in Canada, and
east to Illinois (Map 11). In western Canada it is recorded only from the South Saskatchewan
River drainage.
Map 9. Collection localities for Cheumatopsyche speciosa (Banks) in Canada, with known distribution in North America
by state or province.
Quaest. Ent., 1987,23 (1)
34
Nimmo
Map 10. Known distribution of Cheumatopsyche pinaca Ross in North America, by state.
Map 11. Collection localities for Cheumatopsyche lasia Ross in Canada, with known distribution in North America by
state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
35
Fig. 56-62, Cheumatopsyche speciosa (Banks): 56, genital capsule of male, lateral aspect; 57, left clasper of male,
posterior aspect; 58, aedeagus of male, lateral aspect; 59, segment X of male, posterior aspect; 60, genital segments of
female, lateral aspect; 61, genital segments of female, dorsal aspect; 62, left half of sternite VIII of female, lateral aspect,
cr, clasper receptacle.
Quaest. Ent., 1987, 23 (1)
36
Nimmo
Fig. 63-68, Cheumatopsyche pinaca Ross: 63, genital capsule of male, lateral aspect; 64, left clasper of male, posterior
aspect; 65, aedeagus of male, lateral aspect; 66, segment X of male, posterior aspect; 67, genital segments of female, dorsal
aspect; 68, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
37
Fig. 69-74, Cheumatopsyche lasia Ross: 69, genital capsule of male, lateral aspect; 70, left clasper of male, posterior
aspect; 71, aedeagus of male, lateral aspect; 72, segment X of male, posterior aspect; 73, genital segments of female, dorsal
aspect; 74, genital segments of female, lateral aspect.
Quaest. Ent., 1987,23 (1)
38
Nimmo
THE CAMPYLA GROUP
Cheumatopsyche pasella Ross
Map 12; Fig. 75-80
Cheumatopsyche pasella Ross, 1941:84; Denning, 1943:144; Ross, 1944:1 13; Gordon, 1974:131.
Description. — Male fore-wing length 6.16 mm; chocolate-brown, not irrorate; with large paler areas near
pterostigma, and about distal portions of Cu2 and A. These paler areas absent from female. Hind-wing tinted pale
yellow-brown. Antennae uniformly brown except basal five flagellar annuli with oblique, darker bands; darker in female.
Vertex dark chocolate-brown; darker in female. Spurs pale grey-brown; except latero-apical spur of fore-leg hyaline, finer,
shorter than mesal companion. Female fore-leg latero-apical spur not hyaline, but finer, shorter than mesal companion.
Thorax dark chocolate-brown, to deep red-brown laterally. Legs pale red-brown.
Genitalia. Male. (Fig. 75-78). (Specimen from Huberdeau, Quebec). Males distinguished by darkly sclerotised
cross-shaped pattern which links segment IX with tergum X (Fig. 75); by distal article of clasper very thin in lateral
aspect, curved dorsad (Fig. 76); and by thin, high distal lobes of tergum X with deep, narrow cleft between them and main
body of tergum X (Fig. 75) - these lobes close to each other in posterior aspect.
Genitalia. Female. (Fig. 79-80). (Specimen from Huberdeau, Quebec). Females differentiated by clasper receptacle,
in lateral aspect (Fig. 80), slightly tapered distally; by evident inner opening; by angular incision of clasper receptacle
outer margin directed antero-dorsad; and by small, thin, distally little-widened sclerotised strap of vulval scale.
Biology. — Neves (1979) records from Massachusetts give a flight season of May to August,
no peak mentioned. Ross (1944) indicates that this species prefers faster streams; his two
Illinois records indicate that larvae also inhabit larger, slow-flowing rivers. Gordon & Wallace
(1975) give major larval habitat as fallen trees and branches in flowing waters.
Distribution. — Presently recorded from Oregon, in a narrow zone across northern United
States, then east to Quebec and Maine, and south to Oklahoma and Florida (Map 12). In
Canada it is known only from southern Quebec and southern Ontario.
Cheumatopsyche mickeli Denning
Map 13; Fig. 81-86
Cheumatopsyche mickeli Denning, 1942:50; Ross, 1944:294; Gordon, 1974:132.
Description. — Male fore-wing length 6.86 mm; pale grey-brown, no pattern. Antennae yellow-brown; basal five
flagellar annuli each with oblique, dark band; basal four in female. Vertex dark chocolate-brown, warts paler. Spurs
brown; lateral member of middle and hind-leg pairs shorter than mesal companions. Thorax dark brown, to paler laterally.
Legs pale reddish brown.
Genitalia. Male. (Fig. 81-84). (Specimen from Twenty-mile Ck, Lake Co., Oregon, USA). Males distinguished by
preanal appendage tall, narrow, bowed anterad (Fig. 81); by distal article of clasper tapered fairly abruptly distad,
recurved, in lateral aspect (Fig. 82); and by distal lobes of tergum X with small, squat process anteriorly on base of lobe
(Fig. 81, 84).
Genitalia. Female. (Fig. 85-86). (Specimen from Twenty-mile Ck., Lake Co., Oregon, USA). Females distinguished
by clasper receptacle set very high on lateral wall of segment X (Fig. 86); by inner opening evident in lateral aspect; by
incision of outer margin largish, rounded; and by sclerotised strap of vulval scale inverted-triangular in lateral aspect (Fig.
86), saddle-like over dorsum of scale, not in contact with basal angle of segment X.
Biology. — Anderson’s few records for Oregon indicate a flight season from at least early
June to early September. Nothing more known.
Distribution. — Not yet known from Canada, this species is patchily recorded from Mexico
to Oregon, Idaho, and Wyoming (Map 13).
Cheumatopsyche campyla Ross
Map 14; Fig. 87-92
Cheumatopsyche campyla Ross, 1938b: 152; Denning, 1943:148; Ross, 1944:113; Gordon, 1974:130; Schmid,
1980:144-149.
Arctopsychidae and Hydropsychidae (Trichoptera)
39
Description. — Male fore-wing length 8.35 mm; light grey-brown, with general faint irroration. Hind-wing faintly
tinted brown (stronger tint in female). Antennae pale yellow-brown; basal five flagellar annuli each with oblique, dark
band. Vertex very dark brown. Spur pairs of middle leg with lateral members shorter than mesal companions. Thorax dark
reddish brown, to dull orange-brown laterally (chocolate-brown in female). Legs pale brown to straw.
Genitalia. Male. (Fig. 87-90). (Specimen from Wandering R., Hwy 63, 3 miles S of Wandering River, Alberta).
Males distinguished by very evident dorsal lobes of segment IX, in lateral aspect (Fig. 87); by tall, clavate distal lobes of
tergum X, in lateral aspect (Fig. 87); by distal lobes of tergum X close together in posterior aspect (Fig. 90); and by gently
tapered black band dorsally on segment IX, curved slightly postero-ventrad.
Genitalia. Female. (Fig. 91-92). (Specimen from Wandering R., Hwy 63, 3 miles S of Wandering River, Alberta).
Females distinguished by inner opening of clasper receptacle evident in lateral aspect (Fig. 92); by anterior and posterior
ends of outer margin of clasper receptacle at same level; and by incision of that margin large, angular, directed posterad.
Biology. — As indicated by distribution, this species appears to be ubiquitous in its habitat
preferences. It is recorded from the depths of the Boreal Forest, to deep in the heart of Texas. I
have Canadian records which range from creeks to the largest rivers, some of which are deep,
smooth-flowing waters, others are turbulent. Flight season, based on Canadian records, ranges
from May 5 to Sept. 18 (Ontario) and October 12 (both Nova Scotia and Vancouver Island).
The species may be bivoltine in the southern United States. Larvae are most commonly found
out of the main current, in the backwaters, etc. Ross (1944) indicates that larvae of this species
are very tolerant of pollution.
Distribution. — Virtually throughout the Continent, south of the northern tree line (Map
14). In Canada it has been recorded from south-central British Columbia, northern Alberta,
northern Quebec, Labrador, Newfoundland, and points south.
Cheumatopsyche ela Denning
Map 1 5; Fig. 93-98
Cheumatopsyche ela Denning, 1942:50; Ross, 1944:294; Gordon, 1974:130.
Description. — Male fore-wing length 7.72 mm; overall pale red-brown. Hind-wing very palely tinted brown, with
anal lobe uniform pale grey-brown. Antennae dark brown; basal five flagellar annuli with oblique, dark band. Vertex dark
chocolate-brown. Spurs straw-coloured. Thorax very dark chocolate-brown, to very dark reddish brown laterally. Legs
warm reddish brown.
Genitalia. Male. (Fig. 93-96). (Specimen from St Hippolyte, Quebec). Males distinguished by very distinct dorsal
lobes of segment IX, in lateral aspect (Fig. 93); by distal lobes of tergum X clearly separated from main body of tergum X
by shallow, rounded notch; by distal lobes, in posterior aspect, close together (Fig. 95); and by tergum X clearly delimited
from segment IX by marginal declivity of segment IX.
Genitalia. Female. (Fig. 97-98). (Specimen from St Hippolyte, Quebec). Females distinguished by anterior end of
clasper receptacle outer margin lower than posterior end (Fig. 97); by outer margin with incision angular, directed dorsad;
by receptacle directed antero-dorsad; and by inner opening of receptacle not evident in lateral aspect.
Biology. — Records few, but flight season in eastern United States ranges from April to
July. Roy & Harper (1979) give known flight season in southern Quebec as June 25 to August
7. Little seems to be known of stream types favoured by larvae.
Distribution. — Scattered records from Tennessee and South Carolina to southern Quebec
(Map 15). To date Canadian records are all from the Ottawa-St Lawrence rivers drainage.
Quaest. Ent., 1987, 23 (1)
40
Nimmo
Map 12. Collection localities for Cheumatopsyche pasella Ross in Canada, with known distribution in North America by
state or province.
Map 13. Known distribution of Cheumatopsyche mickeli Denning in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
41
Map 14. Collection localities for Cheumatopsyche campyla Ross in Canada, with known distribution in North America by
state or province.
Map 15. Collection localities for Cheumatopsyche ela Denning in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987, 23 (1)
42
Nimmo
Fig. 75-80, Cheumatopsyche pasella Ross: 75, genital capsule of male, lateral aspect; 76, left clasper of male, posterior
aspect; 77, aedeagus of male, lateral aspect; 78, segment X of male, posterior aspect; 79, genital segments of female, dorsal
aspect; 80, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
43
Fig. 81-86, Cheumatopsyche mickeli Denning: 81, genital capsule of male, lateral aspect; 82, left clasper of male,
posterior aspect; 83, aedeagus of male, lateral aspect; 84, segment X of male, posterior aspect; 85, genital segments of
female, dorsal aspect; 86, genital segments of female, lateral aspect.
Quaest. Ent., 1987,23 (1)
44
Nimmo
Fig. 87-92, Cheumatopsyche campyla Ross: 87, genital capsule of male, lateral aspect; 88, left clasper of male, posterior
aspect; 89, aedeagus of male, lateral aspect; 90, segment X of male, posterior aspect; 91, genital segments of female, dorsal
aspect; 92, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
45
Fig. 93-98, Cheumatopsyche ela Denning: 93, genital capsule of male, lateral aspect; 94, left clasper of male, posterior
aspect; 95, segment X of male, posterior aspect; 96, aedeagus of male, lateral aspect; 97, genital segments of female, lateral
aspect; 98, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
46
Nimmo
THE WABASHA GROUP
Cheumatopsyche wabasha Denning
Map 16; Fig. 99-102
Cheumatopsyche wabasha Denning, 1947:252; Gordon, 1974:130.
Description. — Male fore-wing length 6.7 mm; deep translucent red-brown; very faintly irrorate. Hind-wing paler,
no irroration. Antennae missing; basal segment dark brown. Vertex very dark brown to chocolate. Thorax chocolate to
almost black. Legs dark brown to chocolate.
Genitalia. Male. (Fig. 99-102). (Specimen from Wabasha, Minnesota, USA - Holotype). Gordon used an Oregon
specimen for her illustrations of the male, and description. That specimen was apparently not deformed, as she refers to the
distal deformation of the aedeagus of the male holotype (see Fig. 102 here). She makes no reference to the deformation of
the distal lobes of tergum X as evinced in either lateral or posterior aspects (Fig. 99 & 100 here). Just which, if any, of the
distal lobes borne by the holotype should be regarded as normal for the species is open to doubt. Gordon’s illustrations of
the Oregon specimen seem to show a third type of lobe (posterior aspect). Loss of tergum X distal lobes as reliable
characters is unfortunate, but the holotype, and the Oregon specimens appear to agree in having dorsum of tergum X level,
almost at same height as dorsal lobes of segment IX (Fig. 99). Also, the preanal appendages agree in being located almost
on the base of the distal lobes. The distal article of the clasper is very narrow throughout, in lateral aspect, and sharply
hooked dorsad at distal end; in posterior aspect (Fig. 101) clasper distal article curved posterad, with relatively wide base,
distally acuminate.
Genitalia. Female. Unknown.
Biology. — Unknown except that both Minnesota and Oregon adult records are for July.
Distribution. — Presently known only from Minnesota and Oregon, in the United States
(Map 16).
Map 16. Known distribution of Cheumatopsyche wabasha Denning in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
47
Fig. 99-102, Cheumatopsyche wabasha Denning: 99, genital capsule of male, lateral aspect; 100, segment X of male,
posterior aspect; 101, left clasper of male, posterior aspect; 102, aedeagus of male, lateral aspect.
Quaest. Ent., 1987, 23 (1)
48
Nimmo
THE ROSSI GROUP
Cheumatopsyche logani Gordon & Smith
Map 18; Fig. 103-106
Cheumatopsyche logani Gordon & Smith, 1974:1; Gordon, 1974:128.
Description. — Male fore-wing length 6.94 mm; pale brownish cream; no evident pattern - may be teneral.
Antennae pale straw, no markings on flagellar annuli; scape brown, with cream-coloured areas baso-laterad; pedicel
brown, with posterior area cream. Vertex red-brown, warts almost white. Legs straw-coloured.
Genitalia. Male. (Fig. 103-106). (Specimen from Little Salmon R., Adams Co., Idaho, USA - Holotype). Males
distinguished by distal article of clasper with basal half evenly tapered; distal half uniformly thin, curved dorso-anterad, in
lateral aspect (Fig. 103). Distal article of clasper, in posterior aspect, directed postero-mesad, with tip just visible, directed
mesad (Fig. 105). Dorsum of tergum X sloped postero-ventrad directly from dorsum of segment IX. Preanal appendage
globular, close to base of tergum X distal lobes. Distal lobes of tergum X separated from main body of tergum by shallow,
narrowly v-shaped notch; close together in posterior aspect (Fig. 104).
Genitalia. Female. Unknown.
Biology. — The only flight records available are June 3 and 29, in Washington and Idaho
respectively. Otherwise nothing known.
Distribution. — Presently recorded only from Washington, Idaho, and Montana, in the
United States (Map 18).
Cheumatopsyche smithi Gordon
Map 19; Fig. 107-112
Cheumatopsyche smithi Gordon, 1974:128.
Description. — Male fore-wing length 8.42 mm; warm golden brown; paler areas only at distal end of A. Female
more distinctly irrorate. Antennae brown; basal six flagellar annuli with oblique, dark brown bands; dark yellowish brown
in female. Spurs brownish yellow; lateral member of middle leg pairs notably shorter than mesal companions. Thorax dark
reddish brown, to greyish brown laterally. Legs pale yellowish brown.
Genitalia. Male. (Fig. 107-110). (Specimen from Wandering R., Hwy 63, N of Wandering River, Alberta). Males
distinguished by basal article of clasper very stout, expanded evenly distad, extended dorsad of tergum X; by distal article
of clasper minute by comparison with basal article, tapered distad to fine point, recurved; by distinct dorsal lobes of
segment IX (Fig. 107); by distal lobes of tergum X well separated from main body of tergum X by narrow cleft, in lateral
aspect (Fig. 1 10), with dorsal portion expanded, rounded, flared dorso-laterad; and by small, circular preanal appendages.
Genitalia. Female. (Fig. 111-112). (Specimen from Wandering R., Hwy 63, N of Wandering River, Alberta).
Females distinguished by clasper receptacle expanded distally, in lateral aspect (Fig. 112), with inner end very slightly
cleft; by outer margin of receptacle incised, rounded, incision located at anterior end of margin; by receptacle directed
postero-dorsad; and by sclerotised strap of vulval scale narrow, tapered to fine point distally.
Biology. — Flight season records for Canada range from May 18 to August 12, with
imprecisely defined concentration in June and July. Larvae appear to inhabit a variety of
stream types, from smaller creeks to large rivers, and slower deep waters to fast-flowing waters
on gravel or boulder beds. Records are available from the Vancouver Island rain forest, the
Boreal Forest, prairies, and far southern Ontario.
Distribution. — Recorded to date from Vancouver Island and southern British Columbia, to
southern Ontario, and three States of the Union adjacent to the Canadian border (Map 19). In
Canada most records are from Alberta.
Cheumatopsyche pettiti (Banks)
Map 17; Fig. 113-118
Hydropsyche pettiti Banks, 1908:265; Milne, 1936:73 (with H. analis Banks as synonym of H. morosa).
Hydropsychodes pettiti', Betten, 1934:195.
Cheumatopsyche pettiti', Knowlton & Harmston, 1938:285; Denning, 1943:145; Ross, 1944:294 (as synonym of C. analis
Quaestiones Entomologicae , Volume 23(1)
Nimmo, A. P. (1987, 1-189).- The adult Arctopsychidae and Hydropsychidae (Trichoptera) of
Canada and adjacent United States
CORRIGENDA
pages
49 “(Fig- 17)” in the last paragraph should read “(Map 17)”.
49-50 Captions for maps are in correct places but maps should be rearranged as
follows:
Quaest. Ent., 1987, 23 (4)
Arctopsychidae and Hydropsychidae (Trichoptera)
49
Banks); Gordon, 1974:127, 143 (C. analis as nomina dubia).
Hydropsyche analis Banks, 1903:243; Milne, 1936:73 (as synonym of H. morosa)\ Ross, 1944:1 12.
Hydropsychodes analis ; Betten, 1934:194.
Description. — Male fore-wing length 7.41 mm; pale grey-brown. Hind-wing hyaline. Antennae brown; basal four
flagellar annuli with oblique, dark band; basal five annuli in female. Vertex deep red-brown. Spurs brown; lateral member
of fore-leg pair finer than mesal companion, hyaline; lateral member of middle leg sub-apical pair shorter than mesal
companion. Thorax deep red-brown, to paler laterally. Legs dark straw-coloured.
Genitalia. Male. (Fig. 113-116). (Specimen from R. Maskinonge, Ste Angele, Quebec). Males distinguished by distal
lobes of tergum X very well separated from main body of tergum X, in lateral aspect (Fig. 113); by distal lobes of tergum
X, in posterior aspect (Fig. 116), not particularly close to each other, widened distally, with distal ends flared somewhat
laterad; by dorsal lobes of segment IX clearly evident; and by distal article of clasper more or less confluent with basal
article, in posterior aspect (Fig. 1 14).
Genitalia. Female. (Fig. 117-118). (Specimen from R. Maskinonge, Ste Angele, Quebec). Females distinguished by
clasper receptacle directed antero-dorsad, in lateral aspect (Fig. 1 18); by outer margin of receptacle not incised, continued
anterad on lateral face of segment X by thin, black line; by inner opening of clasper receptacle not evident in lateral aspect;
and by sclerotised strap of vulval scale located well away from segment X, basally thin, distally greatly expanded to poorly
sclerotised area which extends from dorsal to ventral regions of scale.
Biology. — Anderson (1976) summarizes present knowledge of the species. Larvae appear to
prefer smaller streams, but are also recorded from larger rivers. First adults to emerge are very
dark, followed by successively lighter individuals as season progresses. Flight season based on
Canadian records ranges from May 1 1 to October 13, with no pronounced peak.
Distribution. — Known from across North America (Fig. 17), and from northern reaches of
the Boreal Forest south to Texas. Apparently not yet recorded from the southeastern United
States. In Canada this species is known from northern British Columbia, the western and
eastern shores of Hudson’s Bay, Newfoundland, and points south to the United States border.
It appears that this species has also been introduced to Hawaii.
Map 17. Collection localities for Cheumatopsyche pettiti (Banks) in Canada, with known distribution in North America
by state or province.
Quaest. Ent., 1987, 23 (1)
50
Nimmo
Map 18. Known distribution of Cheumatopsyche logani Gordon in North America, by state.
Map 19. Collection localities for Cheumatopsyche smithi Gordon in Canada, with known distribution in North America
by state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
51
Fig. 103 106, Cheumatopsyche logani Gordon: 103, genital capsule of male, lateral aspect; 104, segment X of male,
posterior aspect; 105, left clasper of male, posterior aspect; 106, aedeagus of male, lateral aspect.
Quaest. Ent., 1987, 23 (1)
52
Nimmo
Fig. 107-112, Cheumatopsyche smithi Gordon: 107, genital capsule of male, lateral aspect; 108, left clasper of male,
posterior aspect; 109, aedeagus of male, lateral aspect; 110, segment X of male, posterior aspect; 111, genital segments of
female, dorsal aspect; 1 12, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
53
Fig. 113-118, Cheumatopsyche pettiti (Banks): 113, genital capsule of male, lateral aspect; 114, left clasper of male,
posterior aspect; 115, aedeagus of male, lateral aspect; 116, segment X of male, lateral aspect; 117, genital segments of
female, dorsal aspect; 118, genital segments of female, lateral aspect.
Quaest. Ent., 1987,23 (1)
54
Nimmo
THE HELM A GROUP
Cheumatopsyche helma Ross
Map 20; Fig. 119-124
Cheumatopsyche helma Ross, 1939:68; Ross; 1944:294; Gordon, 1974:135.
Description. — Male fore-wing length 5.03 mm; overall warm, deep red brown; no evident pattern. Antennae
brown; basal five flagellar annuli each with oblique, faintly darker band. Vertex uniform dark brown. Spurs dark brown;
lateral member of middle leg pairs shorter than mesal companions; lateral member of front leg apical pair minute, hyaline
(normal in female). Thorax dark brown, to dull grey-brown laterally. Legs brownish yellow.
Genitalia. Male. (Fig. 119-122). (Specimen from Pineville, Kentucky, USA - Paratype). Males distinguished by
dorsal lobes of segment IX (Fig. 119); by small preanal appendage about mid-point along length of tergum X; by distal
lobes of tergum X, in lateral aspect, triangular, antero-dorsal corner directed anterad; by distal lobes of tergum X, in
posterior aspect, clearly separated by higher, intermediate, angular roof of tergum (Fig. 120); by distal lobes with
acuminate dorso-lateral lobes, in posterior aspect, directed laterad; by segment IX tall, narrow in lateral aspect, with no
posterior projection of postero-lateral edge (compare Fig. 119 & 113); and by distal article of clasper much thinner than
basal article, almost straight in lateral and posterior aspect (Fig. 119, 121), with little taper.
Genitalia. Female. (Fig. 123-124). (Specimen from Gatlinburg, Tennessee, USA). Females distinguished by very
large clasper receptacle, in lateral aspect (Fig. 124), curved dorso-posterad, of uniform width; by inner opening of clasper
recptacle not evident in lateral aspect; by outer margin of receptacle incised at anterior end - incision minute, narrow,
short, directed dorsad; by anterior end of outer margin of receptacle continued by, but not connected to, thin, dark line
across lateral face of segment X; and by large sclerotised strap of vulval scale, with dorsal area of sclerotisation also
present.
Biology. — Blickle & Morse (1966) record adult collection dates from July 8 to 30, in
Maine. Nothing more known at present.
Distribution. — Presently known only from Tennessee, Kentucky, and Maine, in the USA
(Map. 20).
Cheumatopsyche wrighti Ross
Map 21; Fig. 125-128
Cheumatopsyche wrighti Ross, 1947:140; Gordon, 1974:136.
Description. — Male fore-wing length 7.87 mm; uniform warm red-brown; no evident pattern. Antennae brown;
basal five flagellar annuli each with oblique, dark band. Vertex deep red-brown, warts paler. Spurs straw-coloured; lateral
spurs of all pairs on middle and hind legs noticeably shorter than mesal companions. Thorax rich, deep red-brown, to
partly greyish brown laterally. Legs light red-brown, to paler distally.
Genitalia. Male. (Fig. 125-128). (Specimen from Camp Ck, Greene Co., Tennessee, USA - Holotype). Males
distinguished by distal lobes of tergum X very large, appressed anterad along lateral face of tergum X, in lateral aspect
(Fig. 125); by distal lobes of tergum X with posterior edge, in lateral aspect, shouldered; by total lack of dorsal lobes of
segment IX; by distal lobes of tergum X close in posterior aspect (Fig. 126), tapered dorso-mesad; and by recurved distal
article of clasper in posterior aspect (Fig. 127).
Genitalia. Female. Unknown.
Biology. — Neves (1979) records adults from Massachussetts in June- July. The only
Canadian records are from July 5 and 11. Nothing else presently known.
Distribution. — In the United States this species is recorded from Tennessee and the
northeastern seaboard states (Map 21). In Canada there are two records: from Baddeck, Cape
Breton Island, Nova Scotia, and from near Dundee in the eastern half of Prince Edward Island.
Cheumatopsyche h. harwoodi Denning
Map 22; Fig. 129-134
Cheumatopsyche harwoodi Denning, 1949:41.
Cheumatopsyche h. harwoodi ; Gordon, 1974:135.
Arctopsychidae and Hydropsychidae (Trichoptera)
55
Of the two subspecies recognised by Gordon (1974) C. h. harwoodi is the one most likely to
be recorded from Canada.
Description. — Male fore-wing length 6.79 mm; grey-brown. Hind-wing palely tinted grey-brown. Antennae dark
brown; basal six flagellar annuli each with oblique, dark band. Vertex chocolate; posterior warts paler. Spurs dark brown;
lateral member of middle leg pairs much shorter than mesal companions. Thorax chocolate, to mixed chocolate and paler
laterally. Legs dull yellowish brown.
Genitalia. Male. (Fig. 129-132). (Specimen from Credit R., Belfountain, Halton Co., Ontario). Males distinguished
by lack of dorsal lobes on segment IX (Fig. 129); by preanal appendage vertically aligned, small, narrowly elliptical; by
small, rounded notch between main body of tergum X and distal lobes; by distal lobes of tergum X, in lateral aspect (Fig.
129), slightly higher than tergum X, widest dorsally; by distal lobes of tergum X, in posterior aspect (Fig. 131),
moderately separated, with dorso-lateral corners right-angled, and dorso-mesal corners rounded, produced slightly dorsad;
and by basal article of clasper, in posterior aspect (Fig. 130), distally curved gently mesad, with distal half much wider -
distal article of clasper with base narrower than distal end of basal article, gently curved dorsad, tapered to thin, rounded
tip.
Genitalia. Female. (Fig. 133-134). (Specimen from Credit R., Belfountain, Halton Co., Ontario). Females
distinguished by clasper receptacle of medium size, directed dorso-anterad in lateral aspect (Fig. 134), without inner
opening evident; by outer margin of receptacle with deep, angular incision directed dorsad; by anterior end of outer margin
continued antero-ventrad across lateral face of segment IX by thin, black, sinuate line; by sclerotised strap of vulval scale
short, very thin; and by cercus of segment XI very small, short, thin, located immediately at base of dorsal lobe of segment
XI.
Biology. — Flight season data very scarce in literature, but McElravy & Foote (1978)
record possible females from Ohio on August 5. Denning’s original material from Tennessee
was collected on June 6. I have two records from eastern Canada - July 11 and 16. Nothing
else known.
Distribution. — Known from most eastern states of the Union, north of Florida and
Alabama, as far as Maine (Map 22). In Canada the species has been taken in Nova Scotia and
Prince Edward Island.
Map 20. Known distribution of Cheumatopsyche helma Ross in North America, by state.
Quaest. Ent., 1987,23 (1)
56
Nimmo
Map 21. Collection localities for Cheumatopsyche wrighti Ross in Canada, with known distribution in North America by
state or province.
Map 22. Collection localities for Cheumatopsyche h. harwoodi Denning in Canada, with known distribution in North
America by state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
57
Fig. 1 19-124, Cheumatopsyche helma Ross: 1 19, genital capsule of male, lateral aspect; 120, segment X of male, posterior
aspect; 121, left clasper of male, posterior aspect; 122, aedeagus of male, lateral aspect; 123, genital segments of female,
dorsal aspect; 124, genital segments of female, lateral aspect.
Quaest. Ent., 1987, 23 (1)
58
Nimmo
Fig. 125-128, Cheumatopsyche wrighti Ross: 125, genital capsule of male, lateral aspect; 126, segment X of male,
posterior aspect; 127, left clasper of male, posterior aspect; 128, aedeagus of male, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera) 59
Fig. 129-134, Cheumatopsyche h. harwoodi Denning: 129, genital capsule of male, lateral aspect; 130, left clasper of
male, posterior aspect; 131, segment X of male, posterior aspect; 132, aedeagus of male, lateral aspect; 133, genital
segments of female, dorsal aspect; 134, genital segments of female, lateral aspect.
Quaest. Ent., 1987, 23 (1)
60
Nimmo
THE GRACILIS GROUP
Cheumatopsyche gracilis (Banks)
Map 23; Fig. 135-140
Hydropsyche gracilis Banks, 1899:216; Milne, 1936:73 (as synonym of H. morosa).
Hydropsychodes gracilis, Betten, 1934:197.
Cheumatopsyche gracilis-, Ross, 1938c: 15; Denning, 1943:152; Ross, 1944:294; Gordon, 1974:137.
Description. — Male fore-wing length 6.16 mm; pale yellow-brown, with faint, scattered pattern. Antennae
yellow-brown; basal five flagellar annuli each with oblique, dark brown band. Spurs yellow-brown; lateral member of
middle leg pairs notably shorter than mesal companions. Thorax dark brown, to paler laterally. Legs pale yellow-brown.
Genitalia. Male. (Fig. 135-140). (Specimen from Waterton R., nr Standoff, Alberta). Males distinguished by lack of
dorsal lobes on segment IX (Fig. 135); by tergum X sloped postero-ventrad; by preanal appendage large, ovoid; by distal
lobes of tergum not separated from tergum X by gap, in lateral aspect (Fig. 135), antero-dorsal corner angular, turned
slightly dorsad; by distal lobes of tergum X, in posterior aspect (Fig. 137), clearly separated, roughly triangular, with
dorso-lateral corners directed dorso-laterad; and by entire clasper, in posterior aspect (Fig. 136), recurved as unit, without
apparent distinction between basal and distal articles.
Genitalia. Female. (Fig. 139-140). (Specimen from Waterton R., nr Standoff, Alberta). Females distinguished by
massive clasper receptacle (Fig. 140); outer margin of receptacle sinuate, produced ventrad as large, rounded lobe; by
anterior end of outer margin of receptacle continued shortly antero-ventrad as black line; and by receptacle as whole
located very high in segment IX, almost to dorsal crest.
Biology. — Recorded from northern reaches of Boreal Forest, south almost to Texas.
Available records indicate that larvae inhabit a wide range of stream types, from small to very
large, slow to fast and turbulent. Flight season ranges from at least May 12 to August 30 in
Canada, with possible peak in June-July.
Distribution. — Trans-continental, recorded from Boreal Forest south to Utah, Arkansas,
and North Carolina (Map 23). In Canada this species is recorded from south-central British
Columbia to Labrador and Nova Scotia.
Cheumatopsyche vannotei Gordon
Map 24; Fig. 141-144
Cheumatopsyche vannotei Gordon, 1974:138.
Description. — Male fore-wing length 7.02 mm; yellowish brown, evenly irrorate. Hind-wing palely tinted brown.
Antennae brown basally, to light yellow-brown distally. Vertex brown, warts slightly paler. Thorax brown, to pale
yellowish brown laterally. Legs straw to cream-coloured.
Genitalia. Male. (Fig. 141-144). (Specimen from East White Clay Ck, rt. 926, South Chester Co., Pennsylvania,
USA - Holotype). Males distinguished by great elongation, posterad, of all parts of genital capsule, in lateral aspect (Fig.
142, 144); by distal lobes of tergum X, in lateral aspect, almost circular, large; by small, knob-like preanal appendage well
anterad of distal lobe of tergum X; by distal lobes of tergum X, in posterior aspect (Fig. 141), really quite small, well
separated; and by basal article of clasper sufficiently long to extend dorsad of tergum X.
Genitalia. Female. Unknown.
Biology. — Judging from type locality data it appears that larvae favour at least small
streams. Collection date of single known specimen August 3.
Distribution. — Presently known only from Pennsylvania, USA (Map 24).
Arctopsychidae and Hydropsychidae (Trichoptera)
61
Map 23. Collection localities for Cheumatopsyche gracilis (Banks) in Canada, with known distribution in North America
by state or province.
Map 24. Known distribution of Cheumatopsyche vannotei Gordon in North America, by state.
Quaest. Ent., 1987, 23 (1)
62
Nimmo
Fig. 135-140, Cheumatopsyche gracilis (Banks): 135, genital capsule of male, lateral aspect; 136, left clasper of male,
posterior aspect; 137, segment X of male, posterior aspect; 138, aedeagus of male, lateral aspect; 139, genital segments of
female, dorsal aspect; 140, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
63
Fig. 141-144, Cheumatopsyche vannotei Gordon: 141, segment X of male, posterior aspect; 142, genital capsule of male,
lateral aspect; 143, left clasper of male, posterior aspect; 144, aedeagus of male, lateral aspect.
Quaest. Ent., 1987, 23 (1)
64
Nimmo
THE APHANTA GROUP
Cheumatopsyche oxa Ross
Map 25; Fig. 145-150
Cheumatopsyche oxa Ross, 1 938b: 155; Denning, 1943:147; Ross, 1944:110; Gordon, 1974:140.
Description. — Male fore-wing length 7.1 mm; grey-brown, with uniform faint irroration. Hind-wing pale grey.
Antennae brown, without oblique, dark band on any flagellar annulus. Vertex dark chocolate-brown. Spurs pale brown;
lateral member of middle leg pairs, and of hind-leg apical pair, notably shorter than mesal companions. Thorax dark
chocolate-brown. Legs pale brown to yellow.
Genitalia. Male. (Fig. 145-148). (Specimen from creek, Hwy 932, S of Whitecourt, Alberta). Males distinguished by
dorsal lobes present on segment IX (Fig. 145); by no gap between tergum X and distal lobes; by distal lobes, in posterior
aspect (Fig. 148), long, trapezoidal, except distal end slightly expanded, rounded; by distal lobes virtually contiguous; and
by distal article of clasper linear, in posterior aspect (Fig. 146), evenly tapered.
Genitalia. Female. (Fig. 149-150). (Specimen from creek, Hwy 932, S of Whitecourt, Alberta). Females
distinguished by large clasper receptacle, with inner end curved dorsad, of uniform width, in lateral aspect (Fig. 150); by
outer margin of receptacle incised, incision semicircular, directed slightly anterad of dorsal; by inner opening of receptacle
evident, of same width as chimney; and by sclerotised strap of vulval scale acute-triangular, with proximal corner
connected to base of segment X by thin line.
Biology. — Ross (1944) indicates a preference in larvae for small streams, often spring-fed.
This species appears to occur in small, local colonies. Illinois flight dates extend from March to
October. Generally, Canadian locality records support Ross. Canadian flight records extend
from May 17 to September 2, with diffuse peak in July.
Distribution. — Roughly triangular when mapped, with angles in British Columbia, Quebec,
and Georgia (Map 25). The Canadian distribution is from central British Columbia to James
Bay, and the Eastern Townships of Quebec.
Cheumatopsyche aphanta Ross
Map 26; Fig. 151-156
Cheumatopsyche aphanta Ross, 1938b: 1 51; Denning, 1943:151; Ross, 1944:1 1 1; Gordon, 1974:140.
Description. — Male fore-wing length 4.64 mm; red-brown. Hind-wing hyaline with red-brown veins. Antennae
pale red-brown; basal five flagellar annuli each with oblique, dark band. Vertex deep red-brown. Spurs straw-coloured;
lateral member of front and middle leg pairs much shorter than mesal companions. Thorax deep red-brown, to paler
laterally. Legs deep straw-coloured.
Genitalia. Male. (Fig. 151-154). (Specimen from Washington Co., Arkansas, USA). Males distinguished by distinct
dorsal lobes on segment IX (Fig. 151); by distal lobes of tergum X massive in lateral aspect, rounded, not separated from
main body of tergum by gap; by distal article of clasper, in lateral aspect, very much thinner than basal article, with little
taper, hooked sharply dorsad at tip; by distal article of clasper, in posterior aspect, with base almost equal in width to basal
article, acute-triangular, with slight distal curve; and by distal lobes of tergum X, in posterior aspect (Fig. 153), not very
close, spindle-shaped.
Genitalia. Female. (Fig. 155-156). (Specimen from Washington Co., Arkansas, USA). Females distinguished by
clasper receptacle outer margin produced ventrad, and postero-dorsad, as large ventral lobe and posterior flange
respectively, extended posterad of posterior edge of segment X (Fig. 156); by chimney of receptacle inclined slightly
posterad of dorsal, without inner opening visible; and by sclerotised strap of vulval scale expanded evenly, gradually,
distad, strap curved, not in contact with base of segment X.
Biology. — Canadian flight season records extend from June 21 to July 20. Ross (1944)
records the species as common adjacent to small streams and brooks, especially those which are
permanent and spring-fed. Illinois flight season extends from May to late September.
Distribution. — From North Dakota to New York, south to Arkansas in the USA (Map 26).
In Canada this species is presently known only from northeastern New Brunswick and the
Eastern Townships of Quebec.
Arctopsychidae and Hydropsychidae (Trichoptera)
65
Cheumatopsyche halima Denning
Map 27; Fig. 157-162
Cheumatopsyche halima Denning, 1948:400; Gordon, 1974:141.
Description. — Male fore-wing length 6.36 mm; uniform grey-brown. Hind-wing tinted pale grey-brown.
Antennae pale brown; basal five flagellar annuli each with oblique, darker band. Vertex very dark chocolate, to
black-brown. Spurs straw; lateral member of front leg pair much smaller than mesal companion. Thorax very dark
chocolate to black-brown, to slightly paler laterally. Legs dark grey-brown.
Genitalia. Male. (Fig. 157-160). (Specimen from St Hippolyte, Quebec). Males distinguished by lack of dorsal lobes
on segment IX (Fig. 157); by very small, v-shaped notch between tergum X and distal lobes; by distal lobes of tergum X, in
posterior aspect (Fig. 159), triangular, close to each other; and by distal article of clasper, in posterior aspect (Fig. 158),
distinct from basal article, width sharply reduced at junction of the two - distal article curved dorso-laterad.
Genitalia. Female. (Fig. 161-162). (Specimen from St Hippolyte, Quebec). Females distinguished by clasper
receptacle, in lateral aspect (Fig. 162), curved dorso-posterad, with inner opening visible; and by outer margin of
receptacle developed postero-ventrad as large, rounded lobe which partly overlaps posterior edge of segment X.
Biology. — Massachusetts flight season given by Neves (1979) as June to August. Available
Canadian records give a range of June 21 to July 25. Habitat information rare, but larvae are
known to inhabit small streams to small rivers.
Distribution. — Known from Arkansas northeastern states of USA (Map 27). In Canada
this species is recorded from southern Quebec and New Brunswick.
Cheumatopsyche mollala Ross
Map 28; Fig. 163-168
Cheumatopsyche mollala Ross, 1941:81; Ross, 1944:294; Gordon, 1974:142.
Description. — Male fore-wing length 6.47 mm; pale grey-brown, faintly irrorate posterad of Cul+2, and along
costal margin. Hind-wing uniform grey. Antennae brown; basal seven flagellar annuli each with oblique, dark band.
Vertex deep chocolate-brown, warts paler. Spurs yellowish; lateral member of middle leg apical pair, and hind-leg apical
pair, shorter than mesal companions; in female, only laterals of middle leg shorter than mesals. Thorax deep
chocolate-brown throughout; paler laterally in female; warts paler. Legs straw-coloured.
Genitalia. Male. (Fig. 163-166). (Specimen from Lobster Ck, 15 miles SW of Alsea, Benton Co., Oregon, USA).
Males distinguished by small, distinct dorsal lobes on segment IX (Fig. 163); by tergum X slightly humped dorsad in
lateral aspect; by distal lobes of tergum X dorsally acuminate, triangular, without gap between them and main body of
tergum X; by distal lobes, in posterior aspect (Fig. 165), triangular - dorsal angle of each acuminate, directed
dorso-laterad; and by distal article of clasper, in posterior aspect (Fig. 164), with basal three quarters stout, directed
mesad, and distal quarter thin, acuminate, directed dorsad.
Genitalia. Female. (Fig. 167-168). (Specimen from Lobster Ck, 15 miles SW of Alsea, Benton Co., Oregon, USA).
Females distinguished by distinctive, bell-shaped clasper receptacle, in lateral aspect (Fig. 168); by inner portion of
chimney directed dorsad; and by inner opening of recepta le not visible.
Biology. — The sole Canadian record is from May 26. Anderson (1976) gives a flight season
for Oregon from late May to early September, with no definable peak. It is unclear what types
of flowing waters the larvae may favour.
Distribution. — Very spotty (Map 28). Known from Oregon, Idaho, California, and
Arkansas in United States, and eastern Ontario in Canada.
Cheumatopsyche burksi Ross
Map 29; Fig. 169-175
Cheumatopsyche burksi Ross, 1941:83; Ross, 1944:1 13; Gordon, 1974:142.
Description. — Male fore-wing length 7.45 mm; grey-brown; faintly, uniformly irrorate. Antennae yellow-brown;
basal six flagellar annuli each with oblique, faintly darker band; five in female. Vertex deep red-brown, warts slightly
paler. Spurs brown; lateral member of middle and hind-leg pairs noticeably shorter than mesal companions. Thorax deep
red-brown, to yellow-brown laterally. Legs yellow-brown to straw.
Quaest. Ent., 1987, 23 (1)
66
Nimmo
Genitalia. Male. (Fig. 169-172). (Specimen from Tavares, Lake Co., Florida, USA - Paratype). Males distinguished
by distinct dorsal lobes on segment IX (Fig. 169); by small distal lobes of tergum X, in lateral aspect, hardly separable
from main body of tergum; by distal article of clasper with wide basal third surmounted by aristate, dorsally curved distal
two-thirds, in lateral aspect; by distal article of clasper, in posterior aspect (Fig. 170), much as above, except junction
between basal and distal portions more gradual; and by distal lobes of tergum X, in posterior aspect (Fig. 172), well
separated, expanded dorsal portion ovoid.
Genitalia. Female. (Fig. 173-175). (Specimen from Tavares, Lake Co., Florida, USA - Allotype). Females
distinguished by medium-sized clasper receptacle bell-like, but skewed anterad in lateral aspect (Fig. 174); and by
posterior edge of each half of sternite VIII irregular (Fig. 175), with small, triangular process slightly higher than
mid-point.
Biology. — The only flight date available is October 2, in Illinois (Ross, 1944). Nothing
more known at present.
Distribution. — Presently known only from Illinois to Louisiana and Florida, USA (Map
29).
Cheumatopsyche enonis Ross
Map 30; Fig. 176-181
Cheumatopsyche enonis Ross, 1938b: 153; Ross, 1944:294; Gordon, 1974:142.
Cheumatopsyche geolca Denning, 1952:21; Gordon, 1974:142.
Description. — Male fore-wing length 4.99 mm; grey, almost hyaline. Hind-wing hyaline. Antennae brown. Vertex
dark brown. Spurs yellow; lateral member of middle leg pairs shorter than mesal companions. Thorax dark red-brown, to
paler laterally. Legs yellow.
Genitalia. Male. (Fig. 176-179). (Specimen from Dale Ck, Richland, Oregon, USA). Males distinguished by very
prominent dorsal lobes of segment IX (Fig. 176); by small, tapered distal lobes of tergum X directed postero-dorsad in
lateral aspect; by distal lobes fairly close, trapezoidal in posterior aspect (Fig. 179); and by conical distal article of clasper
with meso-ventral spur, in posterior aspect (Fig. 177).
Genitalia. Female. (Fig. 180-181). (Specimen from Dale Ck, Richland, Oregon, USA). Females distinguished by
clasper receptacle, in lateral aspect (Fig. 181), located dorso-anterad in segment X; by receptacle directed anterad; by
receptacle inner opening not visible in lateral aspect [if such an opening exists - not visible in dorsal aspect either (Fig.
180)]; and by vulval scale apparently without sclerotised strap.
Biology. — Anderson (1976) has no comment on habitats, but locality names suggest that
larvae frequent creeks and, at least, smaller rivers. Oregon flight season is given as June to
early September.
Distribution. — Recorded from all western cordilleran states of USA, except Washington,
California, and Arizona (Map 30).
Arctopsychidae and Hydropsychidae (Trichoptera)
67
Map 25. Collection localities for Cheumatopsyche oxa Ross in Canada, with known distribution in North America by
state or province.
Map 26. Collection localities for Cheumatopsyche aphanta Ross in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987, 23 (1)
68
Nimmo
Map 27. Collection localities for Cheumatopsyche halima Denning in Canada, with known distribution in North America
by state or province.
Map 28. Collection localities for Cheumatopsyche mollala Ross in Canada, with known distribution in North America by
state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
69
Map 29. Known distribution of Cheumatopsyche burksi Ross in North America, by state.
Map 30. Known distribution of Cheumatopsyche enonis Ross in North America, by state.
Quaest. Ent., 1987,23 (1)
70
Nimmo
Fig. 145-150, Cheumatopsyche oxa Ross: 145, genital capsule of male, lateral aspect; 146, left clasper of male, posterior
aspect; 147, aedeagus of male, lateral aspect; 148, segment X of male, posterior aspect; 149, genital segments of female,
dorsal aspect; 150, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
71
Fig. 151-156, Cheumatopsyche aphanta Banks: 151, genital capsule of male, lateral aspect; 152, left clasper of male,
posterior aspect; 153, segment X of male, posterior aspect; 154, aedeagus of male, lateral aspect; 155, genital segments of
female, dorsal aspect; 156, genital segments of female, lateral aspect.
Quaest. Ent., 1987, 23 (1)
72
Nimmo
Fig. 157-162, Cheumatopsyche halima Denning: 157, genital capsule of male, lateral aspect; 158, left clasper of male,
posterior aspect; 159, segment X of male, posterior aspect; 160, aedeagus of male, lateral aspect; 161, genital segments of
female, dorsal aspect; 162, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
73
Fig. 163-168, Cheumatopsyche mollala Ross: 163, genital capsule of male, lateral aspect; 164, left clasper of male,
posterior aspect; 165, segment X of male, posterior aspect; 166, aedeagus of male, lateral aspect; 167, genital segments of
female, dorsal aspect; 168, genital segments of female, lateral aspect.
Quaest. Ent., 1987,23 (1)
74
Nimmo
Fig. 169-175, Cheumatopsyche burksi Ross: 169, genital capsule of male, lateral aspect; 170, left clasper of male,
posterior aspect; 171, aedeagus of male, lateral aspect; 172, segment X of male, posterior aspect; 173, genital segments of
female, dorsal aspect; 174, genital segments of female, lateral aspect; 175, left half of sternite VIII of female, lateral
aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
75
Fig. 176-181, Cheumatopsyche enonis Ross: 176, genital capsule of male, lateral aspect; 177, left clasper of male,
posterior aspect; 178, aedeagus of male, lateral aspect; 179, segment X of male, posterior aspect; 180, genital segments of
female, dorsal aspect; 181, genital segments of female, lateral aspect.
Quaest. Ent., 1987,23 (1)
76
Nimmo
Genus Hydropsyche Pictet
Map 31-71; Fig. 5, 182-462
Hydropsyche Pictet, 1834:199; McLachlan, 1878:355, 357; Betten, 1934:183; Milne, 1936:68; Denning, 1943:108; Ross,
1944:86; Ross & Unzicker, 1977:310; Wiggins, 1977:106; Schmid, 1980:58.
Symphitopsyche Ulmer, 1907:32; Ross & Unzicker, 1977:304; Schuster & Etnier, 1978:30.
Description. — Tarsal claws similar on all three pairs of legs; twisted asymmetrically; overhung by heavy, black
hairs. Spur formula 2,4,4. Male sternite V lobe with aperture of large internal gland of two cavities; lobe more or less
developed depending on species; gland small, spherical, in female, with only one slightly developed lobe. Cross-veins
M3 + 4-Cul and Cul-Cu2 of fore-wing clearly separated (Fig. 5a). Hind-wing (Fig. 5b) fl petiolate; median cell closed;
stems of M and Cul parallel, very close.
Genitalia. Male. (Fig. 182-186, 304-308, 332-336, etc.). Segment IX with postero-lateral margin developed posterad
(Fig. 189), with long setae (not shown). Segment X simple or complex, massive with specifically characteristic lobes (Fig.
182, 183). Claspers (inferior appendages) with stout, conical distal article. Aedeagus curved ventrad basally (Fig. 185,
307, 335); bilobed distally, simple or complex; comprised of sclerotised phallotheca with tip cleft in species groups 1 (Fig.
186) and 2 (Fig. 308). No distal cleft in species group 3, but short, erectile endotheca comprised of lobes of varied
complexity (Fig. 335, 336); these lobes variously armed with specifically distinct spines and teeth.
Genitalia. Female. (Fig. 187-188, 309-310, 337-338, etc.). Tergite VIII notched on mid-line, with slightly recurved
lateral angles in some species. Sternite VIII divided apically to two lobes. Segment X with postero-lateral margin produced
as blunt, setate lobe (Fig. 187); with clasper receptacle or lateral depression on dorso-lateral face of segment.
Biology. — Larvae of Hydropsyche spp. inhabit a wide range of flowing waters, from largest
rivers to spring-fed streamlets, with specific restriction to some lesser range of types. Larvae of
H. alternans (Walker) and H. confusa (Walker) have also been reported from wave- washed
shores of larger lakes. Larvae ingest algae, detritus, and animal matter in seasonally
determined proportions.
Hydropsyche , the largest genus of the family, is found in all regions but the Neotropical and
Antarctica. Wiggins (1977) estimates that about 70 species are now known from North
America. Forty-two of these are presented here, most of which are recorded only from east of
the western Cordillera.
Key to known or potential species of Hydropsyche Pictet of Canada
la Males (Fig. 182-186) 2
lb Females (Fig. 187-188) 42
2a (la) Aedeagus entirely sclerotised; without membranous lobes, spines, or teeth
distally (Fig. 185,307) 3
2b Aedeagus with membranous lobes distally (Fig. 335); with or without
spines and/or teeth (Fig. 335, 452, 459) 24
3a (2a) Tip of aedeagus tubular in lateral aspect, truncated at right angles to long
axis; slightly expanded, rounded in some species (Fig. 307, 314, 321)
species group 2 p. 1 18 4
3b Tip of aedeagus, in lateral aspect, bluntly or sharply wedge-shaped, with
dorsally flared lateral flanges (Fig. 185, 191, 202, 209)
species group 1 p. 83 7
4a (3a) Aedeagus, in lateral aspect, with proximal end curved ventrad in
semi-circle, directed posterad (Fig. 307) H. betteni Ross, p. 1 18
4b Aedeagus, in lateral aspect, with proximal end bent only slightly
antero-ventrad (Fig. 314, 321, 328) 5
5a (4b) Distal article of clasper, in posterior aspect, curved smoothly mesad,
expanded distally (Fig. 313) H. confusa (Walker), p. 1 18
Arctopsychidae and Hydropsychidae (Trichoptera)
77
5b Distal article of clasper, in posterior aspect, not curved mesad; narrowed
distally (Fig. 320, 327)
6a (5b) Distal article of clasper, in lateral aspect, smoothly tapered distad, with
rounded tip turned dorsad (Fig. 325) H. depravata Hagen, p. 1 19
6b Distal article of clasper, in lateral aspect, irregularly tapered distad, with
acuminate tip turned slightly dorsad (Fig. 318) H. cuanis Ross, p. 1 19
7a (3b) Cleft in aedeagus tip, in dorsal aspect, simple, v-shaped, without notches or
widenings part-way along sides; cleft long or short, wide or narrow (Fig.
192, 197, 203, 217, 224, 231, 245, 252)
7b Cleft in aedeagus tip not simple v-shape; variously modified from v-shape
(Fig. 186,210,238, 280,294)
8a (7b) Distal article of clasper with tip curved dorsad in lateral aspect (Fig. 206,
276,290)
8b Distal article of clasper with tip not curved dorsad in lateral aspect (Fig.
182,234)
9a (8a) Distal article of clasper, in posterior aspect, blunt (Fig. 292). Gap between
distal lobes of segment X, in dorsal aspect, wide, approximately v-shaped
(Fig. 291) H. valanis Ross, p. 90
9b Distal article of clasper, in posterior aspect, with disto-lateral process (Fig.
208,278)
10a (9b) Cleft in aedeagus tip wide in dorsal aspect (Fig. 210)
H. bidens Ross, p. 84
10b Cleft in aedeagus tip very narrow, with slight widening at inner end (Fig.
280) H. scalaris Hagen, p. 89
11a (8b) Distal lobes of segment X, in lateral aspect (Fig. 182), directed
postero-dorsad, long, narrow H. aerata Ross, p 83
lib Distal lobes of segment X directed posterad, short, triangular (Fig. 234) . .
H. hageni Banks, p. 86
12a (7a) Distal article of clasper, in lateral aspect, with tip transversely truncate to
greater or lesser degree (Fig. 220, 241, 261, 297)
12b Distal article of clasper, in lateral aspect, with disto-ventral corner of tip
produced dorsad, or entire tip curved dorsad (Fig. 189, 194, 199, 213, 227,
248, 255, 269, 283)
13a (12a) Distal lobes of segment X, in dorsal aspect, separated by broad, flat,
v-shaped notch (Fig. 242, 263)
13b Distal lobes of segment X, in dorsal aspect, separated by narrow, deep, v-
or u-shaped notch (Fig. 221, 297)
14a (13a) Distal article of clasper, in lateral aspect, massive, rounded, widened distad
(Fig. 241) H. occidentalis Banks, p. 87
14b Distal article of clasper, in lateral aspect, small, trapezoidal, with slight
projection of disto-ventral corner (Fig. 262) H. placoda Ross, p. 88
15a (13b) Segment X with blade-like process on each lateral surface directed dorsad
(Fig. 220) H. dicantha Ross, p. 85
15b No such process on segment X (Fig. 297) H. venularis Banks, p. 90
16a (12b) Distal lobes of segment X, in dorsal aspect, separated by v-shaped notch
(wide or narrow) (Fig. 190, 195, 200, 249)
6
12
8
9
11
10
13
16
14
15
17
Quaest. Ent., 1987, 23 (1)
78
Nimmo
16b Distal lobes of segment X, in dorsal aspect, separated by notch of varied
shapes except V (Fig. 214, 228, 270, 284) 20
17a (16a) Distal article of clasper with disto-ventral corner, in lateral aspect,
produced as pointed lobe (Fig. 194, 248) 18
17b Distal article of clasper with entire distal end as pointed lobe directed
dorsad (Fig. 189, 199) 19
18a (17a) Tip of aedeagus, in dorsal aspect, cleft deeply, widely (Fig. 197)
H. leonardi Ross, p. 86
18b Tip of aedeagus, in dorsal aspect, with very shallow cleft continued basad
as thin line of closure (Fig. 252) H. orris Ross, p. 87
19a (17b) Segment IX with anterior edge of dorsal concavity well posterad of anterior
edge of segment (Fig. 189) H. alvata Denning, p. 83
19b Segment IX dorsal concavity very close to anterior edge of segment (Fig.
199) H. arinale Ross, p. 84
20a (16b) Notch between distal lobes of segment X, in dorsal aspect, narrowed at
distal opening (Fig. 270, 284) 21
20b Notch between distal lobes of segment X not so narrowed, open (Fig. 214,
228,256) 22
21a (20a) Notch circular (Fig. 284) H. simulans Ross, p. 90
21b Notch elliptical (Fig. 270) H. rossi Flint, Voshell, & Parker, p. 89
22a (20b) Notch composite (Fig. 214) H. calif ornica Banks, p. 84
22b Notch simple (Fig. 228, 256) 23
23a (22b) Notch roughly rectangular (Fig. 256). Distal lobes of segment X long,
narrowed, directed postero-dorsad in lateral aspect (Fig. 255)
H. phalerata Hagen, p. 88
23b Notch u-shaped (Fig. 228). Distal lobes of segment X short, triangular in
lateral aspect (Fig. 227) H.frisoni Ross, p. 85
24a (2b) Distal end of aedeagus with membranous lobes dorsally and
ventro-laterally (Fig. 335, 342, 349, 356, 377, 384, 391, 398, 405) 25
24b Distal end of aedeagus with dorsal membranous lobes only (Fig. 363, 412,
435,452) 33
25a (24a) Ventro-lateral lobes of aedeagus reduced to small papillae or simple holes
in lateral face of distal extremity of aedeagus (Fig. 377, 384, 391, 398,
405) subgroup C p. 138 26
25b Ventro-lateral lobes long, positioned along side of aedeagus, directed
anterad (Fig. 335, 342, 349, 356) subgroup A p. 126 30
26a (25a) Ventro-lateral lobes represented by single lateral hole in sclerotised lateral
wall of distal extremity of aedeagus (Fig. 391, 398) 27
26b Ventro-lateral lobes membranous, very short, in same location as in 26a above 28
27a (26a) Tooth of dorsal lobe massive, with dentate anterior edge (Fig. 381)
H. morosa Hagen, p. 139
27b Tooth of dorsal lobe minute, simple (Fig. 398)
H. slossonae Banks, p. 139
28a (26b) Tooth of dorsal lobe small, directed posterad (Fig. 405)
H. tana Ross, p. 1 40
Tooth of dorsal lobe large, longer than wide (Fig. 377, 384)
28b
29
Arctopsychidae and Hydropsychidae (Trichoptera)
29a (28b) Distal article of clasper, in lateral aspect, triangular (Fig. 374)
H. bronta Ross, p. 138
29b Distal article of clasper, in lateral aspect, rather like clenched hand with
index finger pointing (Fig. 381) H. cheilonis Ross, p. 138
30a (25b) Ventro-lateral lobe of aedeagus with origin at extreme distal end of
aedeagus (Fig. 342, 349)
30b Ventro-lateral lobe of aedeagus with origin ventrad of endotheca (Fig. 335, 353) .
31a Ventro-lateral lobe of aedeagus without distal cluster of spines (Fig.
342) H. piatrix Ross, p. 126
31b Ventro-lateral lobe of aedeagus with distal cluster of spines (Fig. 349) ....
H. vexa Ross, p. 1 27
32a (30b) Distal lobe of segment X, in lateral aspect (Fig. 332), massive, blunt; in
dorsal aspect (Fig. 333) almost linear H. amblis Ross, p. 126
32b Distal lobe of segment X, in lateral aspect (Fig. 353), thin, angled; in
dorsal aspect (Fig. 354), distal ends of lobe curved mesad, gap between
partly enclosed H. walkeri Betten & Mosely, p. 127
33a (24b) Dorsal lobes membranous only, without teeth or spines (Fig. 452,
459) subgroup F p. 160
33b Dorsal lobes membranous, with teeth and/or spines (Fig. 363, 412, 438)
34a (33a) Distal article of clasper, in lateral aspect (Fig. 449), long, tapered from
wide base to slender tip H. oslari Ross, p. 160
34b Distal article of clasper, in lateral aspect (Fig. 456), shorter, blunt, not
tapered H. ventura Ross, p. 160
35a (33b) Dorsal lobe of aedeagus with one large tooth, and separate cluster of spines
(Fig. 439, 445) subgroup E p. 156
35b Dorsal lobe with single tooth only (Fig. 363, 412)
36a (35a) Tooth at distal extremity of lobe; spine cluster laterally (Fig. 438, 439) . . .
H. riola Denning, p. 156
36b Spine cluster at distal extremity; tooth laterally (Fig. 445, 446)
H. sparna Ross, p. 156
37a (35b) Distal tooth of dorsal lobe of aedeagus curved dorsad (Fig. 412, 419, 426,
433) subgroup D p. 148
37b Distal tooth curved or directed ventrad (Fig. 363, 370)
subgroup B p. 134
38a (37a) Distal lobes of segment X, in lateral aspect, curved ventrad (Fig. 409, 416) . . .
38b Distal lobes straight, directed postero-dorsad (Fig. 423, 430)
39a (38a) Distal lobes of segment X, in dorsal aspect (Fig. 410), linear
H. alternans (Walker), p. 148
39b Distal lobes, in dorsal aspect (Fig. 416), curved postero-mesad
H. centra Ross, p. 1 48
H. aenigma Schefter, Wiggins, & Unzicker, p. 148
40a (38b) Distal lobes of segment X, in dorsal aspect (Fig. 424), directed
postero-laterad H. cockerelli Banks, p. 149
40b Distal lobes curved slightly postero-mesad (Fig. 431)
H. jewetti Denning, p. 1 49
79
31
32
34
35
36
37
38
41
39
40
Quaest. Ent., 1987, 23 (1)
80
Nimmo
41a (37b) Distal tooth of dorsal lobe of aedeagus, in lateral aspect (Fig. 363), with
basal quarter wide, distal three-quarters abruptly narrowed to acuminate
spine H. alhedra Ross, p. 134
41b Distal tooth tapered uniformly distad, small, minutely dentate (Fig. 370) .
H. bifida Banks, p. 134
42a (lb) Clasper receptacle without evident inner opening, either in lateral aspect
(Fig. 204, 218, 239, 246, 274, 309, 323) and/or dorsal aspect (Fig. 205,
219,240,275,310,324) 43
42b Clasper receptacle with clearly evident inner opening in dorsal aspect (Fig.
188, 268, 338, 345), in lateral aspect (Fig. 267, 344) in some species 59
43a (42a) Clasper receptacle, in lateral aspect, invagination of lateral wall of segment
X, without ventral prolongation of anterior end of outer margin as curved
declivity (Fig. 218, 274, 309, 323, 358) 44
43b Clasper receptacle, in lateral aspect, curved declivity in dorsal half of
segment X, with or without invagination at some point (Fig. 225, 232, 295, 302) . . .
. . 52
43c Clasper receptacle simple, elongate groove high on segment X (Fig. 246) .
H. Occident alis Banks, p. 87
44a (43a) Clasper receptacle, in lateral aspect, simple, invaginated pouch (Fig. 211,
218,323,358) 45
44b Clasper receptacle an invaginated pouch, with grooves or lobes within
aperture of pouch (Fig. 274, 316, 330) 48
45a (44a) Clasper receptacle, in lateral aspect, small, located dorsally (Fig. 218), or
anteriorly (Fig. 260) on segment X 46
45b Clasper receptacle large, associated with posterior margin of segment X
(Fig. 323, 358), or anterior margin (Fig. 21 1) 47
46a (45a) Clasper receptacle located dorsally on segment X (Fig. 218)
H. californica Banks, p. 84
46b Clasper receptacle located anteriorly on segment X (Fig. 260)
H. phalerata Hagen, p. 88
47a (45b) Clasper receptacle, in lateral aspect, rounded, tubular invagination of
segment X (Fig. 323) H. cuanis Ross, p. 1 19
47b Clasper receptacle long, curved, shallow invagination of segment X (Fig. 358) . . 48
48a (47b) Clasper receptacle close to anterior margin of segment X (Fig. 211); vulval
scale with two sclerotised straps, one angled H. bidens Ross, p. 84
48b Clasper receptacle close to posterior margin of segment X (Fig. 358);
vulval scale with only one strap H. walkeri Betten & Mosely, p. 127
49a (44b) Clasper receptacle, in lateral aspect, only with simple grooves (2) at mouth
of receptacle (Fig. 274, 330) 50
49b Clasper receptacle with lobes (2) at mouth of receptacle (Fig. 309, 316) 51
50a (49a) Outer margin of clasper receptacle simple, not incised (Fig. 274)
H. rossi Flint, Voshell, & Parker, p. 89
50b Outer margin deeply, irregularly incised (Fig. 330)
H. depravata Hagen, p. 1 19
51a (49b) Outer margin of clasper receptacle, in lateral aspect, with simple, sinuate
incision (Fig. 316). Dorsum of segment X wide in lateral aspect
Arctopsychidae and Hydropsychidae (Trichoptera)
81
H. confusa (Walker), p. 1 18
51b Outer margin with acuminate secondary incision on margn (Fig. 309).
Dorsum of segment X narrow H. betteni Ross, p. 1 18
52a (43b) Dorsum of segment X, in lateral aspect, with (Fig. 211, 239, 253, 288, 295)
or without (Fig. 232) slight indentation; thin, black line with origin at
indentation, extended across dorso-lateral area of segment X toward
clasper receptacle 53
52b Without either indentation or line (Fig. 204, 225, 302) 57
53a (51a) Clasper receptacle, in dorsal aspect, simple, domed depression, without
ventral portion overlapped by dorsal portion (Fig. 212, 233, 254) 54
53b Clasper receptacle with overlap of ventral portion by dorsal, secondary
invagination of segment X wall (Fig. 240, 289, 296) 55
54a (53a)Sclerotised strap of vulval scale simple, thin line (Fig. 232)
H.frisoni Ross, p. 85
54b Sclerotised strap large, width irregular (Fig. 253)
H. orris Ross, p. 87
55a (53b) Outer margin of clasper receptacle, in lateral aspect, with distinct,
acuminate tooth, closer to posterior end of margin (Fig. 295)
H. valanis Ross, p. 90.
55b Outer margin of clasper receptacle simple (Fig. 239, 288) 56
56a (55b) Vulval scale with angular secondary sclerotised strap (Fig. 239)
H. hageni Banks, p. 86
56b Vulval scale without secondary strap (Fig. 288) . . H. simulans Ross, p. 90
57a (52b) Vulval scale with angular secondary sclerotised strap (Fig. 302)
H. venularis Banks, p. 90
57b Vulval scale without secondary strap 58
58a (57b) Clasper receptacle, in lateral aspect (Fig. 225), transverse declivity, with
extremities directed ventrad; posterior extremity with slight invagination
H. dicant ha Ross, p. 85
58b Clasper receptacle with shallow dorsal pouch, continued ventro-posterad by
declivity (Fig. 204) H. arinale Ross, p. 84
59a (42b) Floor of clasper receptacle aperture, in lateral aspect, without grooves or
folds (Fig. 267, 281, 337, 407, 428, 461) 60
59b Floor of clasper receptacle aperture with grooves or folds (Fig. 187, 344,
365,372,379,414,421,454) 65
60a (59a) Clasper receptacle, in lateral aspect, associated with declivity (Fig. 267, 281) . .61
60b Clasper receptacle not associated with declivity (Fig. 337, 407, 428, 461) 62
61a (60a) Clasper receptacle outer opening integrated with declivity (Fig. 281)
H. scalaris Hagen, p. 89
61b Clasper receptacle posterad of declivity; receptacle single, circular hole
(Fig. 267) H. placoda Ross, p. 88
62a (60b) Sclerotised strap of vulval scale small; irregularly narrow, or just thin, dark
line (Fig. 337, 407) 63
62b Sclerotised strap of vulval scale large, long, widest at two-thirds of length
(Fig. 428, 461) 64
63a (62a) Clasper receptacle minute, in lateral aspect; directed dorsad, not associated
Quaest. Ent., 1987,23 (1)
82
Nimmo
with thin, dark line down lateral face of segment X (Fig. 407)
H. tana Ross, p. 140
63b Clasper receptacle small, directed anterad; thin, dark line extended across
mouth of receptacle (Fig. 337) H. amblis Ross, p. 126
64a (62b) Clasper receptacle small, directed dorso-anterad, located low down on
lateral face of segment X (Fig. 428) H. cockerelli Banks, p. 149
64b Clasper receptacle large, oriented vertically, directed anterad, located more
dorsally on segment X (Fig. 461) H. ventura Ross, p. 160
65a (59b) Inner opening of clasper receptacle not visible in lateral aspect (Fig. 187,
421,454) 66
65b Inner opening visible in lateral aspect (Fig. 344, 414) 68
66a (65a) Clasper receptacle, in lateral aspect, with single groove on floor of aperture
(Fig. 187) H. aerata Ross, p. 83
66b Clasper receptacle with two grooves (Fig. 421, 454) 67
67a (66b) Clasper receptacle directed dorsad in lateral aspect (Fig. 421), posterad in
dorsal aspect (Fig. 422) H. centra Ross, p. 148
67b Clasper receptacle directed dorso-anterad in lateral aspect (Fig. 454),
mesad in dorsal aspect (Fig. 455) H. oslari Ross, p. 160
68a (65b) Clasper receptacle outer margin, in lateral aspect, with both ends on same
level (Fig. 344,414) 69
68b Clasper receptacle with outer margin anterior end lower than posterior end
(Fig. 372, 379, 393) 70
68c Clasper receptacle outer margin with anterior end higher than posterior
end (Fig. 351, 365, 400, 440, 447) 73
69a (68a) Clasper receptacle, in lateral aspect, curved dorso-posterad (Fig. 344) ....
H. piatrix Ross, p. 126
69b Clasper receptacle, in lateral aspect, directed dorso-anterad (Fig. 414) ...
H. alternans (Walker), p. 148
70a (68b) Clasper receptacle, in dorsal aspect, curved meso-posterad (Fig. 373, 387) 71
70b Clasper receptacle directed mesad in dorsal aspect (Fig. 380, 394) 72
71a (70a) Outer margin of clasper receptacle, in lateral aspect, angularly incised
(Fig. 372) H. bifida Banks, p. 134
71b Outer margin slightly sinuate, not incised (Fig. 386)
H. cheilonis Ross, p. 138
72a (70b) Outer margin of clasper receptacle, in lateral aspect, continued to either
side by thin, dark line (Fig. 379) H. bronta Ross, p. 138
72b No such thin, dark line present (Fig. 393) H. morosa Hagen, p. 139
73a (68c) Cerci, dorsal and ventral lobes of segment XI, in dorsal aspect, visible (Fig.
366,401,448) 74
73b At most, only dorsal lobes, cerci visible in dorsal aspect (Fig. 352, 441) 76
74a (73a) Anterior end of clasper receptacle outer margin continued ventrad as thin,
dark line (Fig. 400) H. slossonae Banks, p. 139
74b No such continuation of clasper receptacle outer margin (Fig. 365, 447) 75
75a (74b) Posterior end of clasper receptacle outer margin, and outer ends of grooves
on floor of receptacle, approximately in line (Fig. 365)
H. alhedra Ross, p. 1 34
Arctopsychidae and Hydropsychidae (Trichoptera)
83
75b These points not in line (Fig. 447) H. sparna Ross, p. 156
76a (73b) Lobe of postero-lateral margin of segment X small, triangular (Fig. 351)
H. vexa Ross, p. 1 27
76b Lobe long, hemi-elliptical (Fig. 440) H. riola Denning, p. 156
SPECIES GROUP 1
This group is characterised by aedeagus sclerotised throughout, without membranous lobes,
with distal tip wedge-shaped in lateral aspect ( e.g ., Fig. 185).
Hydropsyche aerata Ross
Map 31; Fig. 182-188
Hydropsyche aerata Ross, 1 938b: 1 44; Ross, 1944:101; Schuster & Etnier, 1978:77.
Description.— Male fore-wing length 7.02 mm; light brown; faintly, coarsly irrorate posterad of Cul+2.
Hind-wing hyaline. Antennae red-brown, no markings in male; allotype female with at least three basal flagellar annuli
with dark, oblique bands (remainder of antennae missing). Vertex deep red-brown; narrow in male - eyes very large
relative to head; normal in allotype female. Spurs yellow; lateral member of all pairs notably shorter than mesal
companions. Thorax uniformly deep red-brown. Legs yellow.
Genitalia. Male. (Fig. 182-186). (Specimen from Aroma Park, Kankakee R., Illinois, USA). Males distinguished by
distal lobes of segment X approximately rectangular in lateral aspect (Fig. 182), directed postero-dorsad; by distal article
of clasper, in lateral aspect, relatively little tapered, without distal hook; by distal cleft of aedeagus, in dorsal aspect (Fig.
186), not v-shaped, widened laterad at mid-point; and by distal article of clasper, in posterior aspect (Fig. 184),
parallel-sided, truncate distally.
Genitalia. Female. (Fig. 187, 188). (Specimen from Oakwood, Illinois, USA - Allotype). Females distinguished by
clasper receptacle, in lateral aspect (Fig. 187), simple, broad, rounded invagination with inner opening not visible, with one
groove on receptacle floor; by receptacle, in dorsal aspect (Fig. 188), with ventral portion overlapped by dorsal portion; and
by sclerotised strap of vulval scale proximally acuminate, evenly and considerably widened distally.
Biology. — Poorly known. Ross gives flight season as May to late August. Apparently a
species of large rivers with rapids and gravel bottoms.
Distribution. — Not yet known from Canada. In the USA the species is presently only
recorded from Illinois, Indiana, and Michigan (Map 31).
Hydropsyche alvata Denning
Map 32; Fig. 189-193
Hydropsyche alvata Denning, 1949:40; Schuster & Etnier, 1978:125; Flint, Voshell, & Parker, 1979:842.
Description. — Male fore-wing length 8.38 mm; pale red-brown, veins darker, faintly irrorate about
Cul+2/RS/M and distal edge. Hind-wing faintly tinted warm red-brown. Antennae pale red-brown; basal 7 or 8
flagellar annuli each with oblique, dark band. Vertex dark red-brown. Spurs straw-coloured; lateral member of fore- and
middle leg pairs much shorter than mesal companions. Thorax dark red-brown, to paler laterally. Legs straw-coloured.
Genitalia. Male. (Fig. 189-193). (Specimen from Madison Co., Arkansas, USA). Males distinguished by distal lobes
of segment X blunt, triangular in lateral aspect (Fig. 189), short, rounded, separated by u-shaped notch in dorsal aspect
(Fig. 190); by distal cleft of aedeagus very narrowly v-shaped in dorsal aspect (Fig. 192); and by distal article of clasper, in
lateral aspect (Fig. 189), long, curved slightly dorsad, of even width except for distal taper to upturned tooth.
Genitalia. Female. Unknown.
Biology. — Virtually nothing known. Denning’s (1949) original description provides a range
of flight dates from March 3 to July 18.
Distribution.— Not yet recorded from Canada. In the USA the species is known from
Arkansas to Michigan and Virginia (Map 32).
Quaest. Ent., 1987, 23 (1)
84
Nimmo
Hydropsyche arinale Ross
Map 33; Fig. 199-205
Hydropsyche arinale Ross, 1938b: 1 43; Ross, 1944:104; Schuster & Etnier, 1978:86.
Description. — Male fore-wing length 6.32 mm; pale red-brown, with banded pattern in area of anal lobe, with
distinct darker bar mid-way along anal edge; faintly irrorate along Rl. Hind-wing tinted warm red-brown. Antennae
yellow-brown; basal seven flagellar annuli each with oblique, dark band. Vertex pale red-brown. Spurs yellow-brown;
lateral member of fore- and middle leg pairs shorter than mesal companions. Thorax deep red-brown, to paler laterally.
Legs pale red-brown.
Genitalia. Male. (Fig. 199-203). (Specimen from Washington Co., Arkansas, USA). Males distinguished by distal
lobes of segment X, in lateral aspect (Fig. 199), short, broad, rounded, separated, in dorsal aspect (Fig. 200), by shallow,
wide, v-shaped notch; by distal article of clasper, in lateral aspect, unevenly tapered to dorsally upturned distal hook; and
by distal cleft of aedeagus v-shaped in dorsal aspect, with ventral area not notched (Fig. 203).
Genitalia. Female. (Fig. 204-205). (Specimen from Washington Co., Arkansas, USA). Females distinguished by
clasper receptacle, in lateral aspect (Fig. 204), simple depression on lateral wall of segment X, bounded anteriorly by
distinct declivity, with slight invagination at dorsal end of declivity; by clasper receptacle, in dorsal aspect (Fig. 205),
simple, large, domed depression in lateral wall of segment X; and by sclerotised strap of vulval scale large,
acute-triangular.
Biology. — The only available Canadian flight records are July 21-24, in southern Ontario.
Ross (1944) gives flight records from April to September. He also indicates that larvae prefer
smaller, clear streams of many riffles or rapids. Ross also concludes that this species adheres
quite closely to the western fringes of the Oak-Hickory forest.
Distribution. — Recorded from Oklahoma and Kansas to southern Ontario (with gaps), with
one record from Fraserburg, Ontario (Map 33).
Hydropsyche bidens Ross
Map 34; Fig. 206-212
Hydropsyche bidens Ross, 1 938b: 1 42; Denning, 1943:1 18; Ross, 1944:107; Schuster & Etnier, 1978:75.
Description. — Male fore-wing length 11.08 mm; pale brown, faintly irrorate, with darker areas along Cula-lb
and A. Hind-wing hyaline. Antennae yellow-brown; basal ten flagellar annuli each with oblique, dark band. Vertex
red-brown. Spurs pale brown. Thorax red-brown, to slightly paler laterally. Legs yellow-brown to straw.
Genitalia. Male. (Fig. 206-210). (Specimen from lie Perrot, Quebec). Males distinguished by wide, broadly rounded
distal lobes of segment X, in lateral aspect (Fig. 206); by these lobes, in dorsal aspect (Fig. 207), separated by deep,
narrow cleft with rounded interior expansion; and by distal article of clasper, in lateral aspect, slightly expanded at
mid-point, then tapered sharply to dorsally curved, acuminate tooth (tooth visible in posterior aspect (Fig. 208)).
Genitalia. Female. (Fig. 211-212). (Specimen from Lac Monroe, Parc Mont Tremblant, Quebec). Females
distinguished by vulval scale with two sclerites (Fig. 211), one angled; by clasper receptacle high, ventrally tapered
invagination, without inner opening visible (Fig. 211,212); and by clasper receptacle dorsal end associated with thin, dark
line originated postero-dorsad on segment X.
Biology. — Schuster & Etnier (1978) state that this is a species of larger rivers subject to
heavy silt loading. They give flight season dates as April to September, concluding that there is
only one generation per year.
Distribution. — Recorded from Montana, through Manitoba, to Quebec, and south to Texas
(Map 34). In Canada known only from southern Manitoba and southwestern Quebec.
Hydropsyche californica Banks
Map 35; Fig. 213-219
Hydropsyche californica Banks, 1899:217; Betten, 1934:193; Milne, 1936:73; Ross, 1 938c: 1 6; Denning, 1943:115.
Hydropsyche scalaris Hagen; Milne, 1936:73 (as synonym of H. californica).
Description. — Male fore-wing length 9.44 mm; light chocolate-brown, faintly irrorate with larger hyaline areas
peripherally. Hind-wing faintly tinted grey-brown. Antennae pale brown; basal seven flagellar annuli each with oblique,
dark brown band (eight in female). Vertex very dark purplish brown, warts lighter. Spurs brownish yellow; lateral member
Arctopsychidae and Hydropsychidae (Trichoptera)
85
of middle leg pairs, and of hind-leg apical pair, notably shorter than mesal companions. Thorax very dark purplish brown,
to dark chocolate-brown laterally.
Genitalia. Male. (Fig. 213-217). (Specimen from creek, Hwy 19, 32.5 km S of Kelsay Bay jet., Vancouver Island,
British Columbia). Males distinguished by stout basal article of clasper in lateral aspect (Fig. 213); by distal article of
clasper, in lateral aspect, offset somewhat posterad, of even width, with disto-ventral angle produced bluntly distad; by
clasper, in posterior aspect (Fig. 215), smoothly curved mesad; and especially by distal lobes of tergum X, in dorsal aspect
(Fig. 214), composite, with pair of secondary lobes within primary lobes.
Genitalia. Female. (Fig. 218-219). (Specimen from creek, Flwy 19, 32.5 km S of Kelsay Bay jet., Vancouver Island,
British Columbia). Females distinguished by clasper receptacle small, boomerang-like, set high in lateral wall of segment
X, in lateral aspect (Fig. 218); by lack of sclerotised strap on vulval scale; and by cercus thin, appressed to underside of
dorsal lobe of segment XI, well separated from ventral lobe.
Biology. — Very little known. Two records from Canada available for flight season - one for
July 10, the other simply for August. The July 10 material was taken by myself at a lake outlet
into a middling-sized woodland creek over sandy boulder bottom.
Distribution. — Presently known from the western and northwestern cordilleran United
States, and Vancouver Island, with a record from Minnesota (Map 35). In Canada this species
is presently known only from two localities on Vancouver Island.
Hydropsyche dicantha Ross
Map 36; Fig. 220-226
Hydropsyche dicantha Ross, 1938b: 1 46; Ross, 1944:102; Schuster & Etnier, 1978:80; Flint, Voshell, & Parker, 1979:845.
Description. — Male fore-wing length 7.72 mm; overall warm red-brown. Hind-wing very palely tinted reddish
brown. Antennae deep red-brown; distal half of each flagellar annulus paler. Vertex red-brown. Spurs yellow-brown.
Thorax deep red-brown, to paler laterally. Legs red-brown.
Genitalia. Male. (Fig. 220-224). (Specimen from Lac Monroe, Parc Mont Tremblant, Quebec). Males distinguished
by accuminate, hooked lobes on lateral faces of segment X (from which the species gets its name)(Fig. 220); by distal
article of clasper rectangular in lateral aspect; and by large, sigmoid aedeagus (Fig. 223) with distal cleft narrowly
v-shaped (Fig. 222).
Genitalia. Female. (Fig. 225-226). (Specimen from R. du Diable, Parc Mont Tremblant, Quebec). Females
distinguished by clasper receptacle, in lateral aspect (Fig. 225), extended across whole lateral wall of segment X, with
posterior and anterior ventrally directed arms, comprised mostly of sharp declivity with very small invagination at
postero-dorsal angle; by cercus of segment XI large, acute-triangular, directed postero-dorsad along with dorsal and
ventral lobes; by cerci and associated lobes all visible in dorsal aspect (Fig. 226); and by sclerotised strap of vulval scale
long, slender, widened distally, recurved.
Biology. — Schuster & Etnier (1978) indicate that larvae are fairly indiscriminate in choice
of stream type, being known from small, cool, riffled streams, to large, warm rivers. Flight
season extends from late June to late September. Canadian records are slightly earlier.
Distribution. — Presently known from Minnesota to Tennessee, to New Hampshire and
southern Quebec (Map 36). In Canada, recorded from southern fringes of Canadian Shield in
Ontario and Quebec, south to United States border.
Hydropsyche frisoni Ross
Map 37; Fig. 227-233
Hydropsyche frisoni Ross, 1938b: 142; Ross, 1944:105; Schuster & Etnier, 1978:93.
Description. — Male fore-wing length 8.03 mm; light grey-brown, faintly irrorate overall. Hind-wing hyaline.
Antennae yellow-brown; basal seven flagellar annuli each with oblique, dark band; presence of bands in faded paratype
female uncertain. Vertex deep red-brown. Spurs yellow; lateral member of middle leg pairs shorter than mesal
companions. Thorax deep red-brown. Legs light red-brown.
Genitalia. Male. (Fig. 227-231). (Specimen from Plateau Experimental Stn, Cumberland Co., Tennessee, USA).
Males distinguished by distal article of clasper, in lateral aspect (Fig. 227), of uniform width, linear, with distal end
tapered, curved dorsad as small median hook; by aedeagus, in lateral aspect (Fig. 230), with narrow base, distal half
expanded, robust; by distal cleft of aedeagus narrow, v-shaped (Fig. 231); and by gap between distal lobes of tergum X, in
dorsal aspect (Fig. 228) u-shaped.
Quaest. Ent., 1987,23 (1)
86
Nimmo
Genitalia. Female. (Fig. 232-233). (Specimen from White R., Shoals, Indiana, USA - Paratype). Females
distinguished by clasper receptacle as semi-circular declivity directed anterad, in lateral aspect (Fig. 232), with very slight
invagination at dorsal end; and by sclerotised strap of vulval scale thin, dark line.
Biology. — Schuster & Etnier (1978) conclude that larvae prefer small, warm-water rivers
and are intolerant of habitat alteration. Flight season is given as April through August.
Distribution. — Not yet known from Canada. Recorded from Minnesota to Missouri and
Ohio in the United States (Map 37).
Hydropsyche hageni Banks
Map 38; Fig. 234-240
Hydropsyche hageni Banks, 1 905a: 14; Milne, 1936:73 (as synonym of H. scalaris)-, Denning, 1943:119; Ross, 1944:103;
Schuster & Etnier, 1978:102; Flint, Voshell, & Parker, 1979:847.
Description. — Male fore-wing length 1 1.66 mm; dark grey-brown, faintly irrorate posterad of R1 (distally), with
darker areas along veins. Hind-wing tinted grey-brown. Antennae bright pale red-brown; basal nine flagellar annuli each
with oblique, dark band. Vertex red-brown. Spurs pale brown; lateral member of fore- and middle leg pairs shorter than
mesal companions. Thorax red-brown. Legs pale red-brown.
Genitalia. Male. (Fig. 234-238). (Specimen from Clinch R., Hancock Co., Tennessee, USA). Males distinguished by
basal article of clasper, in lateral aspect (Fig. 234), linear, of almost uniform width; by distal article of clasper
irregular-pentagonal in outline; and by gap between distal lobes of tergum X, in dorsal aspect (Fig. 235), slightly narrower
than interior, gap oval.
Genitalia. Female. (Fig. 239-240). (Specimen from Cahaba R., Bibb Co., Alabama, USA). Females distinguished by
clasper receptacle small, narrow invagination at dorsal end of short declivity on lateral face of segment X (Fig. 239); by
vulval scale with two sclerotised straps - one a fine, dark line, other wider, angled; and by ventral portion of clasper
receptacle, in dorsal aspect (Fig. 240), overlapped by dorsal portion.
Biology. — The few Canadian records give a flight season of May 25 to June 26. Ross
(1944) records flight from May to late August. Larvae appear to prefer faster parts of rivers
over boulders and/or bedrock.
Distribution. — Recorded from Manitoba to Alabama, Virginia, and southeastern Ontario
(Map 38). In Canada presently known only from eastern Manitoba, and southeastern Ontario.
Hydropsyche leonardi Ross
Map 39; Fig. 194-198
Hydropsyche leonardi Ross, 1938b: 145; Ross, 1944:294; Flint, Voshell, & Parker, 1979:851.
Description. — Male fore-wing length 9.67 mm; orange-brown, with darker membrane about Cul +2 and patches
posterad of that vein. Hind-wing palely tinted light orange-brown. Antennae brown; some flagellar annuli with oblique,
dark band, but number uncertain as most of flagellum missing. Vertex dark chocolate-brown. Spurs brown; lateral
member of all pairs shorter than mesal companions. Thorax uniformly dark chocolate-brown. Legs light yellow-brown.
Genitalia. Male. (Fig. 194-198). (Specimen from Au Sable R., Crawford Co., Michigan, USA - Paratype). Males
distinguished by large, splayed distal lobes of aedeagus, in dorsal aspect (Fig. 197); by distal lobes of tergum X separated
by small, v-shaped notch, in dorsal aspect (Fig. 195); and by distal article of clasper, in lateral aspect (Fig. 194), robust,
almost trapezoidal, with disto-ventral angle produced as small tooth, and with lateral face concave (see also Fig. 198).
Genitalia. Female. Unknown.
Biology. — Flint, Voshell, & Parker (1979) provide Virginia flight records which range
from April 12 to October 20. Available Canadian records are May 20 and 29. The above
authors report adult emergence from riffles of clean, fast-flowing larger rivers.
Distribution. — Presently known only from Michigan, Virginia, and southeastern Ontario
(at Ottawa)(Map 39).
Arctopsychidae and Hydropsychidae (Trichoptera)
87
Hydropsyche occidentalis Banks
Map 40; Fig. 241-247
Hydropsyche occidentalis Banks, 1900:258; Betten, 1934:194; Milne, 1936:69, 71, 73; Ross, 1 938c: 17; Ross, 1944:294.
Hydropsyche novamexicana Banks, 1 904b: 1 10; Milne, 1936:73 (as synonym of H. occidentalis).
Description. — Male fore-wing length 8.50 mm; pale brownish yellow with scattered darker areas. Antennae
yellow; at least basal six flagellar annuli each with oblique, dark band. Vertex pale reddish brown; yellow-brown in female.
Spurs straw; lateral member of middle leg preapical pair notably shorter than mesal companion. Thorax pale reddish
brown, to yellowish brown laterally. Legs yellow.
Genitalia. Male. (Fig. 241-245). (Specimen from Lethbridge, Alberta). Males distinguished by basal article of clasper
with irregularly slender basal third; distal two-thirds parallel-sided, much wider in lateral aspect (Fig. 241); by distal
article of clasper robust, in lateral aspect; by same article, in posterior aspect (Fig. 243), with disto-dorsal angle projected
mesad from behind disto-ventral edge; and by gap between distal lobes of tergum X, in dorsal aspect (Fig. 242), wide,
v-shaped, rounded, not angular.
Genitalia. Female. (Fig. 246-247). (Specimen from Lethbridge, Alberta). Females distinguished by clasper receptacle
located very high on lateral wall of segment X - a simple slit (Fig. 246); by sclerotised strap of vulval scale linear, widened
distally, of medium size; and by lobes of segment XI large, stubby, directed slightly postero-ventrad.
Biology. — From my records it seems that adults emerge from all manner of flowing waters,
from small, weed-filled, plains streamlets to largest rivers, and from turbulent, rocky foothills
streams and rivers. Flight season ranges from June 2 to August 19. Simmons et al. (1942)
record larvae as being so numerous in a Sierra Nevada water conduit as to require shut-down
for cleaning.
Distribution. — Widespread throughout western Cordillera from Mexico to British
Columbia, east to Great Plains (at least in Canada) of Saskatchewan (Map 40). In Canada,
recorded from south-central British Columbia to prairies and Boreal Forest of Alberta and
Saskatchewan.
Hydropsyche orris Ross
Map 41; Fig. 248-254
Hydropsyche cornuta Ross, 1 938b: 141 (preoccupied by Martynov, 1909).
Hydropsyche orris Ross, 1 938a: 121 (new name); Denning, 1943:1 18; Ross, 1944:105; Schuster & Etnier, 1978:71.
Description. — Male fore-wing length 8.74 mm; pale red-brown; veins darker, foci for irroration pattern.
Hind-wing tinted grey-brown. Antennae red-brown; basal 8-9 flagellar annuli each with oblique, dark band. Vertex deep
red-brown; paler in available female. Spurs red-brown; lateral member of fore- and middle leg pairs much shorter than
mesal companions. Thorax deep red-brown, to slightly paler laterally; warm red-brown in female. Legs red-brown, to straw
distally.
Genitalia. Male. (Fig. 248-252). (Specimen from Washington Co., Arkansas, USA). Males distinguished by distal
article of clasper, in lateral aspect (Fig. 248), of uniform width, curved slightly dorsad, with disto-ventral angle produced
as acuminate tooth; by gap between distal lobes of tergum X, in dorsal aspect (Fig. 249), not deep, v-shaped; and by distal
cleft of aedeagus (Fig. 252) shallow, v-shaped, continued basad by thin line of closure.
Genitalia. Female. (Fig. 253-254). (Specimen from Vicksburg, Mississippi, USA). Females distinguished by clasper
receptacle small, triangular, in lateral aspect (Fig. 253), dorsally directed invagination of dorsal portion of semi-circular
declivity of lateral wall of segment X; by cerci and dorsal lobes of segment XI, only, visible in dorsal aspect (Fig. 254); and
by sclerotised strap of vulval scale narrow at each end, irregularly widened in middle.
Biology. — Schuster & Etnier (1978) conclude that larvae are adapted to large rivers with
high silt loading and high concentration of suspended organic matter. They also conclude that
the species is univoltine, with flight season from April to October.
Distribution. — Presently known from South Dakota to Texas, Georgia, and Michigan
(Map 41). Not yet known from Canada.
Quaest. Ent., 1987, 23 (1)
88
Nimmo
Hydropsyche phalerata Hagen
Map 42; Fig. 255-261
Hydropsyche phalerata Hagen, 1861:287; Betten, 1934:189; Milne, 1936:73 (as synonym of H. morosa)\ Denning,
1943:1 13; Ross, 1944:102; Schuster & Etnier, 1978:78; Flint, Voshell, & Parker, 1979:853.
Description. — Male fore-wing length 7.72 mm; pale grey-brown, no evident pattern. Hind-wing hyaline, to faintly
tinted. Antennae deep red-brown; basal eight flagellar annuli each with oblique, dark band. Vertex deep brown. Spurs
yellow-brown; lateral member of middle leg pairs shorter than mesal companions. Thorax deep brown, to mixed deep
brown and paler brown laterally. Legs yellow-brown, to straw laterally.
Genitalia. Male. (Fig. 255-259). (Specimen from Mississippi R., Hennepin Co., Minnesota, USA). Males
distinguished by distal lobes of tergum X narrow, directed postero-dorsad in lateral aspect (Fig. 255), with gap between, in
dorsal aspect (Fig. 256); by distal cleft of aedeagus, in dorsal aspect (Fig. 259), v-shaped, narrow, of medium depth; and
by distal article of clasper with dorsal edge straight, ventral edge sinuate, with disto-ventral angle slightly, bluntly
produced in lateral aspect (Fig. 255).
Genitalia. Female. (Fig. 260-261). (Specimen from Mississippi R., Hennepin Co., Minnesota, USA). Females
distinguished by clasper receptacle small, triangular, located very close to anterior edge of segment X, in lateral aspect
(Fig. 260); by ventral lobe of segment XI well ventrad of cercus and dorsal lobe; and by sclerotised strap of vulval scale
thin, dark line deeply bowed ventrad.
Biology. — Flint, Voshell, & Parker (1979) give a flight season range of May 25 to
September 27, in Virginia. Ross (1944) gives the Illinois range as late April to September.
According to Schuster & Etnier (1978) larvae prefer very wide rivers with shallow riffle areas,
with silty gravel and small-to-medium-sized rock bottom, with high suspended organic loading,
and with warm water conditions in late Spring and early Fall.
Distribution. — Though recorded from Kansas, this species is primarily confined east of a
line from Minnesota to Florida (Map 42), as far east as Massachusetts and southern Quebec.
In Canada it is presently known only from southern Quebec and southern Ontario.
Hydropsyche placoda Ross
Map 43; Fig. 262-268
Hydropsyche placoda Ross, 1941:87; Denning, 1943:115; Ross, 1944:103; Schuster & Etnier, 1978:127.
Description. — Male fore-wing length 8.66 mm; tinted translucent brown, with darker colour mostly about R1 and
between Cul +2 to A3. Female rather darker overall. Antennae pale orange-brown; basal eight flagellar annuli each with
oblique, dark band. Vertex orange-brown, markedly narrowed due to considerable enlargement of compound eyes; eyes
normal in female. Spurs yellow; lateral member of middle leg pairs notably shorter than mesal companions. Spur formula
apparently 1,4,4. Thorax orange-brown, to yellow-brown laterally. Legs pale brown to straw.
Genitalia. Male. (Fig. 262-266). (Specimen from Pembina R., Sangudo, Alberta). Males distinguished by distal
article of clasper, in lateral aspect (Fig. 262), rectangular, with distal edge scalloped; by gap between distal lobes of
tergum X v-shaped, wide, shallow in dorsal aspect (Fig. 263); and by distal cleft of aedeagus, in dorsal aspect (Fig. 265),
v-shaped, deep, very narrow.
Genitalia. Female. (Fig. 267-268). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Females
distinguished by clasper receptacle simple, circular pit anterad of long, curved declivity very close to anterior edge of
segment X (Fig. 267).
Biology. — Flight season in Canada ranges from May 25 to September 5. Little more known
at present. The species is recorded from St Lawrence R. at Montreal, where are rapids of a very
large river.
Distribution. — Presently known from Alberta and Montana east to Illinois, New York
State, and Quebec (Map 43). In Canada it is known from both Boreal Forest and prairies of the
three Prairie Provinces, and from St Lawrence R. valley of Ontario and Quebec.
Arctopsychidae and Hydropsychidae (Trichoptera)
89
Hydropsyche rossi Flint, Voshell, & Parker
Map 44; Fig. 269-275
Hydropsyche incommoda (not Hagen): Ross, 1944:106; Schuster & Etnier, 1978:92.
Hydropsyche rossi Flint, Voshell, & Parker, 1979:854.
Description. — Male fore-wing length 9.67 mm; grey-brown, irrorate overall. Hind-wing hyaline. Antennae pale
brown; basal nine flagellar annuli each with oblique, dark band. Vertex yellow-brown; warts darker in female. Spurs with
lateral member of middle leg pairs shorter than mesal companions. Thorax deep red-brown, to deep yellow-brown laterally.
Legs yellow. All warts darker in female.
Genitalia. Male. (Fig. 269-273). (Specimen from Waterford, Marshall Co., Mississippi, USA - Paratype). Males
distinguished by distal article of clasper with ventral edge linear, dorsal edge sinuate, disto-ventral angle with short, curved
tooth (Fig. 269); by postero-dorsal corner of distal lobe of tergum X angled (Fig. 269); and by gap between distal lobes, in
dorsal aspect (Fig. 270), small, narrower at opening, elliptical in outline.
Genitalia. Female. (Fig. 274-275). (Specimen from Waterford, Marshall Co., Mississippi, USA - Paratype). Females
distinguished by clasper receptacle, in lateral aspect (Fig. 274), located dorso-anterad on lateral wall of segment X, small,
with outer margin anterior end much lower than posterior end, with no visible inner opening, with two curved grooves on
floor of receptacle; and by sclerotised strap of vulval scale small, acute-triangular, with thin, dark line from apex to base of
segment X - also, small secondary sclerite dorsad of primary.
Biology. — Flight season ranges from March 23 to September 25 according to Flint,
Voshell, & Parker (1979), with reference to States from Illinois south to Arkansas. If this
species is found in Canada the range may be expected to be rather shortened, especially in
Spring. Little more known.
Distribution.— Not yet known from Canada. Recorded in United States from area bounded
by Missouri to Florida, Virginia, and Illinois (Map 44).
Hydropsyche scalaris Hagen
Map 45; Fig. 276-282
Hydropsyche scalaris Hagen, 1861:286; Betten, 1934:190; Milne, 1936:69, 72, 73; Denning, 1943:112; Ross, 1944:106;
Schuster & Etnier, 1978:87; Flint, Voshell, & Parker, 1979:856.
Description. — Male fore-wing length 11.08 mm; pale reddish brown, faintly irrorate, with darker areas along
veins - especially Cul and A3. Antennae yellow-brown; basal nine flagellar annuli each with oblique, dark band. Vertex
yellow-brown. Spurs pale yellow-brown; lateral member of middle leg pairs, and hind-leg apical pair notably shorter than
mesal companions. Thorax deep orange-brown, to paler laterally. Legs pale brown to straw.
Genitalia. Male. (Fig. 276-280). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Males
distinguished by distal article of clasper, in lateral aspect (Fig. 276), much like knife blade; by distal cleft of aedeagus
deep, narrow, with widening part-way along length, in dorsal aspect (Fig. 280); and by gap between distal lobes of tergum
X, in dorsal aspect (Fig. 277), narrow, slightly widened interiorly.
Genitalia. Female. (Fig. 281-282). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Females
distinguished by clasper receptacle, in lateral aspect (Fig. 281), with sharp declivity ventrad of anterior end of outer
margin of receptacle, with posterior end of margin continued to dorsum of segment X by thin, dark line; by inner opening
of receptacle not evident in lateral aspect, inner end of receptacle curved slightly dorso-posterad; and by ventral portion of
receptacle, in dorsal aspect (Fig. 282), overlapped by dorsal portion - inner opening visible.
Biology. — Canadian flight season records range from June 4 to August 28. In Virginia,
Flint, Voshell, & Parker (1979) recorded adults from May 21 to October 20. Schuster &
Etnier (1978) have little more to report on this species except to conclude that larvae prefer
warmer water streams of various sizes.
Distribution. — Recorded from Colorado and New Mexico to Georgia, Maine, and southern
Quebec (Map 45). In Canada, known primarily from southern Quebec and Ontario, but there
is one isolated record from Duck Mountain area of Manitoba, which is in the Boreal Forest.
Quaest. Ent., 1987, 23 (1)
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Nimmo
Hydropsyche simulans Ross
Map 46; Fig. 283-289
Hydropsyche simulans Ross, 1 938b: 1 39; Denning, 1943:117; Ross, 1944:104; Schuster & Etnier, 1978:90.
Description. — Male fore-wing length 10.37 mm; warm yellowish brown, uniformly irrorate except from distal end
of thyridial cell to fl-f5. Antennae pale yellow-brown; basal ten flagellar annuli each with oblique, dark band; basal nine in
female. Vertex yellow-brown. Spurs yellow; lateral member of middle and hind-leg pairs notably shorter than mesal
companions. Thorax red-brown dorsally except mesal line yellow; laterally yellow-brown. Legs yellow to straw.
Genitalia. Male. (Fig. 283-287). (Specimen from Mt Carmel, Illinois, USA - Paratype). Males distinguished by
distal lobes of tergum X, in lateral aspect (Fig. 283), square; by these distal lobes, in dorsal aspect (Fig. 284), with gap
between circular; and by distal article of clasper like knife blade in lateral aspect.
Genitalia. Female. (Fig. 288-289). (Specimen from Washington Co., Arkansas,USA). Females distinguished by outer
margin of clasper receptacle, in lateral aspect (Fig. 288), continued ventrad by declivity, dorsad by thin, dark line; by inner
end of receptacle acuminate in lateral aspect, directed dorso-posterad; by sclerotised strap of vulval scale thin, very little
widened distally; and by ventral area of clasper receptacle, in dorsal aspect (Fig. 289), slightly overlapped by dorsal area.
Biology. — Ross (1944) records emergence as April to late September. Schuster & Etnier
(1978) conclude that larvae prefer larger rivers (30-60 m width) with boulder and coarse gravel
bottom intermixed with silt. They seem to prefer streams with high organic content.
Distribution. — Known to occur from Montana south to Texas, east to southern Ontario
(Map 46), this species is known in Canada from one locality near Sarnia, Ontario.
Hydropsyche valanis Ross
Map 47; Fig. 290-296
Hydropsyche valanis Ross, 1938b: 144; Ross, 1944:105; Schuster & Etnier, 1978:85.
Description. — Male fore-wing length 8.35 mm; warm red-brown, no evident pattern; faintly irrorate in female.
Antennae yellow-brown; basal seven flagellar annuli each with oblique, dark band. Vertex dull yellow-brown, narrow;
compound eyes large, with dorso-mesal edges slanted anterad in dorsal aspect; vertex and eyes normal in female. Spurs
straw; lateral member of middle and hind-leg pairs notably shorter than mesal companions. Thorax uniformly dull
red-brown (dull yellow-brown in female). Legs red-brown, to straw distally; yellow-brown to straw in female.
Genitalia. Male. (Fig. 290-294). (Specimen from Baker, Illinois, USA). Males distinguished by distal article of
clasper, in lateral aspect (Fig. 290), relatively narrow, with tip curved dorsad from entire width of article; by gap between
distal lobes of tergum X, in dorsal aspect (Fig. 291), wide, composite, with inner portion u-shaped; and by distal cleft of
aedeagus, in dorsal aspect (Fig. 294), complex, with four separate modifications throughout its depth.
Genitalia. Female. (Fig. 295-296). (Specimen from Pontiac, Illinois, USA). Females distinguished by clasper
receptacle outer margin with distinct tooth close to posterior end of margin, in lateral aspect (Fig. 295); by receptacle
directed dorso-anterad, without inner opening; and by only dorsal lobes of segment XI visible in dorsal aspect (Fig. 296).
Biology. — Ross (1944) indicates flight season lasts from May to late August. Little more
known, but larvae may prefer very large, warm rivers (see Schuster & Etnier, 1978:86).
Distribution. — Known from Minnesota to Kentucky and Ohio; not yet recorded from
Canada.
Hydropsyche venularis Banks
Map 48; Fig. 297-303
Hydropsyche venularis Banks, 1914:252; Betten, 1934:524; Milne, 1936:73 (as synonym of H. scalaris)\ Ross, 1944:294;
Schuster & Etnier, 1978:96.
Description. — Male fore-wing length 9.98 mm; warm red-brown, faintly irrorate distally; veins Cul+2 and A
darker. In female, irrorate between R1 and RS also. Antennae yellow-brown; basal nine flagellar annuli each with oblique,
dark band; eight in female. Vertex yellow-brown, narrow, eyes larger than in other species; female normal. Spurs yellow;
lateral member of middle leg pairs shorter than mesal companions. Thorax yellow-brown, to red-brown laterally. Legs
straw-coloured.
Genitalia. Male. (Fig. 297-301). (Specimen from Conasauga R., Bradley Co., Tennessee, USA). Males distinguished
by distal article of clasper, in lateral aspect (Fig. 297), with disto-dorsal corner toothed; by distal lobes of tergum X
Arctopsychidae and Hydropsychidae (Trichoptera)
91
rounded-triangular in lateral aspect; and by these distal lobes, in dorsal aspect (Fig. 298), separated by flared, v-shaped
gap.
Genitalia. Female. (Fig. 302-303). (Specimen from Conasauga R., Bradley Co., Tennessee, USA). Females
distinguished by clasper receptacle represented by depression on lateral wall of segment X (Fig. 302); by this depression
bounded anteriorly by declivity; and by vulval scale with angled secondary sclerotised strap on side, besides narrow, short,
primary strap.
Biology. — Larvae occur in medium-sized rivers with large riffle areas; they seem to prefer
vegetation-covered rocks. No flight records available.
Distribution. — Recorded from area bounded by Wisconsin, Missouri, Georgia, and New
York (Map 48). Not yet known from Canada.
Quaest. Ent., 1987,23 (1)
92
Nimmo
Map 31. Known distribution of Hydropsyche aerata Ross in North America, by state.
Map 32. Known distribution of Hydropsyche alvata Denning in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
93
Map 33. Collection localities for Hydropsyche arinale Ross in Canada, with known distribution in North America by state
or province.
Map 34. Collection localities for Hydropsyche bidens Ross in Canada, with known distribution in North America by state
or province.
Quaest. Ent.. 1987,23 (1)
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Nimmo
Map 35. Collection localities for Hydropsyche californica Banks in Canada, with known distribution in North America by
state or province.
Map 36. Collection localities for Hydropsyche dicantha Ross in Canada, with known distribution in North America by
state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
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Map 37. Known distribution of Hydropsyche frisoni Ross in North America, by state.
Map 38. Collection localities for Hydropsyche hageni Banks in Canada, with known distribution in North America by
state or province.
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96
Nimmo
Map 39. Collection localities for Hydropsyche leonardi Ross in Canada, with known distribution in North America by
state or province.
Map 40. Collection localities for Hydropsyche occidentalis Banks in Canada, with known distribution in North America
by state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
97
Map 41 . Known distribution of Hydropsyche orris Ross in North America, by state.
Map 42. Collection localities for Hydropsyche phalerata Hagen in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987,23 (1)
98
Nimmo
Map 43. Collection localities for Hydropsyche placoda Ross in Canada, with known distribution in North America by
state or province.
Map 44. Known distribution of Hydropsyche rossi in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
99
Map 45. Collection localities for Hydropsyche scalaris Hagen in Canada, with known distribution in North America by
state or province.
Map 46. Collection localities for Hydropsyche simulans Ross in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987, 23 (1)
100
Nimmo
Map 47. Known distribution of Hydropsyche valanis Ross in North America, by state.
Map 48. Known distribution of Hydropsyche venularis Banks in North America, by state.
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101
Fig. 182-188, Hydropsyche aerata Ross: 182, genital capsule of male, lateral aspect; 183, genital capsule of male, dorsal
aspect; 184, left clasper of male, posterior aspect; 185, aedeagus of male, lateral aspect; 186, aedeagus of male, dorsal
aspect of tip; 187, genital segments of female, lateral aspect; 188, genital segments of female, dorsal aspect, pr, preanal
appendage; cr, clasper receptacle; ce, cercus, vs, vulval scale.
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102
Nimmo
Fig. 189-198. 189-193, Hydropsyche alvata Denning: 189, genital capsule of male, lateral aspect; 190, genital capsule of
male, dorsal aspect; 191, aedeagus of male, lateral aspect; 192, aedeagus of male, dorsal aspect of tip; 193, left clasper of
male, posterior aspect. 194-198, Hydropsyche leonardi Ross: 194, genital capsule of male, lateral aspect; 195, genital
capsule of male, dorsal aspect; 196, aedeagus of male, lateral aspect; 197, aedeagus of male, dorsal aspect of tip; 198, left
clasper of male, posterior aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
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Fig. 199-205, Hydropsyche arinale Ross: 199, genital capsule of male, lateral aspect; 200, genital capsule of male, dorsal
aspect; 201, left clasper of male, posterior aspect; 202, aedeagus of male, lateral aspect; 203, aedeagus of male, dorsal
aspect of tip; 204, genital segments of female, lateral aspect; 205, genital segments of female, dorsal aspect.
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Nimmo
Fig. 206-212, Hydropsyche bidens Ross: 206, genital capsule of male, lateral aspect; 207, genital capsule of male, dorsal
aspect; 208, left clasper of male, posterior aspect; 209, aedeagus of male, lateral aspect; 210, aedeagus of male, dorsal
aspect of tip; 211, genital segments of female, lateral aspect; 212, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
105
Fig. 213-219, Hydropsyche californica Banks: 213, genital capsule of male, lateral aspect; 214, genital capsule of male,
dorsal aspect; 215, left clasper of male, posterior aspect; 216, aedeagus of male, lateral aspect; 217, aedeagus of male,
dorsal aspect of tip; 218, genital segments of female, lateral aspect; 219, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
106
Nimmo
Fig. 220-226, Hydropsyche dicant ha Ross: 220, genital capsule of male, lateral aspect; 221, genital capsule of male, dorsal
aspect; 222, left clasper of male, posterior aspect; 223, aedeagus of male, lateral aspect; 224, aedeagus of male, dorsal
aspect of tip; 225, genital segments of female, lateral aspect; 226, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
107
Fig. 227-233, Hydropsyche frisoni Ross: 227, genital capsule of male, lateral aspect; 228, genital capsule of male, dorsal
aspect; 229, left clasper of male, posterior aspect; 230, aedeagus of male, lateral aspect; 231, aedeagus of male, dorsal
aspect of tip; 232, genital segments of female, lateral aspect; 233, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
108
Nimmo
Fig. 234-240, Hydropsyche hageni Banks: 234, genital capsule of male, lateral aspect; 235, genital capsule of male, dorsal
aspect; 236, left clasper of male, posterior aspect; 237, aedeagus of male, lateral aspect; 238, aedeagus of male, dorsal
aspect of tip; 239, genital segments of female, lateral aspect; 240, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
109
241
Q.5MM
243
244
Fig. 241-247, Hydropsyche occidentalis Banks: 241, genital capsule of male, lateral aspect; 242, genital capsule of male,
dorsal aspect; 243, left clasper of male, posterior aspect; 244, aedeagus of male, lateral aspect; 245, aedeagus of male,
dorsal aspect of tip; 246, genital segments of female, lateral aspect; 247, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
110
Nimmo
Fig. 248-254, Hydropsyche orris Ross: 248, genital capsule of male, lateral aspect; 249, genital capsule of male, dorsal
aspect; 250, left clasper of male, posterior aspect; 251, aedeagus of male, lateral aspect; 252, aedeagus of male, dorsal
aspect of tip; 253, genital segments of female, lateral aspect; 254, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
111
Fig. 255-261, Hydropsyche phalerata Hagen: 255, genital capsule of male, lateral aspect; 256, genital capsule of male,
dorsal aspect; 257, left clasper of male, posterior aspect; 258, aedeagus of male, lateral aspect; 259, aedeagus of male,
dorsal aspect of tip; 260, genital segments of female, lateral aspect; 261, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
112
Nimmo
Fig. 262-268, Hydropsyche placoda Ross: 262, genital capsule of male, lateral aspect; 263, genital capsule of male, dorsal
aspect; 264, aedeagus of male, lateral aspect; 265, aedeagus of male, dorsal aspect of tip; 266, left clasper of male,
posterior aspect; 267, genital segments of female, lateral aspect; 268, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
113
Fig. 269-275, Hydropsyche rossi Flint, Voshell, & Parker: 269, genital capsule of male, lateral aspect; 270, genital
capsule of male, dorsal aspect; 271, left clasper of male, posterior aspect; 272, aedeagus of male, lateral aspect; 273,
aedeagus of male, dorsal aspect of tip; 274, genital segments of female, lateral aspect; 275, genital segments of female,
dorsal aspect.
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114
Nimmo
Fig. 276-282, Hydropsyche scalaris Hagen: 276, genital capsule of male, lateral aspect; 277, genital capsule of male,
dorsal aspect; 278, left clasper of male, posterior aspect; 279, aedeagus of male, lateral aspect; 280, aedeagus of male,
dorsal aspect of tip; 281, genital segments of female, lateral aspect; 282, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
115
Fig. 283-289, Hydropsyche simulans Ross: 283, genital capsule of male, lateral aspect; 284, genital capsule of male,
dorsal aspect; 285, left clasper of male, posterior aspect; 286, aedeagus of male, lateral aspect; 287, aedeagus of male,
dorsal aspect of tip; 288, genital segments of female, lateral aspect; 289, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
116
Nimmo
Fig. 290-296, Hydropsyche valanis Ross: 290, genital capsule of male, lateral aspect; 291, genital capsule of male, dorsal
aspect; 292, left clasper of male, posterior aspect; 293, aedeagus of male, lateral aspect; 294, aedeagus of male, dorsal
aspect of tip; 295, genital segments of female, lateral aspect; 296, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
117
Fig. 297-303, Hydropsyche venularis Banks: 297, genital capsule of male, lateral aspect; 298, genital capsule of male,
dorsal aspect; 299, left clasper of male, posterior aspect; 300, aedeagus of male, lateral aspect; 301, aedeagus of male,
dorsal aspect of tip; 302, genital segments of female, lateral aspect; 303, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
118
Nimmo
SPECIES GROUP 2
This small group of species characterised by aedeagus sclerotised throughout, with distal
end roughly, transversely truncate, of more or less uniform width throughout; and by clasper
receptacle of females simple, blind invagination of lateral wall of segment X, without inner
opening.
Hydropsyche betteni Ross
Map 49; Fig. 304-310
Hydropsyche betteni Ross, 1 938b: 1 46; Denning, 1943:122; Ross, 1944:99; Schuster & Etnier, 1978:61.
Hydropsyche incommoda not Hagen; Betten, 1934:188.
Description. — Male fore-wing length 10.53 mm; warm red-brown, with darker areas concentrated about Cul to
A3. Hind-wing faintly tinted brown. Antennae red-brown; basal eight flagellar annuli each with oblique, dark band.
Vertex yellow-brown. Spurs pale brown; lateral member of middle and hind-leg pairs shorter than mesal companions.
Thorax deep, rich red-brown, to paler laterally. Legs brownish yellow; red-brown in female.
Genitalia. Male. (Fig. 304-308). (Specimen from St John’s, Newfoundland). Males distinguished by aedeagus, in
lateral aspect (Fig. 307), with base curved ventrad of remainder, in semi-circle; and by gap between distal lobes of tergum
X, in dorsal aspect (Fig. 305), vase-shaped, each side of wider outer limit with distal tooth.
Genitalia. Female. (Fig. 309-310). (Specimen from St John’s, Newfoundland). Females distinguished by clasper
receptacle widened internally (Fig. 309), without inner opening (Fig. 310), with two lobes on floor of outer opening of
receptacle; and by small, curved, slender secondary sclerotised strap laterally on vulval scale.
Biology. — Ross (1944) indicates that larvae prefer small to medium, riffled streams. They
have been recorded in water film of dam spillways. Schuster & Etnier (1978) add that warmer
waters are preferred; also, that this species is one of the more pollution-tolerant in
Hydropsyche. It is, also, often the only Hydropsyche species in given streams. Canadian flight
season extends from May 18 to October 2.
Distribution. — Recorded from Saskatchewan to Arkansas, Georgia, and Newfoundland,
the species appears to be general throughout eastern North America, with extensions into the
Boreal Forest (Map 49). In Canada it is recorded from Saskatchewan, then from northwestern
Ontario to Newfoundland.
Hydropsyche confusa (Walker)
Map 50; Fig. 311-317
Philopotamus confusus Walker, 1852:103.
Hydropsyche confusa ; Milne, 1936:61; Betten & Mosely, 1940:21; Nimmo, 1981:259.
Hydropsyche seperata Banks, 1936:129; Denning, 1943:121; Ross & Spencer, 1952:46 (as synonym of H. guttata Pictet);
Smith, 1979:10; Nimmo, 1981:259 (as synonym of H. confusa (Walker)).
Hydropsyche guttata Pictet; Schuster & Etnier, 1978:126.
Hydropsyche corbetti Nimmo, 1966a:688; Schuster & Etnier, 1978:126 (as synonym of H. guttata Pictet); Nimmo,
1981:259 (as synonym of H. confusa (Walker)).
Description. — Male fore-wing length 8.66 mm; bright grey-brown, faintly irrorate; darker in female. Antennae
brownish cream; basal nine flagellar annuli each with oblique, dark band. Vertex dark grey-brown. Spur formula 1,4,4 in
male; 2,4,4 in female; pale brown; lateral member of middle leg pairs notably shorter than mesal companions. Thorax dark
brown, to orange-brown laterally. Legs dull pale brown.
Genitalia. Male. (Fig. 311-315). (Specimen from Empress, Alberta). Males distinguished by distal article of clasper,
in lateral aspect (Fig. 311), long, slightly bulbous distally, curved dorsad; by distal article, in posterior aspect (Fig. 313),
hooked mesad; and by almost total lack of gap between distal lobes of tergum X, in dorsal aspect (Fig. 312).
Genitalia. Female. (Fig. 316-317). (Specimen from Empress, Alberta). Females distinguished by large, rounded
clasper receptacle, in lateral aspect (Fig. 316), with two lobes on floor of receptacle entrance; and by presence of small,
angled, secondary sclerotised strap on side of vulval scale.
Arctopsychidae and Hydropsychidae (Trichoptera)
119
Biology. — Smith (1979) presents a comprehensive account from Saskatchewan: univoltine,
with extended emergence and flight season; pupae obtained from May 26 to August 24, with
peak in June-July; larvae primarily detritovores and herbivores; appear to prefer larger, more
turbid rivers.
Distribution. — Presently known from British Columbia and Washington to Quebec in east,
and Hudson’s Bay and arctic coasts in north, this species is known in United States only from
northern tier of States (Map 50). Canadian distribution records scattered; this species is
recorded from nearly the southern-most point to Canadian Arctic Coast, though not north of
tree line. Not known from eastern Quebec or Atlantic Provinces.
Hydropsyche cuanis Ross
Map 51; Fig. 318-324
Hydropsyche cuanis Ross, 1938b: 147; Ross, 1944:100; Schuster & Etnier, 1978:70.
Description. — Male fore-wing length 9.36 mm; light orange-brown, with no evident markings. Hind-wing faintly
tinted. Antennae yellow; basal eight flagellar annuli each with oblique, dark band (five in female). Vertex deep reddish
brown, narrowed anterad, with compound eyes of male much larger, relatively, than in most other species; female normal.
Spurs yellow; lateral member of middle and hind-leg pairs notably shorter than mesal companions; applicable in the
female to middle leg pairs only. Thorax rich red-brown, to more orange-brown laterally. Legs pale orange-brown to straw.
Genitalia. Male. (Fig. 318-322). (Specimen from Momence, Illinois, LISA). Males distinguished by basal article of
clasper narrow at base, widened distally, in lateral aspect (Fig. 318); by uniform width of basal article, in posterior aspect
(Fig. 320); by distal article narrowed in two stages, to acuminate tip (Fig. 318); and by gap between tergum X distal lobes
v-shaped.
Genitalia. Female. (Fig. 323-324). (Specimen from Kankakee R., Willmington, Illinois, USA - Paratype). Females
distinguished by sclerotised strap of vulval scale narrow, sinuate (Fig. 323); by clasper receptacle relatively narrow,
rounded, directed dorso-anterad, without grooves or lobes on receptacle floor (Fig. 323); and by receptacle without inner
opening (Fig. 324).
Biology.— Ross (1944) indicates that larvae prefer swift rapids areas of larger rivers. Flight
season apparently commences with May peak, which declines into August.
Distribution.— Limited to mid-west States of USA (Map 51). Not yet known from Canada.
Hydropsyche depravata Hagen
Map 52; Fig. 325-331
Hydropsyche depravata Hagen, 1861:290; Betten, 1934:187; Milne, 1036:70, 72, 73; Ross, 1944:100; Schuster & Etnier,
1978:63.
Description. — Male fore-wing length 9.75 mm; grey-brown, uniformly irrorate. Hind-wing very faintly tinted
grey-brown. A tennae red-brown; basal nine flagellar annuli each with oblique, dark band - eight in female. Vertex dark
brown. Spurs brown - yellow in female; lateral member of all pairs shorter than mesal companions. Thorax dark brown, to
red-brown laterally. Legs straw-coloured.
Genitalia. Male. (Fig. 325-329). (Specimen from Beaver Ck, Knox Co., Tennessee, USA). Males distinguished by
distal article of clasper, in lateral aspect (Fig. 325), evenly tapered from base to rounded tip (tip curved abruptly); by pair
of black sclerites housed in tip of aedeagus, as seen in dorsal aspect (Fig. 329); and by gap between tergum X distal lobes
wide, v-shaped, not deep (Fig. 326).
Genitalia. Female. (Fig. 330-331). (Specimen from Beaver Ck, Knox Co., Tennessee, USA). Females distinguished
by clasper receptacle large, rounded, with pair of grooves on floor of receptacle opening, without inner opening (Fig. 330,
331); and by only dorsal lobes of segment XI visible in dorsal aspect.
Biology. — Schuster & Etnier (1978) state that larvae live in warm-water, small streams
with high organic loading. Apparently found mostly on medium-sized rocks in riffles. Only
flight record available is for July 26, in Saskatchewan.
Distribution. — Primarily known from Indiana to Georgia to Virginia, with isolated record
from prairie of southern Saskatchewan, in Canada (Map 52).
Quaest. Ent., 1987,23 (1)
120
Nimmo
Map 49. Collection localities for Hydropsyche betteni Ross in Canada, with known distribution in North America by state
or province.
Map 50. Collection localities for Hydropsyche confusa (Walker) in Canada, with known distribution in North America by
state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
121
Map 51. Known distribution of Hydropsyche cuanis Ross in North America, by state.
Map 52. Collection localities for Hydropsyche depravata Hagen in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987,23 (1)
122
Nimmo
306
Fig. 304-310, Hydropsyche betteni Ross: 304, genital capsule of male, lateral aspect; 305, genital capsule of male, dorsal
aspect; 306, left clasper of male, posterior aspect; 307, aedeagus of male, lateral aspect; 308, aedeagus of male, dorsal
aspect of tip; 309, genital segments of female, lateral aspect; 310, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
123
Fig. 311-317, Hydropsyche confusa (Walker): 311, genital capsule of male, lateral aspect; 312, genital capsule of male,
dorsal aspect; 313, left clasper of male, posterior aspect; 314, aedeagus of male, lateral aspect; 315, aedeagus of male,
dorsal aspect of tip; 316, genital segments of female, lateral aspect; 317, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
124
Nimmo
Fig. 318-324, Hydropsyche cuanis Ross: 318, genital capsule of male, lateral aspect; 319, genital capsule of male, dorsal
aspect; 320, left clasper of male, posterior aspect; 321, aedeagus of male, lateral aspect; 322, aedeagus of male, dorsal
aspect of tip; 323, genital segments of female, lateral aspect; 324, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
125
Fig. 325-331, Hydropsyche depravata Hagen: 325, genital capsule of male, lateral aspect; 326, genital capsule of male,
dorsal aspect; 327, left clasper of male, posterior aspect; 328, aedeagus of male, lateral aspect; 329, aedeagus of male,
dorsal aspect of tip; 330, genital segments of female, lateral aspect; 331, genital segments of female, dorsal aspect.
Quaest. Ent., 1987,23 (1)
126
Nimmo
SPECIES GROUP 3
This group characterised by aedeagus with toothed and/or spinate membranous lobes
distally.
SUBGROUP A
This subgroup characterised by possession of membranous lobes on aedeagus both dorsally
and ventro-laterally.
Hydropsyche amblis Ross
Map 53; Fig. 332-338
Hydropsyche amblis Ross, 1938a: 120; Ross, 1944:294; Anderson, 1976:65; Schefter & Wiggins, 1986:51.
Description. — Male fore-wing length 9.13 mm; pale cream-brown, with hint of irroration around distal edge.
Hind-wing hyaline. Antennae red-brown; no oblique dark bands. Vertex dark brown, warts cream. Spurs straw-coloured;
lateral member of middle leg pairs, and hind-leg apical pair, notably shorter than mesal companions. Thorax deep
red-brown, to paler laterally. Legs straw coloured.
Genitalia. Male. (Fig. 332-336). (Specimen from East Fork, Humbug Ck, Clatsop Co., Oregon, USA). Males
distinguished by massive, postero-ventrally directed distal lobes of tergum X, in lateral aspect (Fig. 332); by tooth of dorsal
lobe of aedeagus directed postero-dorsad (Fig. 335); by ventro-lateral lobes of aedeagus directed antero-ventrad, with
bundle of spines at tip; and by tip of distal article of clasper, in lateral aspect, with small curved indentation.
Genitalia. Female. (Fig. 337-338). (Specimen from Okop Ck, Eatonville, Washington, USA). Females distinguished
by small, anteriorly directed clasper receptacle traversed by thin, dark line which passes down lateral wall of segment X;
by small, irregular sclerotised strap of vulval scale (Fig. 337); and by clasper receptacle, in dorsal aspect (Fig. 338),
directed antero-mesad, without visible inner openings.
Biology. — Anderson (1976) suggests that larvae prefer small streams. Emergence has been
recorded from early May to mid-August in Oregon.
Distribution. — Presently known only from Lower Mainland of British Columbia, Canada
and from Washington and Oregon, USA (Map 53).
Hydropsyche piatrix Ross
Map 54; Fig. 339-345
Hydropsyche piatrix Ross, 1938b: 1 48; Ross, 1944:97; Schefter & Wiggins, 1986:68.
Symphitopsyche piatrix', Schuster & Etnier, 1978:57.
Description. — Male fore-wing length 7.84 mm; uniform yellowish brown, very faintly irrorate. Hind-wing paler.
Antennae pale yellowish brown, no oblique, dark bands. Vertex brown, warts paler. Spurs brownish straw; lateral member
of middle leg pairs notably shorter than mesal companions. Thorax dark brown, to grey-brown laterally. Legs pale
brownish straw, to straw.
Genitalia. Male. (Fig. 339-343). (Specimen from Mammoth Springs, Arkansas, USA - Paratype). Males
distinguished by tergum X distal lobes small, rounded, curved slightly ventrad in lateral aspect (Fig. 339); by these lobes,
in dorsal aspect, finger-like, curved postero-mesad, gap between elliptical; by ventro-lateral lobe of aedeagus tapered
anterad, without teeth or spines (Fig. 342); and by dorsal lobe not produced, with minute tooth directed slightly
antero-laterad.
Genitalia. Female. (Fig. 344-345). (Specimen from Mammoth Springs, Arkansas, USA). Females distinguished by
clasper receptacle directed dorso-posterad in lateral aspect (Fig. 344), meso-posterad in dorsal aspect (Fig. 345); by vulval
scale with two sclerotised straps in lateral aspect - primary club-shaped, secondary triangular, dorsal; and by segment XI
dorsal lobe, in dorsal aspect (Fig. 345) large enough to obscure all beneath, with mesal shoulder.
Biology. — Very little known except that this species has been taken only at spring-like
waters (Schuster & Etnier, 1978). Only flight dates available are from June and early July.
Distribution. — Very scattered - North Dakota, Missouri, Arkansas, and St Lawrence R.
valley of Quebec (Map 54).
Arctopsychidae and Hydropsychidae (Trichoptera)
127
Hydropsyche vex a Ross
Map 55; Fig. 346-352
Hydropsyche vexa Ross, 1938b: 148; Denning, 1943:124; Ross, 1944:97; Schefter & Wiggins, 1986:81.
Symphitopsyche vexa; Schuster & Etnier, 1978:127.
Description. — Male fore-wing length 7.41 mm; golden brown, faintly irrorate. Antennae pale brown; basal seven
or eight flagellar annuli each with oblique, dark band; five in female, paler. Vertex orange-brown. Spurs yellow; lateral
member of middle leg pairs, and hind-leg apical pair, notably shorter than mesal companions; applies only to middle leg
pairs in female. Thorax orange-brown, to yellow-brown laterally. Legs pale brownish yellow.
Genitalia. Male. (Fig. 346-350). (Specimen from White Earth R., Hwy 28, Alberta). Males distinguished by basal
article of clasper, in lateral aspect (Fig. 346), with slender base, expanded distally; by tergum X distal lobes long, thin,
curved slightly ventrad in lateral aspect (Fig. 346), curved mesad in dorsal aspect (Fig. 347), space between almost
enclosed; by dorsal membranous lobe of aedeagus small, with small, acuminate tooth directed posterad (Fig. 349, 350);
and by ventro-lateral lobe of aedeagus long, straight, directed antero-ventrad, with distal pocket of spines.
Genitalia. Female. (Fig. 351-352). (Specimen from White Earth R., Flwy 28, Alberta). Females distinguished by
clasper receptacle, in lateral aspect (Fig. 351), directed dorsad, of medium size, with two grooves on floor at entrance; and
by clasper receptacle, in dorsal aspect (Fig. 352), curved postero-mesad.
Biology. — Very little known. Flight records extend from May 22 to August 8 in Canada.
Distribution. — Recorded in narrow zone across Continent, from Idaho and Alberta to New
Brunswick and Maine (Map 55). In Canada known only from Prairie Provinces, Montreal, and
New Brunswick.
Hydropsyche walkeri Betten & Mosely
Map 56; Fig. 353-359
Hydropsyche maculicornis Walker, 1852:1 13 (preoccupied by Pictet, 1834 - now in Tinodes).
Hydropsyche walkeri Betten & Mosely, 1940:23 (new name); Ross, 1944:96; Schefter & Wiggins, 1986:83.
Symphitopsyche walkeri ; Schuster & Etnier, 1978:35.
Description. — Male fore-wing length 7.53 mm; pale red-brown. Hind-wing very palely tinted yellow-brown.
Antennae uniformly brown. Vertex dark reddish brown, warts paler. Spurs yellow-brown; lateral member of fore-leg apical
pair much finer than mesal companion, hyaline. Thorax dark reddish brown, to grey-brown laterally. Legs yellowish
brown. Warts, generally, paler.
Genitalia. Male. (Fig. 353-357). (Specimen from St Hippolyte, Quebec). Males distinguished by distal article of
clasper with distal half bent dorsad at approximately 45° to basal half, with tip slightly scalloped (Fig. 353); by tergum X
distal lobes with dorsal edge angled in lateral aspect (Fig. 353), lobes curved mesad, in dorsal aspect (Fig. 354), space
between almost enclosed; by aedeagus with ventro-lateral lobe directed basad along side (Fig. 356); and by dorsal lobes of
aedeagus slightly developed, tooth directed postero-dorsad.
Genitalia. Female. (Fig. 358-359). (Specimen from St Hippolyte, Quebec). Females distinguished by large,
crescent-shaped clasper receptacle located in postero-dorsal angle of segment X (Fig. 358); and by sclerotised strap of
vulval scale large, long, wide except tapered basad.
Biology. — According to Schuster & Etnier (1978) larvae appear to prefer small to medium
sized streams with coarse gravel to small rock bottom, which are rich in organic materials.
Riffle areas appear to be preferred, with smoothly-flowing water. Canadian flight season
extends from May 5 to September 2.
Distribution.— From Saskatchewan to Virginia, Maine, and Quebec (Map 56). In Canada
recorded from northcentral Saskatchewan to east coast of Hudson’s Bay, south to southern
Quebec and Ontario.
Quaest. Ent., 1987, 23 (1)
128
Nimmo
Map 53. Collection localities for Hydropsyche amblis Ross in Canada, with known distribution in North America by state
or province.
Map 54. Collection localities for Hydropsyche piatrix Ross in Canada, with known distribution in North America by state
or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
129
Map 55. Collection localities for Hydropsyche vexa Ross in Canada, with known distribution in North America by state or
province.
Map 56. Collection localities for Hydropsyche walkeri Betten & Mosely in Canada, with known distribution in North
America by state or province.
Quaest. Ent., 1987, 23 (1)
130
Nimmo
Fig. 332-338, Hydropsyche amblis Ross: 332, genital capsule of male, lateral aspect; 333, genital capsule of male, dorsal
aspect; 334, left clasper of male, posterior aspect; 335, aedeagus of male, lateral aspect; 336, aedeagus of male, dorsal
aspect of tip; 337, genital segments of female, lateral aspect; 338, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
131
Fig. 339-345, Hydropsyche piatrix Ross: 339, genital capsule of male, lateral aspect; 340, genital capsule of male, dorsal
aspect; 341, left clasper of male, posterior aspect; 342, aedeagus of male, lateral aspect; 343, aedeagus of male, dorsal
aspect of tip; 344, genital segments of female, lateral aspect; 345, genital segments of female, dorsal aspect.
Quaest. Ent., 1987,23 (1)
132
Nimmo
Fig. 346-352, Hydropsyche vexa Ross: 346, genital capsule of male, lateral aspect; 347, genital capsule of male, dorsal
aspect; 348, left clasper of male, posterior aspect; 349, aedeagus of male, lateral aspect; 350, aedeagus of male, dorsal
aspect of tip; 351, genital segments of female, lateral aspect; 352, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
133
Fig. 353-359, Hydropsyche walkeri Betten & Mosely: 353, genital capsule of male, lateral aspect; 354, genital capsule of
male, dorsal aspect; 355, left clasper of male, posterior aspect; 356, aedeagus of male, lateral aspect; 357, aedeagus of
male, dorsal aspect of tip; 358, genital segments of female, lateral aspect; 359, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
134
Nimmo
SUBGROUP B
This subgroup characterised by aedeagus without ventro-lateral membranous lobes; distal
tooth directed ventrad.
Hydropsyche alhedra Ross
Map 57; Fig. 360-366
Hydropsyche alhedra Ross, 1939:67; Ross, 1944:294; Schefter, Wiggins, & Unzicker, 1986:69; Schefter & Wiggins,
1986:45.
Hydropsyche riola Denning; Schefter, Wiggins, & Unzicker, 1986:69 (as synonym of H. alhedra ).
Hydropsyche racona Denning; Schefter, Wiggins, & Unzicker, 1986:69 (as synonym of H. alhedra).
Symphitopsyche alhedra ; Schuster & Etnier, 1978:45.
See note at end of ‘Introduction’.
Description. — Male fore-wing length 9.09 mm; pale grey-brown, faintly irrorate posterad of M; female paler.
Hind-wing hyaline. Antennae brown; basal seven flagellar annuli each with oblique, dark band; female paler. Vertex dark
red-brown, warts paler; female paler. Spurs straw-coloured; mesal member of middle leg apical pair shorter than lateral
companion. Thorax dark red-brown, to slightly paler laterally. Legs red-brown; female paler.
Genitalia. Male. (Fig. 360-364). (Specimen from St Hippolyte, Quebec). Males distinguished by tergum X distal
lobes, in dorsal aspect (Fig. 361), very slightly directed mesad; by distal tooth of aedeagus dorsal lobe, in lateral aspect
(Fig. 363), with rounded base surmounted by much narrower rounded spine - not visible in dorsal aspect (Fig. 364); and
by lateral lobe on posterior edge of segment IX set very low, connected to dorsum of segment by thin, dark line.
Genitalia. Female. (Fig. 365-366). (Specimen from St Hippolyte, Quebec). Females distinguished by clasper
receptacle directed dorsad in lateral aspect (Fig. 365), directed mesad in dorsal aspect (Fig. 366); by dorsal and ventral
lobes, and cerci of segment XI very close; and by sclerotised strap of vulval scale long, not wide distally, sinuate.
Biology. — Larvae apparently inhabit rapids sections of smaller, cool, clear streams
(Schuster & Etnier, 1978). Emergence in April. Few Canadian flight records range from June
5 to August 29.
Distribution. — Very scattered (Map 57), with isolated records from Tennessee, North
Carolina, to Pennsylvania, Quebec, and southern Manitoba.
Hydropsyche bifida Banks
Map 58; Fig. 367-373
Hydropsyche bifida Banks, 1905a:15; Betten, 1934:193; Milne, 1936:73; Denning, 1943:129; Ross, 1944:97; Schefter &
Unzicker, 1984:331 (as synonym of H. morosai).
Symphitopsyche bifida\ Schuster & Etnier, 1978:30.
See note at end of ‘Introduction’.
Description. — Male fore-wing length 8.03 mm; pale golden brown, faintly irrorate. Antennae brownish yellow;
basal nine flagellar annuli each with oblique, dark band. Vertex yellow-brown. Spurs straw-coloured; lateral member of
middle and hind-leg pairs notably shorter than mesal companions; not so in female. Thorax yellow-brown. Legs yellow, to
pale straw.
Genitalia. Male. (Fig. 367-371). (Specimen from Blindman R., Hwy 2, Ponoka, Alberta). Males distinguished by
tergum X distal lobes set high on distal end of tergum, thin, almost pointed distally, curved slightly ventrad, in lateral
aspect (Fig. 367); by basal article of clasper slender, especially at base, crooked in lateral aspect; by distal article of clasper
with base as wide as basal article, then sharply tapered to distal half which is of even width, narrow, rounded distally; and
by tooth of dorsal lobe of aedeagus curved slightly ventrad, small, clothed in minute dentitions.
Genitalia. Female. (Fig. 372-373). (Specimen from Blindman R., Hwy 2, Ponoka, Alberta). Females distinguished by
clasper receptacle curved antero-dorsad in lateral aspect (Fig. 372), meso-posterad in dorsal aspect (Fig. 373); by mouth of
receptacle with one groove on floor; and by cerci closer to dorsal lobe of segment XI than to ventral lobe.
Biology. — Larvae commonly collected in medium-sized creeks or small rivers with coarse
gravel or small rock substrates, and high organic loading. Canadian flight season extends from
May 22 to October 16, with bulk of records in June- July.
Arctopsychidae and Hydropsychidae (Trichoptera)
135
Distribution. — Widely distributed, from Great Slave Lk. and central British Columbia to
Oklahoma, Tennessee, Vermont, and Quebec (Map 58). In Canada, well recorded across
Prairie Provinces, sparsely in British Columbia, north to near Great Slave Lk. In the east, it is
known from western and southern Quebec, and southern Ontario.
Map 57. Collection localities for Hydropsyche aihedra Ross in Canada, with known distribution in North America by
state or province.
Map 58. Collection localities for Hydropsyche bifida Banks in Canada, with known distribution in North America by state
or province.
Quaest. Ent., 1987,23 (1)
136
Nimmo
Fig. 360-366, Hydropsyche alhedra Ross: 360, genital capsule of male, lateral aspect; 361, genital capsule of male, dorsal
aspect; 362, left clasper of male, posterior aspect; 363, aedeagus of male, lateral aspect; 364, aedeagus of male, dorsal
aspect of tip; 365, genital segments of female, lateral aspect; 366, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
137
Fig. 367-373, Hydropsyche bifida Banks: 367, genital capsule of male, lateral aspect; 368, genital capsule of male, dorsal
aspect; 369, left clasper of male, posterior aspect; 370, aedeagus of male, lateral aspect; 371, aedeagus of male, dorsal
aspect of tip; 372, genital segments of female, lateral aspect; 373, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
138
Nimmo
SUBGROUP C
This group characterised by aedeagus not only with dorsal membranous lobes, but also with
ventro-lateral lobes short, restricted to extreme distal portion of aedeagus; these lobes reduced
in some species to simple, unsclerotised apertures laterally on aedeagus extremity.
Hydropsyche bronta Ross
Map 59; Fig. 374-380
Hydropsyche bronta Ross, 1 938b: 1 49; Denning, 1943:125; Ross, 1944:98; Schefter & Wiggins, 1986:52.
Symphitopsyche ( Ceratopsyche ) bronta; Ross & Unzicker, 1977:305.
Symphitopsyche bronta ; Schuster & Etnier, 1978:37.
Description. — Male fore-wing length 7.29 mm; pale grey-brown, faintly irrorate; female pale orange-brown.
Antennae brownish cream; apparently without oblique, dark bands in male; basal seven flagellar annuli of female each
with faint, oblique, darker band. Spurs yellow-brown; lateral member of middle leg pairs notably shorter than mesal
companions. Thorax and legs bright, pale yellow-brown overall.
Genitalia. Male. (Fig. 374-378). (Specimen from Carrot Ck, Hwy 16, Alberta). Males distinguished by distal article
of clasper, in lateral aspect (Fig. 374), triangular, apex slightly drawn out; by tergum X distal lobes short in lateral aspect
(Fig. 374), curved ventrad; by these lobes, in dorsal aspect (Fig. 375), short, linear, not convergent; and by dorsal lobe of
aedeagus with large, stout, linear, distally acuminate distal tooth or spine directed antero-ventrad (Fig. 377).
Genitalia. Female. (Fig. 379-380). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Females
distinguished by clasper receptacle small, directed antero-dorsad in lateral aspect (Fig. 379), with outer margin of
receptacle continued dorsad and ventrad by thin, dark lines; by inner opening of receptacle clearly evident in lateral aspect;
and by receptacle, in dorsal aspect (Fig. 380), directed mesad.
Biology. — I have taken adults from a great variety of small creeks to medium-sized rivers,
some cool, others warm, some clear, turbulent, others slow, weedy. Flight season ranges from
May 25 to September 2, in Canada, with concentration of records in June and early July.
Distribution. — Widespread from Alberta and Wyoming, to South Carolina and Nova
Scotia (Map 59). In Canada this species commonly recorded from Alberta lower foothills to
eastern Quebec, New Brunswick, and Nova Scotia, northward into southern reaches of Boreal
Forest.
Hydropsyche cheilonis Ross
Map 60; Fig. 381-387
Hydropsyche cheilonis Ross, 1 938b: 1 49; Ross, 1944:98; Schefter & Wiggins, 1986:57.
Symphitopsyche cheilonis; Schuster & Etnier, 1978:33.
Description. — Male fore-wing length 7.80 mm; light purple-brown, slight irroration along A. Hind-wing tinted
grey-brown. Antennae yellow; no banding on annuli. Vertex dark brown anteriorly, yellow posteriorly, warts dark brown.
Spurs grey-brown; lateral member of fore- and hind-leg pairs shorter than mesal companions. Thorax mottled grey-brown
and yellow-brown. Legs grey-brown to dull straw.
Genitalia. Male. (Fig. 381-385). (Specimen from Crossville, Cumberland Co., Tennessee, USA). Males distinguished
by basal article of clasper, in lateral aspect (Fig. 381), sinuate, distal quarter wider than remainder; by distal article of
clasper with trinagular base, with apex drawn out to finger-like process; by tergum X distal lobes, in dorsal aspect (Fig.
382), angled slightly mesad, linear; and by dorsal lobe of aedeagus with long, thin tooth or spine distally - tooth angled
slightly ventrad at mid-point (Fig. 384), slightly expanded distally in dorsal aspect (Fig. 385).
Genitalia. Female. (Fig. 386-387). (Specimen from Beaver Ck, Knox Co., Tennessee, USA). Females distinguished
by vulval scale with thin, curved primary sclerotised strap, and irregular, formless secondary sclerite laterally (Fig. 386);
by clasper receptacle, in lateral aspect (Fig. 386), curved dorso-posterad, with two grooves on floor of outer opening; and
by receptacle, in dorsal aspect (Fig. 387), curved meso-posterad.
Biology. — Larvae seem to prefer small to medium-sized, warm-water streams of slow
current and large riffle areas. Emergence from early April to September.
Arctopsychidae and Hydropsychidae (Trichoptera)
139
Distribution. — Not yet known from Canada. Presently known from Wisconsin to Tennessee
and Virginia (Map 60).
Hydropsyche morosa Hagen
Map 61; Fig. 388-394
Hydropsyche morosa Hagen, 1861:287; Milne, 1936:71, 73; Denning, 1943:127; Ross, 1944:98; Wiggins, 1977:106;
Schefter & Unzicker, 1984:331; Schefter & Wiggins, 1986:62.
Hydropsyche chlorotica Hagen, 1861:290; Ross, 1 938c: 1 6 (as synonym of H. morosa).
Symphitopsyche morosa ; Schuster & Etnier, 1978:41.
See note at end of ‘Introduction’.
Description. — Male fore-wing length 9.63 mm; pale yellow-brown, faintly irrorate distally and along distal third
of Rl. Hind-wing hyaline. Antennae brown; with dark bands at each end of flagellar annuli. Vertex brown. Spurs brown.
Thorax red-brown, to paler laterally. Legs light yellow-brown.
Genitalia. Male. (Fig. 388-392). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Males
distinguished by distal article of clasper, in lateral aspect (Fig. 388), with wide base which supports long, finger-like distal
process; by tergum X distal lobes, in dorsal aspect (Fig. 389), well separated, gently curved, thin, distally rounded; and by
dorsal lobe of aedeagus with massive distal tooth, dorsal edge of which is spinate.
Genitalia. Female. (Fig. 393-394). (Specimen from Huberdeau, Quebec). Females distinguished by clasper
receptacle, in lateral aspect (Fig. 393), linear, directed dorso-anterad; by outer opening of receptacle with two lobes on
floor of opening; and by receptacle, in dorsal aspect (Fig. 394), directed mesad.
Biology. — Apparently a species of medium-sized rivers with large riffle areas of small to
medium rocks clothed in weed. Canadian flight season extends from May 16 to September 21.
Distribution.— Known from Cape Breton Island, Nova Scotia, west to Alberta, south to
Tennessee (Map 61). In Canada the species is known from southern Ontario and Quebec, with
scattered records to north and west.
Hydropsyche slossonae Banks
Map 62; Fig. 395-401
Hydropsyche slossonae Banks 1905a:14; Betten, 1934:185 (as synonym of H. alternans)', Milne, 1936:69, 72, 73; Denning,
1943:131; Ross, 1944:99; Schefter & Wiggins, 1986:70.
Symphitopsyche slossonae, Schuster & Etnier, 1978:47.
Description. — Male fore-wing length 9.36 mm; grey-brown, more or less irrorate. Antennae yellow-brown; basal
seven flagellar annuli each with oblique, dark band; five in female. Vertex brown anteriorly, to pale yellow-brown
posteriorly; dark brown with white warts in female. Spurs yellow; lateral member of middle leg pairs notably shorter than
mesal companions. Thorax brown, to brownish yellow laterally; dark brown to grey-brown laterally in female. Legs pale
brown, except hind-legs pale straw; uniform yellow-brown in female.
Genitalia. Male. (Fig. 395-399). (Specimen from Rapids Ck, Trans-Canada Hwy, Gap, Alberta). Males distinguished
by massive distal lobes of tergum X, in lateral aspect (Fig. 395), tapered rather abruptly in distal half; by these lobes, in
dorsal aspect (Fig. 396), lyre-like; and by dorsal lobe of aedeagus small, abruptly tapered anterad, with minute distal tooth
(Fig. 398).
Genitalia. Female. (Fig. 400-401). (Specimen from creek, Hwy 932, 6 miles S of Whitecourt, Alberta). Females
distinguished by clasper receptacle, in lateral aspect (Fig. 400), oriented vertically, curved dorso-posterad distally, with
two grooves on floor of outer opening; and by dorsal and ventral lobes of segment XI, and cerci, all visible in dorsal aspect
(Fig. 401).
Biology. — A species of cold-water streams; little more known of larval habitat preferences.
Canadian flight records range from June 6 to September 5, with peak of sorts in late June and
July.
Distribution. — Widespread from eastern seaboard of North America to northwestern
North America (Great Slave Lake and central British Columbia) (Map 62). Recorded in
Canada from Newfoundland to Great Slave Lk, central British Columbia, south to United
States border.
Quaest. Ent., 1987, 23 (1)
140
Nimmo
Hydropsyche tana Ross
Map 63; Fig. 402-408
Hydropsyche tana Ross, 1938b: 151; Ross, 1944:294; Schefter & Wiggins, 1986:77.
Description. — Male fore-wing length 8.42 mm; grey-brown, fairly uniformly irrorate. Antennae brownish straw;
basal seven flagellar annuli each with oblique, dark band. Vertex dark brown, posterior warts paler. Spurs yellow; lateral
member of middle and hind-leg pairs much shorter than mesal companions. Thorax dark brown, to grey-brown laterally.
Legs straw-coloured.
Genitalia. Male. (Fig. 402-406). (Specimen from creek, Hwy 37, 92 km N of Kitwanga, British Columbia). Males
distinguished by tergum X distal lobes, in lateral aspect (Fig. 402), massive, with narrower tip directed postero-ventrad; by
these lobes, in dorsal aspect (Fig. 403), curved postero-mesad; by distal article of clasper tapered slightly distad, with
dorsal edge of tip scalloped (Fig. 402); and by dorsal lobe of aedeagus (Fig. 405) with tooth (wide-based, with distal spine)
directed postero-laterad (Fig. 406).
Genitalia. Female. (Fig. 407-408). (Specimen from Teton R., Tetonia, Teton Co., Idaho, USA). Females
distinguished by clasper receptacle minute, set very high on lateral wall of segment X (Fig. 407); and by outer margin of
receptacle bowed distinctly ventrad.
Biology. — Nothing known, except known flight dates range from July 1 to 16.
Distribution. — Presently known only from western Montana, Idaho, British Columbia, and
Vancouver Island (Map 63). In Canada, recorded from just north of Skeena R., southern
Vancouver Island, and interior British Columbia.
Map 59. Collection localities for Hydropsyche bronta Ross in Canada, with known distribution in North America by state
or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
141
1
Map 60. Known distribution of Hydropsyche cheilonis Ross in North America, by state.
Map 61. Collection localities for Hydropsyche morosa Hagen in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987, 23 (1)
142
Nimmo
Map 62. Collection localities for Hydropsyche slossonae Banks in Canada, with known distribution in North America by
state or province.
Map 63. Collection localities for Hydropsyche tana Ross in Canada, with known distribution in North America by state or
province.
Arctopsychidae and Hydropsychidae (Trichoptera)
143
Fig. 374-380, Hydropsyche bronta Ross: 374, genital capsule of male, lateral aspect; 375, genital capsule of male, dorsal
aspect; 376, left clasper of male, posterior aspect; 377, aedeagus of male, lateral aspect; 378, aedeagus of male, dorsal
aspect of tip; 379, genital segments of female, lateral aspect; 380, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
144
Nimmo
Fig. 381-387, Hydropsyche cheilonis Ross: 381, genital capsule of male, lateral aspect; 382, genital capsule of male,
dorsal aspect; 383, left clasper of male, posterior aspect; 384, aedeagus of male, lateral aspect; 385, aedeagus of male,
dorsal aspect of tip; 386, genital segments of female, lateral aspect; 387, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
145
Fig. 388-394, Hydropsyche morosa Hagen: 388, genital capsule of male, lateral aspect; 389, genital capsule of male,
dorsal aspect; 390, left clasper of male, posterior aspect; 391, aedeagus of male, lateral aspect; 392, aedeagus of male,
dorsal aspect of tip; 393, genital segments of female, lateral aspect; 394, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
146
Nimmo
Fig. 395-401, Hydropsyche slossonae Banks: 395, genital capsule of male, lateral aspect; 396, genital capsule of male,
dorsal aspect; 397, left clasper of male, posterior aspect; 398, aedeagus of male, lateral aspect; 399, aedeagus of male,
dorsal aspect of tip; 400, genital segments of female, lateral aspect; 401, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
147
Fig. 402-408, Hydropsyche tana Ross: 402, genital capsule of male, lateral aspect; 403, genital capsule of male, dorsal
aspect; 404, left clasper of male, posterior aspect; 405, aedeagus of male, lateral aspect; 406, aedeagus of male, dorsal
aspect of tip; 407, genital segments of female, lateral aspect; 408, genital segments of female, dorsal aspect.
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148
Nimmo
SUBGROUP D
This group characterised by aedeagus with dorsal membranous lobes only, distal teeth, or
spines, of which are curved antero-dorsad.
Hydropsyche aenigma Schefter, Wiggins, & Unzicker
Hydropsyche aenigma Schefter, Wiggins, & Unzicker, 1986:78; Schefter & Wiggins, 1986:44.
This species, described from New York State in 1986, is very close to H. alternans (Walker)
and H. centra Ross. Time constraints precluded inclusion of illustrations here, but comparative
illustrations of all three species are given by the authors.
Hydropsyche alternans (Walker)
Map 64; Fig. 409-415
Philopotamus alternans Walker, 1852:104.
Hydropsyche alternans-, Vorhies, 1909:707 (sp. indet.); Betten, 1934:185 (prob. H. bifida)-, Nimmo, 1981:261 ( H .
recurvata as synonym); Schefter & Wiggins, 1986:48.
Hydropsyche slossonae v ar. recurvata Banks, 1914:253.
Hydropsyche recurvata-, Betten, 1934:190; Milne, 1936:73 (as synonym of H. slossonae ); Denning, 1943:126; Ross,
1944:99; Schmid, 1980:Fig. 131-140; Nimmo, 1981:261 (as synonym of H. alternans).
Symphitopsyche recurvata-, Schuster & Etnier, 1978:34.
Hydropsyche codona Betten, 1934:187; Milne, 1936:73 (as synonym of H. slossonae)-, Ross, 1 938c: 1 8 (as synonym of H.
recurvata).
Description. — Male fore-wing length 9.44 mm; grey-brown, clearly irrorate. Hind-wing faintly tinted brown.
Antennae pale brown; basal eight flagellar annuli each with oblique, dark band. Vertex dark brown anteriorly,
yellow-brown posteriorly. Spurs pale yellow-brown, to straw; lateral member of middle leg pairs notably shorter than mesal
companions. Thorax dark brown, to paler laterally.
Genitalia. Male. (Fig. 409-413). (Specimen from Wandering R., Hwy 63, Wandering River, Alberta). Males
distinguished by distal article of clasper, in posterior aspect (Fig. 411), acute-triangular; by small, membranous, dorsally
directed lobe located dorsally on aedeagus, between two dorsal lobes (Fig. 412); and by tergum X distal lobes narrow, well
separated, acuminate, in dorsal aspect(Fig. 410).
Genitalia. Female. (Fig. 414-415). (Specimen from Wandering R., Hwy 63, Wandering River, Alberta). Females
distinguished by clasper receptacle, in lateral aspect (Fig. 414), directed antero-dorsad, like inverted vase due to swelling;
by outer opening of receptacle with two grooves on floor; and by receptacle, in dorsal aspect (Fig. 415), directed
antero-mesad.
Biology. — Commonly collected from fast, cold waters, but known from warmer waters also.
Known from small creeks, to largest rivers, suggesting wide tolerance of habitat types. Ross
(1944) also records larvae from wave-washed shores of large lakes. Flight season ranges from
May 8 to October 16 in Canada, with peak in June/ July.
Distribution. — Very widespread, known from Alaska to Newfoundland, south to southern
British Columbia, St Lawrence R. valley in Canada, and to most states in United States, about
the Great Lakes (Map 64).
Hydropsyche centra Ross
Map 65; Fig. 416-422
Hydropsyche centra Ross, 1938b: 150; Ross, 1944:294; Anderson, 1976:66; Schefter & Wiggins, 1986:55.
Description. — Male fore-wing length 9.36 mm; uniform pale yellowish brown, faintly irrorate posterad of
Cul+2; irroration not evident in females seen. Hind-wing hyaline; pale reddish brown in female. Antennae pale
yellow-brown; basal seven flagellar annuli each with oblique, dark band. Vertex almost black, warts paler; dark red-brown
to chocolate in female. Spurs pale yellow-brown; lateral member of middle leg pairs shorter than mesal companions.
Thorax very deep red-brown, to paler laterally. Legs pale yellow to straw.
Arctopsychidae and Hydropsychidae (Trichoptera)
149
Genitalia. Male. (Fig. 416-420). (Specimen from Leaburg Dam, Mackenzie R., Lake Co., Oregon, USA). Males
distinguished by lobe on postero-ventral edge of segment IX long, narrow at base, rounded distally (Fig. 416); by tergum X
distal lobes, in dorsal aspect (Fig. 417), curved mesad with oval gap between lobes; and by teeth or spines of dorsal lobes of
aedeagus, in lateral aspect (Fig. 419), large, with short, straight base, with distal portion long, curved antero-dorsad from
base, acuminate.
Genitalia. Female. (Fig. 421-422). (Specimen from Lakelse, 18 miles of S Terrace, British Columbia). Females
distinguished by small clasper receptacle, in lateral aspect (Fig. 421), directed dorsad, distally rounded, without inner
opening visible; and by receptacle, in dorsal aspect (Fig. 422), directed meso-anterad.
Biology. — Little known. Flight season records for Oregon (Anderson, 1976) range from
late April to late September, with peak in May/ June. The few Canadian records fall within this
range.
Distribution.- — Presently known only from Oregon, Washington, British Columbia,
Vancouver Island (Map 65). In British Columbia the species has been recorded as far north as
Skeena R. basin.
Hydropsyche cockerelli Banks
Map 66; Fig. 423-429
Hydropsyche cockerelli Banks, 1905a; 14; Betten, 1934:193; Milne, 1936:70, 71, 73; Ross, 1944:294; Schefter, Wiggins, &
Unzicker, 1986:73; Schefter & Wiggins, 1986:58.
See note at end of ‘Introduction’.
Description. — Male fore-wing length 8.74 mm; pale greyish brown, faintly irrorate; female more orange-brown.
Hind-wing faintly tinted brown. Antennae pale yellow-brown; basal eight flagellar annuli each with oblique, dark band.
Vertex dark brown. Lateral member of middle and hind-leg spur pairs rather shorter than mesal companions. Thorax dark
reddish brown, to brown laterally. Legs yellowish cream.
Genitalia. Male. (Fig. 423-427). (Specimen from Waterton R., Hwy 5, Alberta). Males distinguished by tergum X
distal lobes, in lateral aspect (Fig. 423), directed postero-dorsad, with disto-dorsal angle produced as rounded, triangular
point; by these lobes, in dorsal aspect (Fig. 424), curved postero-laterad; and by distal article of clasper, in lateral aspect
(Fig. 423), skewed acute-triangular, with tip hooked slightly dorsad.
Genitalia. Female. (Fig. 428-429). (Specimen from Waterton R., Hwy 5, Alberta). Females distinguished by very
small clasper receptacle, in lateral aspect (Fig. 428), directed dorso-anterad, with outer margin located at mid-point of
receptacle; by receptacle, in dorsal aspect (Fig. 429), directed sharply anterad; and by sclerotised strap of vulval scale
large, widest at two-thirds distance from proximal end.
Biology. — Larvae seem to exhibit wide latitude in choice of habitat. Recorded from small,
sluggish prairie streams, from very large cordilleran rivers, and all sizes of foothills streams.
Flight season ranges from May 20 to September 5 in Canada, with no very obvious peak.
Anderson (1976) records the peak of emergence in Oregon as August/September.
Distribution. — Recorded from New Mexico, California, to southern Yukon (Map 66). In
Canada, widely recorded from plains and near-foothills areas of Alberta, with scattered records
from British Columbia, and Whitehorse, Yukon.
Hydropsyche jewetti Denning
Map 67; Fig. 430-434
Hydropsyche jewetti Denning, 1965:78; Schefter, Wiggins, & Unzicker, 1986:73 (as synonym of H. cockerelli).
See note at end of ‘Introduction’.
Description. — Male fore-wing length 10.14 mm; golden brown, faintly irrorate. Hind-wing very pale gold.
Antennae pale brownish yellow; basal nine flagellar annuli each with oblique, dark band. Vertex deep chocolate, slightly
paler posteriorly. Spurs pale yellow-brown; lateral member of middle leg pairs markedly shorter than mesal companions.
Thorax dark chocolate-brown, to lighter laterally. Legs pale straw.
Genitalia. Male. (Fig. 430-434). (Specimen from 1-mile Ck, Hwy 5, N of Princeton, British Columbia). Males
distinguished by tergum X distal lobes, in lateral aspect (Fig. 430), directed postero-dorsad, slightly cleft distally to two
rounded lobes; by distal article of clasper, in lateral aspect, acute-triangular; by tergum X distal lobes, in dorsal aspect
(Fig. 431), short, acuminate, with tips turned slightly mesad; and by distal teeth of aedeagus dorsal lobes small, curved
Quaest. Ent., 1987,23 (1)
150
Nimmo
only slightly at base (Fig. 433).
Genitalia. Female. Unknown.
Biology. — Almost nothing known. 1-mile Creek is a small, rocky stream flanked by Poplar
and farmland. Date of collection was July 13. Newell & Potter (1973) give June/July as
Montana flight season.
Distribution. — Presently known only from western Montana and south-central British
Columbia (Map 67).
Map 64. Collection localities for Hydropsyche alternans (Walker) in Canada and Alaska, with known distribution in
North America by state or province.
Map 65. Collection localities for Hydropsyche centra in Canada, with known distribution in North America by state or
province.
Arctopsychidae and Hydropsychidae (Trichoptera)
151
Map 66. Collection localities for Hydropsyche cockerelli Banks in Canada, with known distribution in North America by
state or province.
Map 67. Collection localities for Hydropsyche jewetti Denning in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987,23 (1)
152
Nimmo
Fig. 409-415, Hydropsyche alternans (Walker): 409, genital capsule of male, lateral aspect; 410, genital capsule of male,
dorsal aspect; 411, left clasper of male, posterior aspect; 412, aedeagus of male, lateral aspect; 413, aedeagus of male,
dorsal aspect of tip; 414, genital segments of female, lateral aspect; 415, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
153
Fig. 416-422, Hydropsyche centra Ross: 416, genital capsule of male, lateral aspect; 417, genital capsule of male, dorsal
aspect; 418, left clasper of male, posterior aspect; 419, aedeagus of male, lateral aspect; 420, aedeagus of male, dorsal
aspect of tip; 421, genital segments of female, lateral aspect; 422, genital segments of female, dorsal aspect.
Quaest. Ent., 1987,23 (1)
154
Nimmo
Fig. 423-429, Hydropsyche cockerelli Banks: 423, genital capsule of male, lateral aspect; 424, genital capsule of male,
dorsal aspect; 425, left clasper of male, posterior aspect; 426, aedeagus of male, lateral aspect; 427, aedeagus of male,
dorsal aspect of tip; 428, genital segments of female, lateral aspect; 429, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
155
Fig. 430-434, Hydropsyche jewetti Denning: 430, genital capsule of male, lateral aspect; 431, genital capsule of male,
dorsal aspect; 432, left clasper of male, posterior aspect; 433, aedeagus of male, lateral aspect; 434, aedeagus of male,
dorsal aspect of tip.
Quaest. Ent., 1987, 23 (1)
156
Nimmo
SUBGROUP E
This subgroup characterised by aedeagus dorsal lobes with one large tooth (distally &
laterally), and cluster of small spines (laterally & distally).
Hydropsyche riola Denning
Map 68; Fig. 435-441
Hydropsyche riola Denning, 1942:49; Denning, 1943:133; Ross, 1944:294; Schefter, Wiggins, & Unzicker, 1986:69 (as
synonym of H. alhedra).
Symphitopsyche riola\ Schuster & Etnier, 1978:44.
See note at end of ‘Introduction’.
Description. — Male fore-wing length 9.52 mm; bright grey-brown, with alternate areas of colour and hyaline
membrane; female more generally irrorate. Antennae brown; basal eight flagellar annuli each with oblique, dark band.
Vertex deep brown anteriorly, to deep orange-brown posteriorly. Spurs yellow-brown; lateral member of middle and
hind-leg pairs notably shorter than mesal companions. Thorax deep brown. Legs brownish yellow to straw.
Genitalia. Male. (Fig. 435-439). (Specimen from creek, Hwy 932, 6 miles S of Whitecourt, Alberta). Males
distinguished by tergum X distal lobes, in dorsal aspect (Fig. 436), close together, long, evenly tapered, curved slightly
mesad, with long elliptical gap between; by basal article of clasper, in posterior aspect (Fig. 437), with very narrow basal
half; and by aedeagus dorsal lobe, in lateral aspect (Fig. 438), with long, slender, acuminate tooth distally, and cluster of
spines mid-way to tooth, on posterior face of lobe.
Genitalia. Female. (Fig. 440-441). (Specimen from creek, Hwy 932, 6 miles S of Whitecourt, Alberta). Females
distinguished by clasper receptacle, in dorsal aspect (Fig. 441), directed mesad; by receptacle, in lateral aspect (Fig. 440),
traversed by thin, dark line from dorsum of segment X; and by sclerotised strap of vulval scale long, sinuate, widened
distally.
Biology. — My records indicate that larvae are not especially restricted in stream types
occupied. Adult collecting sites include sluggish, silty prairie streams; turbulent,
boulder-bottomed small rivers; smooth-flowing, earth-banked boreal creeks; and weed-filled
streamlets. Flight season records range from May 25 to August 8, in Canada.
Distribution. — Recorded in narrow zone across North America from Massachusetts to
Alaska (Map 68). Other than in Saskatchewan and Alberta, the known distribution is
disjointed.
Hydropsyche sparna Ross
Map 69; Fig. 442-448
Hydropsyche sparna Ross, 1 938b: 1 50; Denning, 1943:134; Ross, 1944:97; Schefter & Wiggins, 1986:74.
Symphitopsyche sparna-, Schuster & Etnier, 1978:52.
Description. — Male fore-wing length 8.27 mm; pale golden brown, no patter evident. Hind-wing faintly tinted
golden brown. Antennae yellow; no dark banding on flagellar annuli. Vertex brownish yellow; orange-brown in female.
Spurs yellow; lateral member of middle leg pairs markedly shorter than mesal companions. Thorax brownish yellow;
orange-brown in female. Legs yellow to straw.
Genitalia. Male. (Fig. 442-446). (Specimen from Flanders, Morris Co., New Jersey, USA). Males distinguished by
distal article of clasper, in lateral aspect (Fig. 442), abruptly narrowed, from wide base, to long, thin, slightly curved
process; by tergum X distal lobes, in dorsal aspect (Fig. 443), directed posterad, slightly twisted, with wide, u-shaped gap
between; and by aedeagus dorsal lobe, in lateral aspect (Fig. 445), with distal cluster of spines, and tooth (wide-based,
abruptly narrowed to distal spine) located at mid-point on posterior face.
Genitalia. Female. (Fig. 447-448). (Specimen from Flanders, Morris Co., New Jersey, USA). Females distinguished
by clasper receptacle, in lateral aspect (Fig. 447), with thin, dark line from dorsum of segment X terminated at dorsal
extremity of receptacle; by receptacle, in dorsal aspect (Fig. 448), directed postero-mesad; and by vulval scale sclerotised
strap thin, angled.
Biology. — Larvae exhibit wide ecological tolerance, from sluggish, small, organically rich
streams, to fast, clear, cold trout-stream waters (Schuster & Etnier, 1978). Canadian flight
season ranges from May 17 to September 26, with bulk of records from August.
Arctopsychidae and Hydropsychidae (Trichoptera)
157
Distribution. — Predominantly an eastern species, recorded from Georgia to Newfoundland,
west across the Appalachians, with extension from Michigan to Manitoba (Map 69). In
Canada, known from Duck Mountain Provincial Park, on western boundary of Manitoba, to
Labrador and Newfoundland. From the above it may be surmised that the species is confined to
woodland waters.
Map 68. Collection localities for Hydropsyche riola Denning in Canada and Alaska, with known distribution in North
America by state or province.
Map 69. Collection localities for Hydropsyche sparna Ross in Canada, with known distribution in North America by state
or province.
Quaest. Ent., 1987, 23 (1)
158
Nimmo
Fig. 435-441, Hydropsyche riola Denning: 435, genital capsule of male, lateral aspect; 436, genital capsule of male, dorsal
aspect; 437, left clasper of male, posterior aspect; 438, aedeagus of male, lateral aspect; 439, aedeagus of male, dorsal
aspect of tip; 440, genital segments of female, lateral aspect; 441, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
159
Fig. 442-448, Hydropsyche sparna Ross: 442, genital capsule of male, lateral aspect; 443, genital capsule of male, dorsal
aspect; 444, left clasper of male, posterior aspect; 445, aedeagus of male, lateral aspect; 446, aedeagus of male, dorsal
aspect of tip; 447, genital segments of female, lateral aspect; 448, genital segments of female, dorsal aspect.
Quaest. Ent., 1987, 23 (1)
160
Nimmo
SUBGROUP F
This group characterised by aedeagus dorsal membranous lobes without teeth or spines in
any form.
Hydropsyche oslari Banks
Map 70; Fig. 449-455
Hydropsyche oslari Banks, 1 905a: 1 3; Betten, 1934:194; Milne, 1936:70, 71, 73; Ross, 1944:294; Schefter & Wiggins,
1986:66.
Hydropsyche partita Banks, 1914:252; Milne, 1936:73.
Description. — Male fore-wing length 9.91 mm; pale grey-brown, uniformly irrorate, with scattered larger areas of
uniform colour. Antennae brownish cream; basal seven flagellar annuli each with oblique, dark band. Vertex dark brown
anteriorly, to yellow-brown posteriorly. Spurs yellow to cream; lateral member of middle leg pairs notably shorter than
mesal companions; not so noticeable in female. Thorax rich red-brown, to yellow-brown laterally. Legs yellowish cream.
Genitalia. Male. (Fig. 449-453). (Specimen from Oldman R., Hwy 922, Alberta). Males distinguished by distal
article of clasper, in lateral aspect (Fig. 449), evenly, gradually narrowed from base to long, dorsally curved distal portion;
by distal article, in posterior aspect (Fig. 451), with very long, slender, linear tip; and by dorsal lobes of aedeagus with
dorsally directed median lobe between two minute lateral lobes (Fig. 452, 453).
Genitalia. Female. (Fig. 454-455). (Specimen from Oldman R., Hwy 922, Alberta). Females distinguished by clasper
receptacle, in lateral aspect (Fig. 454), located high on lateral wall of segment X, directed dorso-anterad, without evident
inner opening; by receptacle, in dorsal aspect (Fig. 455), directed mesad; and by vulval scale sclerotised strap long,
gradually widened distally, angled at mid-point (Fig. 454).
Biology. — I have records of adults from shallow, small, rock and gravel streams of
negligible turbulence, to larger rivers of swift, deep, turbulent waters (clear or turbid).
Canadian flight season ranges from June 4 to September 14, with diffuse peak through
July/August.
Distribution. — Western species, ranging from Mexico to western Yukon (Map 70). In
Canada, widely recorded from open plains, foothills, and Boreal Forest of Alberta, from
throughout British Columbia, from Vancouver Island, with one record from Dawson City,
Yukon.
Hydropsyche ventura Ross
Map 71; Fig. 456-462
Hydropsyche ventura Ross, 1941:92; Ross, 1944:294; Schefter & Wiggins, 1986:80.
Symphitopsyche ventura-, Schuster & Etnier, 1978:55.
Description. — Male fore-wing length 8.38 mm; uniformly very pale reddish brown; anal edge with alternate dark
and pale areas. Hind-wing very palely tinted. Antennae uniform pale straw. Vertex deep red-brown, warts paler. Spurs
straw-coloured. Thorax deep red-brown overall. Legs straw-coloured.
Genitalia. Male. (Fig. 456-459). (Specimen from St Hippolyte, Quebec). Males distinguished by distal article of
clasper, in lateral aspect (Fig. 456), of almost uniform width, with triangular tip; by tergum X distal lobes, in dorsal aspect
(Fig. 457), long, slender, well spaced, curved somewhat mesad about roughly circular gap between; and by aedeagus dorsal
lobes, in dorsal aspect (Fig. 460), each bilobed, with posterior short, stout lobe, and anterior long, thin lobe.
Genitalia. Female. (Fig. 461-462). (Specimen from St Hippolyte, Quebec). Females distinguished by clasper
receptacle, in lateral aspect (Fig. 461), oriented vertically, directed anterad, crescentic, with inner opening not visible; by
receptacle, in dorsal aspect (Fig. 462), directed mesad, with inner opening on small secondary tubercle; and by vulval scale
sclerotised strap with long, slender, slightly widened proximal portion, semi-circular distal portion.
Biology. — Little known. Schuster & Etnier (1978) describe one particular site in detail, but
make no generalisations. They give known flight season as early April to September.
Distribution. — Recorded from Tennessee to Newfoundland, with scattered records from
southern Quebec, eastern Ontario (Map 71).
Arctopsychidae and Hydropsychidae (Trichoptera)
161
Map 70. Collection localities for Hydropsyche oslari Ross in Canada, with known distribution in North America by state
or province.
Map 71. Collection localities for Hydropsyche ventura Ross in Canada, with known distribution in North America by
state or province.
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162
Nimmo
Fig. 449-455, Hydropsyche oslari Ross: 449, genital capsule of male, lateral aspect; 450, genital capsule of male, dorsal
aspect; 451, left clasper of male, posterior aspect; 452, aedeagus of male, lateral aspect; 453, aedeagus of male, dorsal
aspect of tip; 454, genital segments of female, lateral aspect; 455, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
163
Fig. 456-462, Hydropsyche ventura Ross: 456, genital capsule of male, lateral aspect; 457, genital capsule of male, dorsal
aspect; 458, left clasper of male, posterior aspect; 459, aedeagus of male, lateral aspect; 460, aedeagus of male, dorsal
aspect of tip; 461, genital segments of female, lateral aspect; 462, genital segments of female, dorsal aspect.
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164
Nimmo
Genus Potamyia Banks
Map 72; Fig. 6, 463-468
Potamyia Banks, 1900:259; Betten, 1934:197; Ross, 1944:85; Wiggins, 1977:1 16; Schmid, 1980:59.
Description. — Head globular, convex; malar space large. Antennae very fine, long, especially in male; basal
article globular. Fore-wings (Fig. 6a) with few hairs. Hind-wings (Fig. 6b) large, especially in male; somewhat pointed
distally. Hind-wing fl present; stems of veins M and Cul very close, parallel; cross-veins M3 + Cul and Cul-Cu2 very
close together. Fore- wing without cross-veins Sc-Rl and R1-R2 + R3. Spur formula 0,4,4 male; 1,4,4 female. Fore-leg
tarsi of male strongly spinate; claws asymmetrical, overhung by stout, black setae.
Genitalia. Male. (Fig. 463-466). Very similar to genitalia of Hydropsyche and Cheumatopsyche.
Genitalia. Female. (Fig. 467-468). Again, very similar to genitalia of Hydropsyche and Cheumatopsyche. Sternite
VIII longitudinally divided throughout. Clasper receptacle not present (Fig. 467).
The above characters are a melange of characters shared with the Macronematinae,
Hydropsyche , and Cheumatopsyche, plus several peculiar to Potamyia.
Biology. — Larvae prefer large, warmish rivers, and appear to congregate on rocks in sandy,
silt-free conditions in slower currents.
Potamyia is confined to Siberia and eastern North America east of Montana. Of two known
species one, P. flay a, is recorded from North America.
Potamyia flava (Hagen)
Map 72; Fig. 6, 463-468
Macronema flavum Hagen, 1861:285.
Potamyia flava\ Banks, 1900:259; Betten, 1934:198; Denning, 1943:136; Ross, 1944:85; Wiggins, 1977:Fig. 6.11; Schmid,
1980:Fig. 141-143.
Hydropsyche flava\ Milne, 1936:73.
Hydropsyche kansasensis Banks, 1905a:15; Milne, 1936:73.
Description. — Male fore-wing length 9.20 mm; uniform light brownish yellow; no pattern. Antennae light
yellow-brown; basal two flagellar annuli each with oblique, paler band; less evident in female. Vertex light yellow-brown;
uniform brownish yellow in female. Spurs pale, dull yellow; in female, lateral member of middle leg pairs, and hind-leg
apical pair, notably shorter than mesal companions. Thorax pale, dull yellow-brown throughout. Legs dull straw; reddish
yellow in female.
Genitalia. Male. (Fig. 463-466). (Specimen from Hamilton, Illinois, USA). Males distinguished by high, narrow
segment IX in lateral aspect (Fig. 463); by small, acuminate, dorso-laterally directed distal lobes of tergum X (Fig.
463-466); and by distal article of clasper, in lateral aspect, widened distally.
Genitalia. Female. (Fig. 467-468). (Specimen from Washington Co., Arkansas, USA). Females distinguished by lack
of any sign of clasper receptacle on lateral wall of segment X (Fig. 467); and by lack of vulval scale sclerotised strap.
Biology. — See under genus above. The only Canadian flight date is June 6, in Manitoba.
Distribution.— Widespread throughout eastern United States (Map 72), as far west as
Montana; not known from New England States. In Canada Schmid (1980) reports this species
from southern Ontario. I have examined material taken on the banks of the Assiniboin River,
Manitoba (just W of Winnipeg).
Arctopsychidae and Hydropsychidae (Trichoptera)
165
Map 72. Collection localities for Potamyia flava (Hagen) in Canada, with known distribution in North America by state
or province.
Quaest. Ent., 1987, 23 (1)
166
Nimmo
Fig. 463
-468, Potamyia flava (Hagen): 463, genital capsule of male, lateral aspect; 464 left clasper of male posterior
165, aedeagus of male, lateral aspect; 466, genital capsule of male, dorsal aspect; 467, gemtal segments of female.
aspect; 465, aedeagus of male, tatum e --
lateral aspect; 468, genital segments of female, dorsal aspect
Arctopsychidae and Hydropsychidae (Trichoptera)
167
SUBFAMILY DIPLECTRONINAE ULMER
Diplectroninae Ulmer, 1951:303; Ross, 1956:10; Marlier, 1962:135; Wiggins, 1977:93; Schmid, 1980:61.
Following characterisation of adults derived from Marlier (1962).
Description. — Medium to large (wing-span 12-20 mm). Wings generally large; wide basally, rounded or angular
distally. Antennae equal in length to fore-wings, or slightly longer. Cephalic warts less prominent than in
Macronematinae; posterior pair large, oval; anterior pair quite small; interantennal wart round. Thorax large, robust. Male
fore-leg tarsal claws normal, equal, not overhung by setal tufts. Middle leg tarsi of female not expanded, not flattened.
Spur formula 2,4,4. Fore-wing Sc and R1 (Fig. 4a) complete, unfused, or joined distally; often robust. Discoidal and
median cells small, subequal, closed. Forks fl-fV present; fl, fill, and fV petiolate. Thyridial cell closed, in contact with
median. Hind-wing (Fig. 4b) often widened at mid-point; Sc and R1 distinct throughout; discoidal cell closed, elongate;
fll, fill, and fV present; median cell open.
Genitalia. Male. (Fig. 469-472, 475-479). Much as usual for Hydropsychidae; simple, very little modified. Tergum X
roof-like dorsad of aedeagus; not seperable from segment IX; with or without wart-like preanal appendages. Aedeagus
simple (Fig. 478) or complex (Fig. 471), curved, basally expanded, without spines. Claspers (inferior appendages) long, of
two articles; distal article smaller than basal article.
Genitalia. Female. (Fig. 473-474, 480-481). Much as for Hydropsychinae. With or without clasper receptacles; if
present, not prominent. Cerci prominent.
Genus Aphropsyche Ross
Map 73; Fig. 469-474
Aphropsyche Ross, 1941:78; Ross, 1944:83.
Following generic characterisation derived from Ross (1941).
Description. — Head somewhat prognathoid; eyes widely separated, located anterad. Antennae short, somewhat
robust; pedicel only half as long as scape, of thickness similar to flagellum. Vertex convex; mesal ridges inconspicuous;
postero-lateral warts large. Pronotum with pair of large, close warts. Wings similar in shape, evenly rounded distally;
venation generalised, typical for group; radial veins of hind-wing straight.
Genitalia. Male. (Fig. 469-472). Genital capsule basically as for Hydropsychinae, but considerably less regular in
outline (Fig. 469); with postero-ventral edge produced posterad. Clasper articles not obviously distinguishable. Aedeagus
complex (Fig. 471), with paired (Fig. 472), long, slender, acuminate dorsal lobes; with expanded, rounded, single ventral
lobe surmounted by smaller, complex intromittent structure.
Genitalia. Female. (Fig. 473-474). Much as for Hydropsychinae. Cercus large, prominent. No clasper receptacle
evident. Sternite VIII cleft mesally, more or less throughout length.
Biology. — As larvae not yet definitely associated (see Wiggins, 1977), nothing can usefully
be included here.
Four species presently known: two in Asia, two in eastern North America. One is likely to be
recorded in eastern Canada.
Aphropsyche doringa Milne
Map 73; Fig. 469-474
Aphropsyche doringa Milne, 1936:68-69, 73; Ross, 1944:294.
Aphropsyche aprilis Ross, 1941:78; Ross, 1944:83; Flint, 1966:374 (as synonym of A. doringa).
Description. — Male fore-wing length 8.81 mm; uniform purple-grey, with stigma and anal edge darker.
Hind-wing uniform purple-grey. Female overall darker, more uniform in fore-wing. Antennae uniform dull brown in male;
dark purple-brown in female. Vertex deep chocolate to black, warts red-brown, intense black in female with reddish black
warts. Spurs purple-brown. Thorax, dorsally, as vertex in either sex; laterally, deep red-brown (to yellow-brown coxae) -
deep red-brown throughout in female. Legs yellow-brown proximally, purple-brown distally; dull red-brown throughout in
female.
Genitalia. Male. (Fig. 469-472). (Specimen from Reid Ck, Blount Co., Tennessee, USA). Males distinguished on
basis of aedeagus alone (Fig. 471-472), with paired dorsal lobes, single ventral lobe. Other features are: clasper apparently
of one article, curved gently dorsad, of uniform width except for tapered distal end; anterior edge of segment IX highly
irregular in lateral aspect (Fig. 469); postero-ventral edge of segment IX with posteriorly projected shelf; and tergum X
distal lobes high, black, flared postero-laterad, with slender, acuminate dorsal spine each.
Quaest. Ent., 1987, 23 (1)
168
Nimmo
Genitalia. Female. (Fig. 473-474). (Specimen from Reid Ck, Blount Co., Tennessee, USA). Females distinguished by
vulval scale sclerotised, except membranous distally (Fig. 474); by anterior edge of segment X linear (Fig. 473); and by
cercus large, intermediate between dorsal and ventral lobes of segment XI, with minute distal article.
Biology. — Almost unknown. Etnier & Schuster (1979) report adult occurrence in
Tennessee as April 21 to May 13. Neves (1979) reports adults in Massachusetts in June.
Distribution. — Not yet known from Canada, with only scattered records from eastern
United States (Map 73), from as far north as New Hampshire.
Map 73. Known distribution of Aphropsyche doringa Milne in North America, by state.
Arctopsychidae and Hydropsychidae (Trichoptera)
169
Fig. 469-474, Aphropsyche doringa Milne: 469, genital capsule of male, lateral aspect; 470, left clasper of male, posterior
aspect; 471, aedeagus of male, lateral aspect; 472, aedeagus of male, dorsal aspect; 473, genital segments of female, lateral
aspect; 474, genital segments of female, ventral aspect.
Quaest. Ent., 1987, 23 (1)
170
Nimmo
Genus Diplectrona Westwood
Map 74; Fig. 475-481
Diplectrona Westwood, 1840:49; McLachlan, 1978:374; Betten, 1934:182; Milne, 1936:68; Ross, 1944:84; Bull. Zool.
Nomen, 1965:288 (validation: opinion 758); Wiggins, 1977:102; Schmid, 1980:61.
Aphelocheira Stephens, 1836:167, 179; Bull. Zool. Nomen., 1965:288 (suppresion: opinion 758).
Description. — Antennae as long as fore-wings, fine, notched; annuli as thick as long, swollen at mid-point; basal
annuli each with oblique, dark band. Maxillary palpi with second article much longer than basal article, third and fourth
articles shorter. Vertex with four large warts. Middle leg tarsi of female normal, not flattened, not enlarged. Sternite V
with bulge prolonged by very long, thin tube, slightly longer in male than female. Fore-wing (Fig. 4a) widened at chord.
Hind-wing (Fig. 4b) large, rounded, distally blunt. Fore-wing fl and fill petiolate; discoidal cell small, median and
thyridial cells large; Cu2 and A markedly curved, postcostal cell much enlarged; cross-veins M3 + 4-Cul and Cul-Cu2
close. Hind-wing with Sc very thick, R1 very thin, both sinuate, subcostal cell very narrow basally, very large distally;
discoidal cell narrow; fl and fill petiolate.
Genitalia. Male. (Fig. 475-479). Segment IX without postero-lateral edge poorly developed (Fig. 475). Segment X
not clearly delimited from IX, with two pairs of lobes (one mesal, the other lateral) (Fig. 476). Claspers (inferior
appendages) with very long basal article. Aedeagus (Fig. 478) stout, two endothecal lobes distally (Fig. 479).
Genitalia. Female. (Fig. 480-481). Sternite VIII completely divided. Segment X with barely visible clasper receptacle
(Fig. 480), produced ventrad to form distinct sclerotised strap along lateral face of vulval scale.
Biology. — Larvae found in rapid areas of small, cool streams. They have been found in
moss on submerged rocks, and in leaf accumulations. Univoltine, with egg hatch in late
Summer and Autumn; larvae, never abundant, mature in early summer.
Diplectrona spp. known from all regions except Ethiopian and Neotropical Regions, and
Antarctic continent. Three species known in North America - two eastern, one western. One
known from Canada.
Diplectrona modesta Banks
Map 74; Fig. 475-481
Diplectrona modesta Banks, 1908:266; Betten, 1934:182; Milne, 1936:68, 73; Ross, 1944:84; Wiggins, 1977:102; Schmid,
1980:Fig. 150-157.
Description. — Male fore-wing length 8.74 mm; uniform grey-brown. Hind-wings palely tinted grey-brown.
Antennae red-brown. Vertex dark red-brown. Spurs brown, dark brown in female; lateral member of fore-leg pair minute
relative to mesal companion; equal in female. Thorax dark red-brown, to paler laterally. Legs dark straw-brown.
Genitalia. Male. (Fig. 475-479). (Specimen from St Hippolyte, Quebec). Males distinguished by distal article of
clasper of uniform width, much narrower than basal article (Fig. 475, 477); by dorsum of segment IX long; and by tergum
X with two pairs of lobes, in dorsal aspect (Fig. 476).
Genitalia. Female. (Fig. 480-481). (Specimen from Milton, Ontario). Females distinguished by clasper receptacle
located high on lateral wall of segment X as minute, circular pit (Fig. 480); and by vulval scale sclerotised strap long,
widened distally, attached to basal angle of segment X by long, irregular root.
Biology. — As above, except to add that known flight season in Canada extends from May
30 to September 26.
Distribution. — Eastern seaboard of North America from Florida to Cape Breton Island,
Nova Scotia, northwest to American mid-western states and northwestern Ontario. There are
records from Arkansas and Oklahoma (Map 74). In Canada, recorded from near Dryden,
northwestern Ontario, to Baddeck, Cape Breton Island, Nova Scotia, south to United States
border.
Arctopsychidae and Hydropsychidae (Trichoptera)
171
Map 74. Collection localities for Diplectrona modesta Banks in Canada, with known distribution in North America by
state or province.
Quaest. Ent., 1987,23 (1)
172
Nimmo
Fig. 475-481, Diplectrona modesta Banks: 475, genital capsule of male, lateral aspect; 476, genital capsule of male, dorsal
aspect; 477, left clasper of male, posterior aspect; 478, aedeagus of male, lateral aspect; 479, aedeagus of male, dorsal
aspect of tip; 480, genital segments of female, lateral aspect; 481, genital segments of female, dorsal aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
17?
SUBFAMILY MACRONEMATINAE ULMER
Macronematinae Ulmer, 1 905a:4 1 ; Betten, 1934:200; Milne, 1936:67; Wiggins, 1977:93; Schmid, 1980:56.
Oestropsinae Brauer, 1868. Denning, 1943:106, 155 (for Macronematinae).
As only one genus is at all likely to concern us here, the generic characterisation will suffice
for purposes of this text. Those wishing full details of the subfamily should consult Betten
(1934:200) and Wiggins (1977:93).
Genus Macrostemum Kolenati
Maps 75-77; Fig. 482-496
Macrostemum Kolenati, 1859; Flint & Bueno-Soria, 1982:358.
Macronema Pictet, 1836:400 (part); McLachlan, 1878:353 (part); Betten, 1934:203 (part); Milne, 1936:72; Denning,
1943:156 ( Macronemum)\ Ross, 1944:114 ( Macronemum ); Bull. Zool. Nomen., 1962:80 (validation - opinion 623);
Wiggins, 1977:110; Schmid, 1980:56.
Description. — Distinct secondary sexual dimorphism present. Head globular, vertex convex, malar space large.
Antennae very fine, 1.5 times length of fore-wing, 1.3 times in female. Anterior warts of vertex very large, slightly less so
in female. Maxillary palpi with two basal articles short; male flagellum 1.3 times longer than in female. Middle leg tarsi of
female much enlarged, flattened. Hind-leg tibia of male with long setae. Male sternite V with finger-like lobe; minute
bump in female. Wings sparsely hirsute; fore- and hind-wings of different shapes; fore-wing slenderly elliptical; hind-wing
triangular, 1.5 times size of fore-wing, not so large in females; female hind-wing with distinct costal angle. Hind-wing
venation somewhat irregular, incomplete. Fore-wing Sc and R1 joined prior to distal edge; fl-FV present, fl petiolate;
discoidal, median cells small, thyridial cell very long, postcostal cell very large; A1 very long. Only fll, fill, and fV present
in hind-wing; R1 and R2 + 3 joined with Sc; discoidal cell open; M with origin at base of RS. Fore-wing variously banded
transversely with light and dark colour.
Genitalia. Male. (Fig. 483-487, 490-494). Segment IX, in lateral aspect (Fig. 490), elongate; short dorsally and
ventrally. Segment X comprised of two large lobes (Fig. 491), produced postero-laterad; meso-internal area membranous.
Claspers (inferior appendages) long, of more or less uniform thickness, of two subequal articles. Aedeagus fairly stout
basally, greatly swollen distally, simple (Fig. 493, 494).
Genitalia. Female. (Fig. 488-489, 496). Sternite VIII cleft twice on posterior edge, to form three lobes, or once to
form two lobes (Fig. 489). Segment X roughly triangular in lateral aspect (Fig. 488, 496); ventral angle produced, vulval
scale enclosed. Segment XI slightly developed; normal two papillate lobes of hydropsychids slender, distinct; space between
lobes occupied by several smaller, papillate, membranous processes. Cerci small, of two articles.
Biology. — Larvae inhabit larger rivers, and ingest fine particulate detritus, phytoplankton,
and bacteria.
Macrostemum is a widespread genus, found in Africa, Asia, Australia, with three species
known from eastern North America.
Key to known or potential species of Macrostemum Kolenati of Canada
la Wings pale yellow, with narrow, transverse brown stripes (more diffuse in
female) M. transversum (Walker), p. 174
lb Wings with longitudinal stripes (purplish brown) on basal part, transverse
stripes distad of these, with irregular patch of brown on distal quarter of
wing (more diffuse in females, with smaller patches of brown) 2
2a (lb) Compound eye large relative to head in lateral aspect (Fig. 482), vertex low
M. Carolina (Banks), p. 174
2b Compound eye small relative to head in lateral aspect (Fig. 495), vertex
high M. zebratum (Hagen), p. 174
Quaest. Ent., 1987, 23 (1)
174
Nimmo
Macrostemum Carolina (Banks)
Map 75; Fig. 482
Macronema Carolina Banks, 1909:342; Betten, 1934:204; Milne, 1936:73, 74; Ross, 1944:1 16 ( Macronemum ).
Macrostemum Carolina-, Flint & Bueno-Soria, 1982:369.
Description. — Male fore-wing length 10.06 mm; purplish brown with large areas of yellow-brown disposed in
longitudinal bars (basal) or transverse bars (mid-way to tip), with irregular patch in distal quarter. Female pattern more
diffuse; dark areas paler distally, with distal quarter almost all pale yellow-brown. Hind-wing translucent pale brown,
almost hyaline along costal edge. Antennae dark reddish brown. Vertex dark purplish brown. Spurs yellowish brown;
lateral member of middle leg pairs, and hind-leg apical pair, notably shorter than mesal companions. Thorax virtually
black, to very deep purplish brown laterally. Legs brown to yellow. Eyes large relative to head, malar space below not wide,
rather narrow (Fig. 482). Vertex low in lateral aspect - this point will separate M. Carolina from M. zebratum.
Genitalia. Male. (Not illustrated - identical to those of M. zebratum). (Specimen from Washington Co., Arkansas,
USA). M. Carolina and M. zebratum may be separated from M. transversum by segment IX and tergum X with
membranous partial gap between, in lateral aspect (Fig. 490); and by tergum X distal lobes, in dorsal aspect (Fig. 491),
with mesal edges parallel, gap between u-shaped.
Genitalia. Female. (Not illustrated - identical to those of M. zebratum). (Specimen from Washington Co., Arkansas,
USA). M. Carolina and M. zebratum may be distinguished from M. transversum by bases of cerci and lobes of segment XI
enclosed laterally by postero-lateral edge of segment X; and by segment X wide from top to bottom (Fig. 496).
Biology. — South Carolina flight season is May to September. Wallace & Scherberger
(1974) discuss larval retreat and net in detail.
Distribution. — Not yet recognised from Canada. In United States, recorded from most
states from Oklahoma east to Florida, Illinois, and New York (Map 75).
Macrostemum transversum (Walker)
Map 76; Fig. 483-489
Hydropsyche transversa Walker, 1852:1 14
Macronema transversum-, McLachlan, 1866:264; Betten, 1934:205; Milne, 1936:72, 74; Ross, 1944:117 {Macronemum)-,
Kimmins & Denning, 1951:120.
Macrostemum transversum-, Flint & Bueno-Soria, 1982:369.
Macronema polygrammatum McLachlan, 1871:129; Betten, 1934:204, 205 (A/, polygrammaticum)-, Kimmins &
Denning, 1951:106 (as synonym of M. transversum).
Description. — Male fore-wing length 12.95 mm; pale yellow-brown with deep red-brown patches and bars; distal
area of yellow-brown as transverse bar. Female fore-wing with dark patches more diffuse; distal transverse bar less
distinct, extended to costal edge of wing. Hind-wing hyaline. Antennae pale orange-brown. Vertex very wide, short; deep
red-brown, warts paler. Spurs yellow; lateral member of all pairs shorter than mesal companions; most noticeable on
middle leg. Thorax deep red-brown throughout; venter clothed in long, very fine, hyaline pubescence. Legs straw-coloured;
clothed in long, very fine, hyaline hairs.
Genitalia. Male. (Fig. 483-489). (Specimen from Altahana R., Appling Co., Georgia, USA). Males distinguished by
pattern of sinuate, branched lines on segment IX and tergum X in lateral aspect (Fig. 483); by tergum X distal lobes, in
dorsal aspect, separated by v-shaped gap (Fig. 484); and by no distinct division between segment XI and tergum X.
Genitalia. Female. (Fig. 488-489). (Specimen from Altahana R., Appling Co., Georgia, USA). Females distinguished
by cerci and segment XI lobes entirely posterad of any overlap by lateral wall of segment X (Fig. 488); and by segment X
narrow from top to bottom, in lateral aspect.
Biology. — No flight season data available. Wallace & Sherberger (1975) provide a detailed
account of larval retreat and feeding net.
Distribution. — Not yet known from Canada. United States records scattered (Map 76),
including Washington D.C., Virginia, Georgia, Indiana, and Ohio.
Macrostemum zebratum (Hagen)
Map 77; Fig. 490-496
Macronema zebratum Hagen, 1861:285; Betten, 1934:205; Milne, 1936:72, 74; Denning, 1943:157; Ross, 1944:115
{Macronemum)-, Wiggins, 1977:110; Schmid, 1 980: Fig. 124-130.
Arctopsychidae and Hydropsychidae (Trichoptera)
175
Macrostemum zebratum-, Flint & Bueno-Soria, 1982:369.
Phryganea ( Leptocerus ) hieroglyphica Harris - nom. nud ., invalid; Hagen, 1873:297 (as synonym of Macrotiema
zebratum).
Phryganea ( Leptocerus ) variegata Harris - nom. nud., invalid; Hagen, 1873:297 (as synonym of Macronema zebratum).
Description, — Identical to M. Carolina (see M. Carolina above). May be distinguished from M. Carolina by eyes
small relative to head (Fig. 495), malar space small, vertex high in lateral aspect. Also, M. zebratum is larger species, with
fore-wing length of male 14.04 mm.
Genitalia. Male. (Fig. 490-494). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Identical to M.
Carolina.
Genitalia. Female. (Fig. 496). (Specimen from lie Ste Helene, St Lawrence R., Montreal, Quebec). Identical to M.
Carolina.
Biology. — Wallace (1975) reports on larval feeding, and net structure. Larvae inhabit
rivers from large to very largest, found primarily in rapids. Canadian flight season ranges from
June 1 to September 8.
Distribution. — Other than a curious record from Utah, this species is confined to eastern
North America (Map 77), east of line from North Dakota to Georgia. Recorded northeastward
to Maine. In Canada, abundantly recorded from southern Quebec and Ontario, with one
isolated record from Ignace, far northwestern Ontario.
Map 75. Known distribution of Macrostemum Carolina (Banks) in North America, by state.
Quaest. Ent., 1987, 23 (1)
176
Nimmo
Map 76. Known distribution of Macrostemum transversum (Walker) in North America, by state.
Map 77. Collection localities for Macrostemum zebratum (Hagen) in Canada, with known distribution in North America
by state or province.
Arctopsychidae and Hydropsychidae (Trichoptera)
177
Fig. 482-489. 482, Macrostemum Carolina (Banks): head of male, lateral aspect. 483-489, Macrostemum transversum
(Walker): 483, genital capsule of male, lateral aspect; 484, genital capsule of male, dorsal aspect; 485, aedeagus of male,
dorsal aspect of tip; 486, left clasper of male, posterior aspect; 487, aedeagus of male, lateral aspect; 488, genital segments
of female, lateral aspect; 489, sternite VIII of females, ventral aspect.
Quaest. Ent., 1987,23 (1)
178
Nimmo
Fig. 490-496, Macrostemum zebratum (Hagen): 490, genital capsule of male, lateral aspect; 491, genital capsule of male,
dorsal aspect; 492, left clasper of male, posterior aspect; 493, aedeagus of male, lateral aspect; 494, aedeagus of male,
dorsal aspect of tip; 495, head of adult male, lateral aspect; 496, genital segments of female, lateral aspect.
Arctopsychidae and Hydropsychidae (Trichoptera)
179
ACKNOWLEDGEMENTS
The manuscript of this paper was prepared under a contract awarded by Supply & Services
Canada, on behalf of Agriculture Canada. I am grateful for that.
It is with much appreciation that I acknowledge administration of the above contract by the
University of Alberta, and provision of accomodation by the Department of Entomology.
George E. Ball, then Chairman of the Department, who was technically co-ordinator of the
work, left me to get on with it. His confidence is appreciated.
To my wife Susan, and two daughters, respectively entomological widow and orphans more
often than they cared for, I can only offer my heartfelt thanks for their understanding and
co-operation during protracted absences in the field, or while at the microscope or typewriter.
I wish to extend my warmest thanks to the following institutions and/or individuals for their
various assistances during the course of preparing the manuscript. Their assistance ranged from
loan of requested material, through allowing personal visits to their collections, provision of
comments and information, assistance in tracking down material, to discussion of problems.
They are:
Museum of Comparative Zoology, Harvard University, Boston, Massachusetts; J.D.
Unzicker, Entomology, Illinois Natural History Survey, Urbana, Illinois; G.B. Wiggins,
Entomology, Royal Ontario Museum, Toronto, Ontario; F. Schmid, Biosystematics Research
Institute, Ottawa; Museum of Zoology, University of Michigan, Ann Arbor, Michigan; R.
Neves, U.S. Fish & Wildlife Service, Blacksburg, Virginia; Walter Krivda, The Pas, Manitoba;
Pacific Forest Research Centre, Victoria, British Columbia; O.S. Flint jr., Entomology,
Smithsonian Institution, Washington, D.C.; D. Givens, Entomology, Smithsonian Institution,
Washington, D.C.; P.Schefter, Entomology, Royal Ontario Museum, Toronto, Ontario; D.G.
Denning, Moraga, California; J.B. Wallace, Entomology, University of Georgia, Athens,
Georgia; Academy of Natural Sciences, Entomology, Philadelphia, Pennsylvania; Lymen
Entomology Museum, McDonald College, Ste Anne de Bellevue, Quebec; P. Barnard,
Entomology, British Museum (Natural History), London, England; D.H. Kavanaugh,
Entomology, California Academy of Science, San Francisco, California; G.G.E. Scudder,
Zoology, University of British Columbia, Vancouver, British Columbia; University of Guelph,
Environmental Sciences, Guelph, Ontario; N.H. Anderson, Entomology, Oregon State
University, Corvallis, Oregon; University of Arkansas, Entomology, Fayetteville, Arkansas;
D.A. Etnier, Zoology & Entomology, University of Tennessee, Knoxville, Tennessee;
University of Minnesota, Entomology, Fisheries, & Wildlife, St Paul, Minnesota; S.D. Smith,
Biological Sciences, Central Washington University, Ellensburg, Washington; R.R. Hooper,
Provincial Museum of Natural History, Regina, Saskatchewan; A.E. Gordon, Entomology,
Cornell University, Ithaca, New York; R. Blickle, Entomology, University of New Hampshire,
Durham, New Hampshire; N. Williams, Life Sciences, Scarborough College, West Hill,
Ontario; D. Smith, Biological Sciences, University of Saskatchewan, Saskatoon, Saskatchewan;
B. Wright, Nova Scotia Museum, Halifax, Nova Scotia; P.P. Harper, Sciences Biologiques,
Universite de Montreal, Montreal, Quebec; D.S. Larson, Biology, Memorial University of
Newfoundland, St John’s, Newfoundland; K.W. Stewart, Biological Sciences, North Texas
State University, Denton, Texas.
For a permit to collect I thank the Government of The Yukon.
While on field work during the two summers of the project I was the grateful recipient of the
hospitality of colleagues and friends scattered across Canada and some northern States of the
Quaest. Ent., 1987, 23 (1)
180
Nimmo
Union, and too numerous to mention individually. I thank them all. One only shall I mention
individually. Horace Drury of Fairbanks, Alaska, most kindly took a day to fly me to several
points in the Brooks Range of northern Alaska. Never mind that flying is one of his many
accomplishments and pastimes, it was a marvelously generous gesture on his part for which I
can offer little in return but my gratitude.
Funding for publication of this paper was provided from Natural Sciences & Engineering
Research Council grant #A-1399, held by G.E. Ball, for which assistance I am more than
grateful.
And finally, in the hope that it is unneccesary, I apologise to anyone not acknowledged here,
who should have been.
REFERENCES
Those entries distinguished by an asterisk (*) were used only for accumulation of
distributional data, and are not directly referred to in the text.
Anderson, N.H. 1976. The distribution and biology of the Oregon Trichoptera. Agriculture
Experiment Station, Oregon State University, Technical Bulletin 134:viii+152 pp.
Corvallis, Oregon.
Banks, N. 1899: Descriptions of new North American neuropteroid insects. Transactions of the
American Entomological Society 25:199-218.
Banks, N. 1900. New genera and species of nearctic neuropteroid insects. Transactions of the
American Entomological Society 26:239-259.
Banks, N. 1903. Some new neuropteroid insects. Journal of the New York Entomological
Society 11:236-243.
Banks, N. 1904a. A list of neuropteroid insects, exclusive of Odonata, from the vicinity of
Washington, D.C. Proceedings of the Washington Entomological Society 6:201-217.
Banks, N. 1904b. Neuropteroid insects from New Mexico. Transactions of the American
Entomological Society 32:97-110.
Banks, N. 1905a. Descriptions of new nearctic neuropteroid insects. Transactions of the
American Entomological Society 32:1-20.
Banks, N. 1905b. Descriptions of new species of nearctic neuropteroid insects from the Black
Mountains, N.C. Bulletin of the American Museum of Natural History 21:215-218.
Banks, N. 1907. Descriptions of new Trichoptera. Proceedings of the Washington
Entomological Society 8:117-133.
Banks, N. 1908. Neuropteroid insects - notes and descriptions. Transactions of the American
Entomological Society 34:255-267.
Banks, N. 1909. Two new Caddis Flies. Entomological News 20:342.
Banks, N. 1914. American Trichoptera - notes and descriptions. Canadian Entomologist
46:252-258.
Banks, N. 1918. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology,
Harvard 62:19-22. Cambridge, Masachusetts.
Banks, N. 1936. Notes on some Hydropsychidae. Psyche, Cambridge 43:126-130. Cambridge,
Masachusetts.
Banks, N. 1943. Notes and descriptions of nearctic Trichoptera. Bulletin of the Museum of
Comparative Zoology, Harvard 92:341-369. Cambridge, Masachusetts.
Betten, C. 1934. The Caddis Flies or Trichoptera of New York State. New York State
Arctopsychidae and Hydropsychidae (Trichoptera)
181
Museum Bulletin, Albany 292:1-576.
Betten, C. & M.E. Mosely. 1940. The Francis Walker types of Trichoptera in the British
Museum (Natural History), London, vi + 248 pp.
Blickle, R.L. & W.J. Morse. 1966: The Caddisflies (Trichoptera) of Maine excepting the
family Hydroptilidae. Maine Agricultural Experiment Station, University of Maine,
Bulletin T-24:12 pp.
Brauer, F. 1868. Neue von Herrn Dr G.Semper gesammelte Neuroptera. Verhandlungen der
Zoologisch-Botanischen Gesellschaft in Wien 18:263-268.
*Bueno-Soria, J. & O.S. Flint, jr. 1978. Catalogo sistematico de los Tricopteros de Mexico
(Insecta: Trichoptera), con algunos registros de Norte, Centro y Sudamerica. Anales del
Instituto de Biologia, Universidad de Mexico, Series Zoologia 49:189-218.
Bull. Zool. Nomen. 1962. Opinion 623. Macronema Pictet, 1836 (Insecta, Trichoptera);
validated under the Plenary Powers. Bulletin of Zoological Nomenclature, London
19:80-81.
Bull. Zool. Nomen. 1965. Opinion 758. Diplectrona Westwood, 1839 (Insecta, Trichoptera);
validation and designation of a type-species under the Plenary Powers. Bulletin of
Zoological Nomenclature, London 22:288-289.
Burmeister, H.C.C. 1839. Neuroptera. pp. 757-1050. In: Handbuch der Entomologie, Berlin.
Carpenter, F.M. 1933. Trichoptera from the mountains of North Carolina and Tennessee.
Psyche, Cambridge 40:32-47. Cambridge, Massachusetts.
Curtis, J. 1835. British Entomology vol. 12. London.
Denning, D.G. 1942. Descriptions of new Trichoptera from the United States. Canadian
Entomolgist 74:46-51.
Denning, D.G. 1943. The Hydropsychidae of Minnesota (Trichoptera). Entomologica
Americana 23:101-171.
Denning, D.G. 1947. New species of Trichoptera from the United States. Entomological News
58:249-257.
Denning, D.G. 1948. New species of Trichoptera. Annals of the Entomological Society of
America 41:397-401.
Denning, D.G. 1949. New species of nearctic Caddis Flies. Bulletin of the Brooklyn
Entomological Society 44:37-48.
Denning, D.G. 1952. Descriptions of several new species of Caddis Flies. Canadian
Entomologist 84:17-22.
*Denning, D.G. 1956. Chapter 10 Trichoptera. pp. 237-270. In. Aquatic insects of California
with keys to North American genera and California species, ed. R.L.Usinger. University of
California Press, Berkeley & Los Angeles.
Denning, D.G. 1965. New Hydropsychidae (Trichoptera). Journal of the Kansas
Entomological Society 38:75-84.
*Denning, D.G. & J.W. Beardsley. 1967. The collection of Cheumatopsyche analis in Hawaii.
Proceedings of the Entomological Society of Washington 69:56-57.
*Edwards, S.W. 1973. Texas Caddisflies. Texas Journal of Science 24:491-516.
*Ellis, R.J. 1962. Adult Caddisflies (Trichoptera) from Houghton Creek, Ogemaw County,
Michigan. Occasional Papers of the Museum of Zoology, University of Michigan No. 624,
16 pp.
*Ellis, R.J. 1978. Seasonal abundance and distribution of adult Caddisflies of Sashin Creek,
Baranof Island, southeastern Alaska. Pan-Pacific Entomologist 54:199-206.
Quaest. Ent., 1987,23 (1)
182
Nimmo
*Etnier, D.A. 1965. An annotated list of the Trichoptera of Minnesota, with description of a
new species. Entomological News 74:141-152.
*Etnier, D.A. 1973. Extensions of the known ranges of northern Trichoptera into the southern
Appalachians. Journal of the Entomological Society of Georgia 8:272-274.
Etnier, D.A. & G.A. Schuster. 1979. An annotated list of Trichoptera (Caddisflies) of
Tennessee. Journal of the Tennessee Academy of Science 54:15-22.
Fischer, F.C.J. 1963. Hydropsychidae, Arctopsychidae. Trichopterorum Catalogus 4:vi + 225
pp. Nederlandsche Entomologische Vereeniging, Amsterdam.
Fischer, F.C.J. 1972. Polycentropodidae, Psychomyiidae, Xiphocentronidae, Hydropsychidae,
Arctopsychidae. Trichopterorum Catalogus 13:vii+172 pp. (Supplement to vol. 3 & 4).
Nederlandsche Entomologische Vereeniging, Amsterdam.
Flint, O.S., jr. 1961. The immature stages of the Arctopsychinae occurring in eastern North
America (Trichoptera: Hydropsychidae). Annals of the Entomological Society of America
54:5-11.
Flint, O.S., jr. 1966. Notes on certain nearctic Trichoptera in the Museum of Comparative
Zoology. Proceedings of the United States National Museum 1 1 8(3530):373— 390.
Flint, O.S., jr. & J. Bueno-Soria. 1982. Studies of neotropical Caddisflies, XXXII: The
immature stages of Macronema variipenne Flint & Bueno, with the division of Macronema
by the resurrection of Macrostemum (Trichoptera: Hydropsychidae). Proceedings of the
Biological Society of Washington 95:358-370.
Flint, O.S., jr., J.R. Voshell, jr., & C.R. Parker. 1979. The Hydropsyche scalaris group in
Virginia, with the description of two new species (Trichoptera: Hydropsychidae).
Proceedings of the Biological Society of Washington 92:837-862.
Gordon, A.E. 1974. A synopsis and phylogenetic outline of the nearctic members of
Cheumatopsyche. Proceedings of the Academy of Natural Sciences of Philadelphia
126:117-160.
Gordon, A.E. & S.D. Smith. 1974. A new species of Cheumatopsyche (Trichoptera,
Hydropsychidae) from the northwestern United States. Notulae Naturae of the Academy of
Natural Sciences of Philadelphia No. 450, 3 pp.
Gordon, A.E. & J.B. Wallace. 1975. Distribution of the family Hydropsychidae (Trichoptera)
in the Savannah River basin of North Carolina, South Carolina and Georgia. Hydrobiologia
46:405-423.
Hagen, H. 1861. Synopsis of the Neuroptera of North America, with a list of the South
American species. Miscellaneous Collections of the Smithsonian Institution pp. 249-298,
328-329.
Hagen, H. 1873. Trichoptera. pp. 293-298. In. Report on the Pseudo-Neuroptera and
Neuroptera of North America in the collection of the late T.W. Harris. Proceedings of the
Boston Society of Natural History 15.
*Harris, S.C. & R.B. Carlson. 1977. New records for Trichoptera in North Dakota.
Entomological News 88:217.
*Harris, S.C. & R.B. Carlson. 1978. Distribution of Hydropsychidae (Trichoptera) in sandhill
streams of southeastern North Dakota. Annual Proceedings of the North Dakota Academy
of Science 31:23-27.
*Harris, S.C., P.K. Lago, & R.B. Carlson. 1980. Preliminary survey of the Trichoptera of
North Dakota. Proceedings of the Entomological Society of Washington 82:39-43.
Kimmins, D.E. 1963. On the Trichoptera of Ethiopia. Bulletin of the British Museum (Natural
Arctopsychidae and Hydropsychidae (Trichoptera)
183
History), Entomology 13:1 19-170.
Kimmins, D.E. & D.G. Denning. 1951. The McLachlan types of North American Trichoptera
in the British Museum. Annals of the Entomological Society of America 44:1 1 1-140.
Knowlton, G.F. & F.C. Harmston. 1938. Notes on Utah Plecoptera and Trichoptera.
Entomological News 49:284-286.
Kolenti, F.A. 1859. Genera et species Trichopterorum. Pars altera, pp. 143-296. Moscow.
*Lager, T.M., M.D. Johnson, S.N. Williams, & J.J. McCulloch. 1979. A preliminary report on
the Plecoptera and Trichoptera of northeastern Minnesota. Great Lakes Entomologist
12:109-114.
*Leonard, J.W. & F.A. Leonard. 1949. An annotated list of Michigan Trichoptera. Occasional
Papers of the Museum of Zoology, University of Michigan No. 522, 35 pp. Ann Arbor,
Michigan.
Ling, S.-w. 1938. A few new Caddis Flies in the collection of the California Academy of
Sciences. Pan-Pacific Entomologist 14:59-69.
*Longridge, J.L. & W.L. Hilsenhoff. 1973. Annotated list of Trichoptera (Caddisflies) in
Wisconsin. Wisconsin Academy of Sciences, Arts, & Letters 61:173-183.
Marlier, G. 1962. Genera des Trichopteres de l’Afrique. Annales du Musee royal de Fafrique
Centrale, Tervuren, Belgium No. 109, 263 pp.
Martynov, A.V. 1909. Die Trichopteren des Kaukasus. Zoologisches Jahrbiicher, Systematic,
Okologie und Geographie der Tiere, Jena 27:509-558.
Martynov, A.V. 1925. [Trichoptera]. In. [Practical Entomology]. Leningrad, lxvii + 388 pp. (In
Russian).
*Masteller, E.C. & O.S. Flint, jr. 1979. Light trap and emergence trap records of Caddis Flies
(Trichoptera) of the Lake Erie region of Pennsylvania and adjacent Ohio. Great Lakes
Entomologist 12:165-177.
*McConnochie, K. & F.E. Likens. 1969. Some Trichoptera of the Hubbard Brook
Experimental Forest in central New Hampshire. Canadian Field-Naturalist 83:147-154.
*McElravy, E.P., T.L. Arsuffi, & B. A. Foote. 1977. New records of Caddisflies (Trichoptera)
for Ohio. Proceedings of the Entomological Society of Washington 79:599-604.
McElravy, E.P. & B.A. Foote. 1978. Annotated list of Caddisflies (Trichoptera) occurring
along the upper portion of the west branch of the Mahoning River in northeastern Ohio.
Great Lakes Entomologist 11.143-154.
McLachlan, R. 1866. Descriptions of new or little known genera and species of exotic
Trichoptera, with observations on certain species described by Mr F.Walker. Transactions
of the Royal Entomological Society of London (3) 5:247-275.
McLachlan, R. 1868. Contributions to a knowledge of European Trichoptera. Transactions of
the Royal Entomological Society of London (3) 7:289-308.
McLachlan, R. 1871. On new forms, etc.., of extra-European trichopterous insects. Journal of
the Linnean Society, London 11:98-141.
McLachlan, R. 1878. A monographic revision and synopsis of the Trichoptera of the European
fauna. Part 7:349-428.
Milne, L.J. 1936. Studies in North American Trichoptera. 3:56-128. Cambridge,
Massachusetts.
Milne, L.J. & M.J. Milne. 1938. The Arctopsychinae of continental America north of Mexico
(Trichoptera). Bulletin of the Brooklyn Entomological Society 33:97-1 10.
Milne, M.J. 1940. Immature North American Trichoptera. Psyche, Cambridge 46:9-19.
Quaest. Ent., 1987,23 (1)
184
Nimmo
Cambridge, Massachusetts.
*Morse, J.C., J.W. Chapin, D.D. Herlong, & R.S. Harvey. 1980. Aquatic insects of upper
Three Runs Creek, Savannah River Plant, South Carolina. Part I: Orders other than
Diptera. Journal of the Georgia Entomological Society 15:73-101.
*Morse, W.J. & R.L. Blickle. 1953. A check list of the Trichoptera (Caddis Flies) of New
Hampshire. Entomological News 64:68-73.
Navas, L. 1918. Tricopteros nuevos de Espana (Quinta serie). Broteria 16:7-20.
Navas, L. 1933. Insecta Orientalia, 12 ser. Memorie-atti della Pontificia Accademia Sciencia
nuovi Lincei, Roma (2) 17:75-108.
Neave, F. 1929. Reports of the Jasper Park lakes investigations 1925-26. IV. Aquatic insects.
Contributions to Canadian Biology & Fisheries, N.S. 4:187-195.
Neves, R.J. 1979. A checklist of Caddisflies (Trichoptera) from Massachusetts. Entomological
News 90:167-175.
*Newell, R.L. & G.W. Minshall. 1977. An annotated list of the aquatic insects of southeastern
Idaho, Part II: Trichoptera. Great Basin Naturalist 37:253-257.
Newell, R.L. & D.S. Potter. 1973. Distribution of some Montana Caddisflies (Trichoptera).
Proceedings of the Montana Academy of Sciences 33:12-21.
Nimmo, A.P. 1966a. A list of Trichoptera taken at Montreal and Chambly, Quebec, with
descriptions of three new species. Canadian Entomologist 98:688-693.
Nimmo, A.P. 1966b. The arrival pattern of Trichoptera at artificial light near Montreal,
Quebec. Quaestiones Entomologicae 2:217-242.
Nimmo, A.P. 1971. The adult Rhyacophilidae and Limnephilidae (Trichoptera) of Alberta and
eastern British Columbia and their post-glacial origin. Quaestiones Entomologicae 7:3-234.
Nimmo, A.P. 1981. Francis Walker types of, and new synonymies for, North American
Hydropsyche species (Trichoptera, Hydropsychidae). Psyche, Cambridge 88:259-263.
Cambridge, Massachusetts.
*Nimmo, A.P. & G.G.E. Scudder. 1978. An annotated checklist of the Trichoptera (Insecta)
of British Columbia. Syesis 11:117-133.
*Nimmo, A.P. & G.G.E. Scudder. 1983. Supplement to an annotated checklist of the
Trichoptera (Insecta) of British Columbia. Syesis 16:71-83.
*Parker, C.R. & J.R. Voshell. 1981. A preliminary checklist of the Caddisflies (Trichoptera)
of Virginia. Journal of the Georgia Entomological Society 16:1-7.
Pictet, F.J. 1834. Recherches pour servir a l’histoire et a l’anatomie des Phryganides. 235 pp.
Geneva.
Pictet, F.J. 1836. Description de quelques nouvelles especes de Neuropteres du Musee de
Geneve. Memoires de la Societe de physique et d’histoire naturelle de Geneve 7:396-403.
*Resh, V.H. 1975. A distributional study of the Caddisflies of Kentucky. Transactions of the
Kentucky Academy of Science 36:6-16.
Ross, H.H. 1938a. Descriptions of new North American Trichoptera. Proceedings of the
Entomological Society of Washington 40:1 17-124.
Ross, H.H. 1938b. Descriptions of nearctic Caddis Flies. Bulletin of the Illinois Natural
History Survey, Urbana 21:iv+ 183 pp.
Ross, H.H. 1938c. Lectoptypes of North American Caddis Flies in the Museum of
Comparative Zoology. Psyche, Cambridge 45:1-61. Cambridge, Massachusetts.
Ross, H.H. 1939. New species of Trichoptera from the Appalachian region. Proceedings of the
Entomological Society of America 41:65-72.
Arctopsychidae and Hydropsychidae (Trichoptera)
185
Ross, H.H. 1941. Descriptions and records of North American Trichoptera. Transactions of the
American Entomological Society 67:35-126, 13 plates.
Ross, H.H. 1944. The Caddis Flies, or Trichoptera, of Illinois. Bulletin of the Illinois Natural
History Survey, Urbana 23, 326 pp.
Ross, H.H. 1947. Descriptions and records of North American Trichoptera, with synoptic
notes. Transactions of the American Entomological Society 73:125-168, 8 plates.
Ross, H.H. 1956. Evolution and classification of the mountain Caddisflies. vii + 213 pp.
University of Illinois Press, Urbana.
Ross, H.H. & G.J. Spencer. 1952. A preliminary list of the Trichoptera of British Columbia.
Proceedings of the Entomological Society of British Columbia 48:43-51.
Ross, H.H. & J.D. Unzicker. 1977. The relationships of the genera of American
Hydropsychinae as indicated by phallic structures (Trichoptera, Hydropsychidae). Journal
of the Georgia Entomological Society 12:298-312.
Roy, D. & P.P. Harper. 1979. Liste preliminaire des Trichopteres (Insectes) du Quebec.
Annales de la Societe Entomologique du Quebec 24:148-172.
Schmid, F. 1968. La famille des Arctopsychides. Memoires de la Societe Entomologique du
Quebec No. 1, 84 pp.
Schmid, F. 1979. On some new trends in Trichopterology. Bulletin of the Entomological
Society of Canada 1 1:48-57.
Schmid, F. 1980. Genera des Trichopteres du Canada et des Etats adjacentes. Les Insectes et
Arachnides du Canada, part 7, 296 pp. Agriculture Canada, Ottawa.
Schefter, P.W. & J.D. Unzicker. 1984. A review of the Hydropsyche morosa-bifida complex
in North America (Trichoptera: Hydropsychidae). pp. 331-335. In: Proceedings of the
Fourth International Symposium on Trichoptera. ed. J.C. Morse. Junk, The Hague.
Schefter, P.W. & G.B. Wiggins. 1986. A systematic study of the nearctic larvae of the
Hydropsyche morosa group (Trichoptera: Hydropsychidae). Life Sciences Miscellaneous
Publications, Royal Ontario Museum, Toronto. 94 pp.
Schefter, P.W., G.B. Wiggins, & J.D. Unzicker. 1986. A proposal for assignment of
Ceratopsyche as a subgenus of Hydropsyche , with new synonyms and a new species
(Trichoptera: Hydropsychidae). Journal of the North American Benthological Society
5:67-84.
Schuster, G.A. & D.A. Etnier. 1978. A manual for the identification of the larvae of the
Caddisfly genera Hydropsyche Pictet and Symphitopsyche Ulmer in eastern and central
North America (Trichoptera: Hydropsychidae). Environmental Monitoring & Support
Laboratory, Cincinnati EPA-600/4-78-060, xii+129 pp.
Simmons, P., D.F. Barnes, C.K. Fischer, & G.F. Kaloostian. 1942. Caddisfly larvae fouling a
water tunnel. Journal of Economic Entomology 35:77-79.
Smith, D. 1979. The larval stage of Hydropsyche separata Banks (Trichoptera:
Hydropsychidae). Pan-Pacific Entomologist 55:10-20.
Smith, S.D. 1968. The Arctopsychinae of Idaho (Trichoptera: Hydropsychidae). Pan-Pacific
Entomologist 44: 1 02-1 1 2.
Smith, S.D. 1969. Two new species of Idaho Trichoptera with distributional and taxonomic
notes on other species. Journal of the Kansas Entomological Society 42:46-53.
Snodgrass, R.E. 1957. A revised interpretation of the external reproductive organs of male
insects. Smithsonian Miscellaneous Collections 1 35(6): 1 1 1 +60 pp.
Stephens, J.F. 1836. Illustrations of British Entomology. 6:146-234.
Quaest. Ent., 1987,23 (1)
186
Nimmo
Tarter, D.C. & P.L. Hill. 1979. Caddisflies (Trichoptera) of the Cranberry Glades in West
Virginia. Entomological News 90:205-206.
Ulmer, G. 1905a. Zur Kenntnis Aussereuropaischer Trichopteren. Stettiner Entomologischer
Zeitung 66:1-1 19.
Ulmer, G. 1905b. Trichopteren aus Java. Mitteilungen aus dem Naturhistorischen Museum in
Hamburg 22:89-200.
Ulmer, G. 1905c. Neue und wenig bekannte Trichopteren der Museen zii Brussel und Paris.
Annales de la Societe Entomologique de Belgique 49:17-41.
Ulmer, G. 1907. Neue Trichopteren. Notes from the Leyden Museum 29:1-53.
Ulmer, G. 1951. Kocherfliegen (Trichoptera) von den Sunda-Inseln (Teil I). Archiv fur
Hydrobiologie, Supplement 19, 528 pp.
Unzicker, J.D., L. Aggus, & L.O. Warren. A preliminary list of the Arkansas Trichoptera.
Journal of the Georgia Entomological Society 5:167-174.
Vorhies, C.T. 1909. Studies on the Trichoptera of Wisconsin. Transactions of the Wisconsin
Academy of Sciences, Arts, and Letters 16:647-738.
Walker, F. 1852. Catalogue of the specimens of neuropterous insects in the collections of the
British Museum. Part 1. 192 pp. British Museum (Natural History), London.
Wallace, J.B. 1975a. The larval retreat and food of Arctopsyche; with phylogenetic notes on
feeding adaptations in Hydropsychidae larvae (Trichoptera). Annals of the Entomological
Society of America 68:167-173.
Wallace, J.B. 1975b. Food partitioning in net-spinning Trichoptera larvae: Hydropsyche
venularis, Cheumatopsyche etrona, and Macronema zebratum (Hydropsychidae). Annals
of the Entomological Society of America 68:463-472.
Wallace, J.B. & F.F. Sherberger. 1974. The larval retreat and feeding net of Macronema
Carolina Banks (Trichoptera: Hydropsychidae). Hydrobiologia 45:177-184.
Wallace, J.B. & F.F. Sherberger. 1975. The larval dwelling and feeding structure of
Macronema transversum (Walker)(Trichoptera: Hydropsychidae). Animal Behaviour
23:592-596.
Wallengren, H.D.J. 1891. Skandinaviens Neuroptera. Andra afdelningen. Svenska Akademien
Handlingar 24(10): 173 pp.
Westwood, J.O. 1840. An introduction to the modern classification of insects. Generic Synopsis
2:49-51. London.
Wiggins, G.B. 1977. Larvae of the North American Caddisfly genera (Trichoptera). xi + 401
pp. University of Toronto Press, Toronto.
Arctopsychidae and Hydropsychidae (Trichoptera)
187
INDEX TO NAMES OF TAXA
(Synonyms in italics)
FAMILY GROUP TAXA
Arctopsychidae, 2, 4
Arctopsychinae, 4
Diplectroninae, 22, 167
Hydropsychidae, 2, 4, 22, 167
Hydropsychinae, 22-23, 167
Macronematinae, 22, 164, 167, 173
Oestropsinae, 173
Phryganeidae, 22
GENERA AND SUBGENERA
Aphelocheira Stephens, 170
Aphropsyche Ross, 167
Arctopsyche McLachlan, 4-5, 14
Cheumatopsyche Wallengren, 2, 23, 164
Diplectrona Westwood, 170
Hydropsyche Pictet, 2, 23, 76, 164
Hy dropsy chodes Ulmer, 23
Macronema Pictet, 173
Macrostemum Kolenati, 22, 173
Parapsyche Betten, 4, 14
Potamyia Banks, 164
Symphitopsyche Ulmer, 76
Ulmeria Navas, 23
SPECIES AND SUBSPECIES
aenigma Schefter, Wiggins, & Unizicker,
Hydropsyche, 79, 148
aerata Ross, Hydropsyche, 77, 82-83
alhedra (Ross), Symphitopsyche, 134
alhedra Ross, Hydropsyche, 4, 80, 82, 134,
156
almota Ross, Parapsyche, 14-15
alternans (Walker), Hydropsyche, 76, 79,
82, 139, 148
alternans Walker, Philopotamus, 148
alvata Denning, Hydropsyche, 78, 83
amblis Ross, Hydropsyche, 79, 82, 126
analis (Banks), Hydropsy chodes, 49
analis Banks, Cheumatopsyche, 49
analis Banks, Hydropsyche, 48-49
aphanta Ross, Cheumatopsyche, 25-26
apicalis (Banks), Parapsyche, 14-15
apicalis Banks, Arctopsyche, 1 5
aprilis Ross, Aphropsyche, 167
arinale Ross, Hydropsyche, 78, 81, 84
betteni Ross, Hydropsyche, 76, 81, 1 18
bidens Ross, Hydropsyche, 77, 80, 84
bifida (Banks), Symphitopsyche, 134
bifida Banks, Hydropsyche, 4, 80, 82, 134
bronta (Ross), Symphitopsyche, 138
bronta Ross, Hydropsyche, 79, 82, 138
burksi Ross, Cheumatopsyche, 25-26, 65
californica Banks, Hydropsyche, 78, 80, 84
campyla Ross, Cheumatopsyche, 26-27,
38
Carolina (Banks), Macrostemum, 173-175
Carolina Banks, Macronema, 174
centra Ross, Hydropsyche, 79, 82, 148
cheilonis (Ross), Symphitopsyche, 138
cheilonis Ross, Hydropsyche, 79, 82, 138
chlorotica Hagen, Hydropsyche, 139
cockerelli Banks, Hydropsyche, 4, 79, 82,
149
codona Betten, Hydropsyche, 148
confusa (Walker), Hydropsyche, 76, 81,
118
confusus Walker, Philopotamus, 118
corbetti Nimmo, Hydropsyche, 118
cornuta Ross, Hydropsyche, 87
cuanis Ross, Hydropsyche, 77, 80, 119
depravata Hagen, Hydropsyche, 77, 80,
119
dicantha Ross, Hydropsyche, 77, 81, 85
doringa Milne, Aphropsyche, 22, 167
ela Denning, Cheumatopsyche, 25, 27, 39
elsis Milne, Parapsyche, 14, 16
enonis Ross, Cheumatopsyche, 25-26, 66
flava (Hagen), Hydropsyche, 164
flava (Hagen), Potamyia, 23, 164
flavum Hagen, Macronema, 164
frisoni Ross, Hydropsyche, 78, 81, 85
geolca Denning, Cheumatopsyche, 66
gracilis (Banks), Cheumatopsyche, 25-26,
60
gracilis (Banks), Hydropsychodes, 60
Quaest. Ent., 1987,23 (1)
188
Nimmo
gracilis Banks, Hydropsyche, 60
grandis (Banks), Arctopsyche, 5
guttata Pictet, Hydropsyche, 118
h. harwoodi Denning, Cheumatopsyche,
25, 27, 54
hageni Banks, Hydropsyche, 77, 81, 86
halima Denning, Cheumatopsyche, 25-26,
65
harwoodi Denning, Cheumatopsyche, 54
helma Ross, Cheumatopsyche, 25, 27, 54
hieroglyphica Harris, Phryganea, 175
incommoda not Hagen, Hydropsyche, 89,
118
inermis Banks, Arctopsyche , 5
irrorata Banks, Arctopsyche, 5-6
jewetti Denning, Hydropsyche, 4, 79, 149
kansasensis Banks, Hydropsyche, 1 64
ladogensis (Kolenati), Arctopsyche, 5, 7
ladogensis Kolenati, Aphelocheira, 7
lasia Ross, Cheumatopsyche, 24, 27, 32
leonardi Ross, Hydropsyche, 78, 86
logani Gordon, Cheumatopsyche, 26, 48
maculicornis Walker, Hydropsyche, 127
mickeli Denning, Cheumatopsyche, 25-26,
38
minuscula (Banks), Cheumatopsyche, 24,
26- 27
minuscula (Banks), Hy dropsy chodes, 27
minuscula Banks, Hydropsyche, 27
modesta Banks, Diplectrona, 22, 170
mollala Ross, Cheumatopsyche, 24, 27, 65
montrealensis Nimmo, Cheumatopsyche,
27- 28
morosa (Hagen), Symphitopsyche, 139
morosa Hagen, Hydropsyche, 4, 48-49,
60,78,82, 88, 139
novamexicana Banks, Hydropsyche, 87
occidentalis Banks, Hydropsyche, 77, 80,
87
oregonensis Ling, Arctopsyche, 1 4
orris Ross, Hydropsyche, 78, 81, 87
oslari Ross, Hydropsyche, 79, 82, 160
oxa Ross, Cheumatopsyche, 24, 27, 64
partita Banks, Hydropsyche, 160
pasella Ross, Cheumatopsyche, 25, 27, 38
pettiti (Banks), Cheumotopsyche, 25-26,
48
pettiti (Banks), Hydropsy chodes, 48
pettiti Banks, Hydropsyche, 48
phalerata Hagen, Hydropsyche, 78, 80, 88
phryganoides Banks, Arctopsyche, 5
piatrix (Ross), Symphitopsyche, 126
piatrix Ross, Hydropsyche, 79, 82, 126
pinaca Ross, Cheumatopsyche, 24, 26, 32
placoda Ross, Hydropsyche, 77, 81, 88
polygrammaticum (McLachlan),
Macronema, 174
polygrammatum McLachlan,
Macronema, 174
racona Denning, Hydropsyche, 1 34
recurvata (Walker), Hydropsyche, 148
recurvata (Walker), Symphitopsyche, 148
riola (Denning), Symphitopsyche, 156
riola Denning, Hydropsyche, 4, 79, 83,
134, 156
rossi Flint, Voshell, & Parker,
Hydropsyche, 78, 80, 89
scalaris Hagen, Hydropsyche, 77, 81, 84,
86, 89
seperata Banks, Hydropsyche, 1 18
simulans Ross, Hydropsyche, 78, 81, 90
slossonae (Banks), Symphitopsyche, 139
slossonae Banks, Hydropsyche, 78, 82,
139, 148
smithi Gordon, Cheumatopsyche, 25, 27,
48
sordida (Hagen), Cheumatopsyche, 24, 26,
28
sordida (Hagen), Hydropsy chodes, 28
sordida Hagen, Hydropsyche, 28
sparna (Ross), Symphitopsyche, 156
sparna Ross, Hydropsyche, 79, 83, 156
speciosa (Banks), Cheumatopsyche, 24,
26,32
speciosa (Banks), Hydropsy chodes, 32
speciosa Banks, Hydropsyche, 32
tana Ross, Hydropsyche, 78, 82, 140
transversa Walker, Hydropsyche, 174
transversum (Walker), Macronema, 174
transversufn (Walker), Macrostemum,
173-174
valanis Ross, Hydropsyche, 77, 81, 90
Arctopsychidae and Hydropsychidae (Trichoptera)
189
vannotei Gordon, Cheumatopsyche, 24, 60
variegata Harris, Phryganea, 175
ventura (Ross), Symphitopsyche, 160
ventura Ross, Hydropsyche, 79, 82, 160
venularis Banks, Hydropsyche, 77, 81, 90
vexa (Ross), Symphitopsyche , 127
vexa Ross, Hydropsyche, 79, 83, 127
wabasha Denning, Cheumatopsyche, 24,
46
walkeri Betten & Mosely, Hydropsyche,
79-80, 127
walkeri Betten & Mosely,
Symphitopsyche , 127
wrighti Ross, Cheumatopsyche, 25, 54
zebratum (Hagen), Macrostemum,
173-175
Quaest. Ent., 1987,23 (1)
Book Reviews
191
BOOK REVIEW
Campos-Ortega, J. A. and V. Hartenstein. 1985. The Embryonic Development of Drosophila
melanogaster. Springer-Verlag. Berlin, Heidelberg, New York, Tokyo. 227 pp., 85 figs., 2
tables, subject index. $120.00Can.
Flies of the species Drosophila melanogaster are arguably the best understood of all higher
animals because of their enormous popularity with geneticists since 1909 when T. H. Morgan
first started culturing them in his “fly room” at Columbia University. To-day they are the
organisms of choice for legions of highly ingenious and pecunious developmental biologists
interested in understanding how genes control cell determination and differentiation in higher
animals.
In the last decade, a flood of papers have appeared on the genetic basis of cell determination
and pattern formation in Drosophila embryos-most recently on the mode-of-action of genes
controlling segmentation of the germ band (segmentaion genes) and longitudinal specification
of segment identity (homoeotic genes). Most of these papers have been based on use of genetic
dissection methods and have involved analysis of mutant embryonic and larval phenotypes
(more recently, use of recombinant DNA and immunocytochemical techniques has
predominated). However, in spite of this explosion in experimental results, the standard
description of normal embryogenesis against which they are compared has remained
Sonnenblick’s and Poulson’s superb but now out-of-date chapters in Demerec’s (1950) Biology
of Drosophila (updated but with little added by Fullilove and Jacobson [1978] and Bownes
[1982]). Campos-Ortega and Hartenstein’s new book supplants these classic accounts with
style and depth (it is dedicated to Donald F. Poulson).
Eggs of D. melanogaster average 0.42 x 0.15 mm and require about one day to develop from
fertilization to hatching at 25°C. This development is described in great detail in the eight
chapters of this book. Chapter 1 (6 pp.) presents a brief, illustrated (12 drawings), summary of
embryogenesis in this species and chapter 2 (75 pp.) a system for staging it. The authors divide
embryogenesis into 17 stages ranging in length from 10 min (each of stages 6 and 7) to 11.7 hr
(stage 17) and summarize the events occurring in each stage. These events are illustrated with
71 photomicrographs of living whole mounts and 123 of fixed embryos embedded in plastic and
sectioned longitudinally or horizontally.
Origin and differentiation of most organ systems are considered in Chapter 3 (79 pp.) -
including the gut, Malpighian tubules, salivary glands, gonads, somatic and visceral
musculature, dorsal vessel, fat body, epidermis, central (CNS), peripheral (PNS) and
stomatogastric (SNS) nervous systems and the tracheal system. The chapter is illustrated with
42 drawings, 56 photomicrographs and seven transmission electron micrographs - the most
impressive of the drawings being Fig. 3.14 (a lateral view of a first instar larva with every
external cuticular structure illustrated), Figs. 3.23-3.25 (detailed, 3-dimensional
reconstructions of all external sensilla and their projections to specific ganglia of the CNS in
both ventral and lateral aspect), and Fig. 3.30 (six 3-dimensional reconstructions illustrating
neurogenesis of the CNS). (The origin and differentiation of the CNS is remarkably similar to
that recently described in great detail in various grasshopper embryos by Goodman, Bate and
their colleagues).
The pattern of embryonic cell division is summarized in Chapter 4 (7 pp., 6 drawings) and is
shown to occur during two periods. The first takes place prior to cell formation at blastoderm
and involves 1 3 rounds of mitosis. The first seven zygotic divisions are synchronous and yield a
Quaest. Ent., 1987, 23 (1)
192
Book Reviews
syncytium of 128 nuclei distributed as an ellipsoid within the yolk. With the next 3 mitoses
(8-10), the resulting nuclei approach the peripheral periplasm to form the syncytial blastoderm
with 200 nuclei remaining behind in the yolk as vitellophags and 17-18 entering the posterior
pole plasm to form pole cells (germ line cells). Three further (11-13) and now parasynchronous
mitoses increase the number of syncytial blastoderm nuclei to about 5000 at which time cell
membranes form between adjacent nuclei. During the second period of mitosis, from
blastoderm formation to hatching, most cells divide only two or three more times - exceptions
being the teloblastic nerve mother cells (neuroblasts) of the CNS, sensillar stem cells and pole
cells.
Chapter 5 (13 pp., 16 drawings) considers the morphogenetic movements involved in
changing the simple cellular monolayer of the blastoderm into the complex, 3-dimensional
structure of the young embryo. These movements occur either by growth and infolding of
certain regions of the blastoderm or by movement of cells or groups of cells into the interior and
can involve cell proliferation, changes in cell shape and size, and shifting of individual cells in
relation to each other. Gastrulation, germ band elongation and (later) shortening, head
involution and dorsal closure are among the more important of these movements in Drosophila
embryogenesis. With gastrulation, 1250 cells invaginate from the ventral midline of the embryo
to form mesoderm and endoderm leaving 3750 cells about the yolk to form the rest of the larval
body (ectoderm). The germ band elongates 220% around the posterior end of the egg during
germ ban elongation and later, during its shortening, each thoracic and abdominal segment
shortens and broadens - a shape change attained through shape changes of their individual
cells.
Cephalogenesis (Chapter 6-13 pp., 9 drawings, 11 photomicrographs) includes head
involution, a phenomenon unique to higher Diptera (Cyclorrhapha). At stage 11, the
developing head is similar to, though less complex than, heads of other insect embryos at
comparable stages of development and contains a large procephalon and three gnathal
segments each bearing a pair of short, blunt, appendages. Portions of all these segments shift
cephalad during the remainder of embryogenesis and invaginate into the front of the embryo to
form the atrium of the foregut in which is later secreted the cuticle of the larval
cephalopharyngeal skeleton. The segmental source of the various parts of this skeleton are
indicated (the mouth hooks, for example, derive from the maxillary appendages).
Certain aspects of segmentation are considered in more detail in Chapter 7 (12 pp., 8
drawings, 2 tables) including formation of segmental borders, epidermal sensory organs, muscle
innervation, and homologies between the muscles and sense organs of each segment.
The final chapter (8- 8 pp., 2 drawings) presents a new fate map of the Drosophila
blastoderm. This was derived by use of three methods: 1 . The enzyme horse radish peroxidase
(HRP) was injected into donor eggs before cellularization. This tracer spreads freely
throughout each embryo and becomes incorporated into all its cells during cellularization. Just
after cellularization, labelled cells are taken from these donors by means of a micropipette and
transplanted singly into unlabelled host embryos. The progeny of each transplanted cell is then
traced during subsequent development of the host embryo. 2. Progressively older embryos were
removed from their choria and vitelline membranes and stained with fuchsin (every nucleus can
be counted and drawn in such embryos) 3. Planimetric reconstructions of complete series of
sections of embryos of similar age were prepared by “rolling off’ folded and invaginated tissues.
Use of these methods enabled the authors to obtain reliable quantitative data for each
embryonic stage. Fig. IB illustrates this fate map for one side of the blastoderm and indicates
Book Reviews
193
the number of blastoderm cells contributing to each larval organ derived from that side (for
example, 150 blastoderm cells give rise to one half the proctodeum). The rest of this chapter
considers cell number in each organ and the number of blastoderm cells giving rise to it.
Production quality of the book is outstanding, for paper, printing and binding, and the
halftones of the photomicrographs are of uniformly high standard. This last was because all
sections were cut from eggs dechorionated and fixed according to Zalokar and Erk’s (1977)
phase-partition method (which causes essentially no disruption of egg contents) and embedded
in plastic. There are a few typographical errors and some awkward grammar but not much,
considering that the senior author is Spanish and the junior author German. My principal
criticisms are its outrageous price, the total absence of scanning electron micrographs
(available from the work of Turner and Mahowald [1976-1979] and cited in the book), and the
almost total absence of reference to embryos of other insects (only Goodman et aV s work on
grasshopper neurogenesis, Anderson’s (1962, 1972) synopses, and Wheeler’s (1891, 1983) and
Weisman’s (1863) classical descriptive studies are mentioned). Their discussion of head
segmentation in chapter 6, for example, would have been vastly improved had they referred to
Rempel’s (1975 Quaest. Ent. 11: 7-25) critical review of this topic. (This subject has recently
been considered at great length for this species by Jurgens et al., Rous’s Archives of
Developmental Biology 195: 359-377. [1986]). There are only 1 12 references.
However, this book was written for Drosophila workers-not for comparative embryologists
or entomologists-and as such will be indispensible to all individuals investigating the embryos
of these insects (most recent papers cite it). There are no descriptive studies of the embryos of
other insects that even approach this study in detail, and it is now the standard of excellence
against which all future such investigations will be compared.
B. S. Heming
Dept, of Entomology
Univ. of Alberta
Quaest. Ent., 1987,23 (1)
194
Book Reviews
BOOK REVIEW: INSECTS FOR ALL SEASONS
TAUBER, M.J., C.A. TAUBER AND S. MASAKI. 1986. Seasonal Adaptations of Insects.
Oxford University Press, New York. 411 pages, 16 text figures, bibliography, author index,
species index, subject index. Price $39.95 USA.
When I consider writing up an investigation somewhat removed from the area of my most
recent endeavor, I begin by reading a book that reviews the general topic. I have two main
objectives: (1) to increase confidence about my grasp of the literature, and (2) to help forge a
more synthetic entomological backbone for the paper that I would like to write. With these
goals in mind, I purchased Seasonal Adaptations of Insects off the shelf in Blackwell’s
Bookstore, despite its scandalous British price. The book did not offer immediate help with
either of my objectives, but instead laid bare the breadth of my ignorance about an important
and interesting field of entomological research. The treatment is well organized and solidly
documented with meaty discussions of work relevant to all major groups of insects. It is an
excellent synthetic synopsis of a complex literature that has grown by leaps and bounds during
the past 15 years. The authors lead the reader to delight in, rather than despair over, the
incredible variety of phenological adaptations in insects, and suggest the outlines for a more
holistic approach to empirical studies of insect life cycles. After reading the book, I share the
authors’ enthusiastic optimism about the future of such work. I also think that I got my
money’s worth!
The first five chapters provide a basis for understanding seasonal adaptations in the context
of the “diapause syndrome”. Chapter 1 provides a brief but useful historical summary about
work on insect life cycles and clearly focuses the reader on the authors’ two main objectives: (1)
to describe the variety of seasonal adaptations employed by insects, and (2) to provide a
synthetic basis for ecological and evolutionary studies of diapause. Chapter 2 provides a
classification of the ways that insects respond to seasonal change and places diapause within a
broader adaptive context. Chapter 3 describes the course of diapause with reference to the
endless variations known among insects.
The fourth chapter is the keystone of the book, laying out the authors’ concept of diapause.
Their “diapause syndrome” is a more global and dynamic concept than those that are generally
employed, and I found it more satisfying. For Tauber, Tauber and Masaki, diapause is not
primarily concerned with surviving adverse conditions, but instead, with effecting broadly
adaptive synchrony between insect population dynamics and seasonal variation in the
environment, both biotic and abiotic. In their view then, the anticipatory nature of diapause is
paramount, and seasonal synchronization is linked explicitly to neurohormonal bases. This
broad concept removes any possible confusion between dormancy and diapause, isolates the
evolution and development of cold hardiness as a parallel problem, and incorporates dormancy,
migration and various polymorphisms in the adaptive context of the entire life cycle.
Chapter 5 is a brief but effective summary of how environment regulates seasonal cycles.
Evidence for the central position of photoperiod in many species is crisply summarized, the
primary inductive and secondary modifying effects of temperature are discussed, and these are
considered in relation to other environmental characteristics known to influence the
manifestation of diapause. The basis for constructing photoperiod response curves and for the
concept of critical photoperiod are critically reviewed. Although the authors show that
substantial progress has been made by assessing responses to static photoperiods, they argue
that we must adopt a more dynamic approach in order to get crucial information about
Book Reviews
195
diapause intensity, and in order to understand the effects of changing daylengths experienced
by natural populations. The chapter is an excellent guide to the important primary literature,
and work about groups that I know well was efficiently and accurately abstracted.
In Chapter 6, the authors explicitly tackle the vast diversity of insect life cycles by extending
the general principles summarized in the first part of the book to insects of highly specialized
adaptive zones or extreme habitats. The section about parasitoids underscores the diverse range
of ways that the insect-parasitoid relationship can affect the life cycles of either partner. We
also learn about how the unique aspects of social life are tied up in regulation of phenology
among the social Hymenoptera. Unfortunately, there is only a passing descriptive reference to
one paper about termites, and uncharacteristically, the authors do not mention whether more
extensive information is available. The treatment of diapause in tropical insects is fascinating.
Two important points are well established: (1) numerous tropical species show diapause, and
(2) diapause is clearly adaptive in at least some tropical populations and not simply a result of
introgression from more temperate populations. The chapter concludes with treatments of
phenology in desert and arctic insects. Diapause is an important component of life in such
environments and the diapause syndrome of arctic and desert species has been molded in weird
and wonderous ways. However, as the authors point out, we have much to learn about the
details of insect life cycles in both environments.
Chapters 7-9 consider the diapause syndrome as an evolutionary phenomenon. In Chapter
7, the authors partition genetic variation for seasonal adaptation into continuous variation in
quantitative traits and disjunct variation in qualitative traits. They offer solid arguments about
the fitness implications of quantitative traits such as intrapopulation variation of critical
photoperiod, but also show that some variation has little direct relation to fitness and may have
little adaptive value. They provide general guidelines for working out the genetics of diapause
and document them with reviews of information now available. It appears that most
quantitative traits are under polygenic control with genetic diversity maintained by variable
selection pressures and intrapopulation genetic exchange. The relative importance of particular
diversity maintaining processes are still a matter of dispute and may likely vary among taxa.
Although Mendelian inheritance has been shown in only a few insect taxa where seasonal traits
vary discretely, the authors believe that this will become more common as more detailed studies
are made. The authors also discuss how genetic variation in polygenic traits can lead to
polymorphisms with the incorporation of a threshold mechanism allowing continuous variation
to be expressed discretely. This chapter provides a good starting place for anyone contemplating
an evolutionary study of insect life cycles.
Chapter 8 provides an evolutionary scenario for diapause that links the diverse seasonal
adaptations summarized in the earlier chapters and explains their control by a host of seasonal
token stimuli. This general model allows for the obvious multiple evolution of diapause in the
context of three steps. First, the authors assume the evolution of a time measuring system of
some sort. The authors review the several models that have been published and show that there
is still more to learn before we can make an informed choice among them. Second, their
scenario requires the evolution of environmentally controlled neuroendocrinological responses
in an adaptive context. The third stage involves the coupling of the neuroendocrine responses to
seasonal environmental stimuli. The authors go on to show how their scheme provides a useful
way of thinking about the evidence for evolution of diapause that has accumulated from studies
employing artificial selection, work about colonizing species, and a host of comparative studies.
In an illuminating critical discussion, they outline the benefits and limitations of the sorts of
Quaest. Ent., 1987, 23 (1)
196
Book Reviews
broad comparative studies that have been made, and finish with a plea for detailed comparisons
of environmental and physiological control mechanisms in groups of related species with well
established phylogenies. This approach can reveal the polarity of diapause evolution in specific
lineages. A simple, but yet instructive example is provided by discussion of the Taubers’ own
work with green lacewings in the Chrysopa carnea species complex.
The general evolution of life histories is viewed in the context of seasonal adaption in
Chapter 9. Up to now most studies of insect life history have considered isolated features or
suites of characters that are obviously co-adapted. Their diapause syndrome concept
underscores that adaptation to seasonality influences life cycle timing and many other features
of insect life history, and the authors rightly urge us to take a broader view of life history
evolution in insects. Two important proximate objectives are readily accessible. First, we need
to understand the limits of genetic variation for life history traits and relate the pattern of such
variation to environmental correlates such as are envisioned by Southwood’s (1977, J. Anim.
Ecol. 46:337-365) notion of the “habitat templet”. Genetic variation for life history has obvious
geographical correlates and these are outlined in the context of a few well reviewed studies.
Geographical variation in life cycles may also be related to speciation in many taxa. Second, we
can search for pattern in how life history traits evolve together in the context of seasonality with
particular attention to how fitness tradeoffs are managed by natural selection. Life cycles are
indeed adaptive, but they are also constrained by their underlying genetic systems. We need to
know more about how variation for life history traits is maintained before we can make
progress toward this objective The final chapter effectively discharges the entomologist’s
standard social obligation by showing how understanding of life cycles is crucial to pest
management. It is largely a summary of current applications and a discussion of several
approaches that are under development now. There is not much new information or conceptual
synthesis useful to pest managers. However, for those desiring a summary of the anthorpo-
centric justification for studying insect life cycles, the chapter will be useful. It is ideal reading
for intoductory students in pest management and I will use it in my own teaching.
The book may be criticized because it largely ignores developments in the trendy literature
of life history evolution. However, such criticism would miss the point. The authors and most
entomologists are compelled by both interest and practical necessity to explain both highly
complex life cycles and more life history variation than fits comfortably into general theoretical
frameworks that have been developed to date. This book clearly establishes that life history
variation within insect species, and often within populations, is as important as general patterns
at higher taxonomic levels. Understanding how such variation is maintained and distributed
geographically and taxonomically can become a major unifying theme in future work about
insect life history.
The book is carefully written and produced. It is admirable for its clarity and scarcity of
typographical error. Each chapter section is clearly summarized and this is very handy for both
preview and review. The bibliography is immense, including nearly 2000 entries, most of which
are cited in constructive arguments in the text. The indicies are superb and will make the book
valuable as a reference on any entomological bookshelf. Seasonal Adaptions of Insects is a
perfect basis for graduate seminar courses, and for allowing professional entomologists to catch
up on an exploding literature.
Although the book confronts complexity squarely and without apology, the organization and
presentation of material makes it easily digestible. Of the two objectives mentioned in the
introductory chapter, the authors clearly succeed with the first; the complexities of insect life
Book Reviews
197
history are superbly described and well documented. They light the way for an assault on the
second objective by urging researchers to consider the genetic basis of insect life history, and to
view life history in the more holistic context of the diapause syndrome. It is fair to point out
that this task was begun by collections of authors at the last few International Congresses of
Entomology [Dingle, H. (ed.), 1978, Evolution of Insect Migragion and Diapause,
Springer- Verlag; Brown, V.K. and I. Hodek (eds.), 1983, Diapause and Life Cycle Strategies
in Insects. Junk]. However, Tauber, Tauber and Masaki’s new book is the first attempt to
review the relations between insect life history and seasonality within a general entomological
framework. Their book is highly successful and entomologists of all persuasions will profit from
reading it. I hope that it will be widely read, and trust that it will inspire attention to the many
fascinating problems that it raises.
John R. Spence
Department of Entomology
University of Alberta
Quaest. Ent., 1987, 23 (1)
Quaestiones,
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 23
NUMBER 2
SPRING 1987
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 23 Number 2 1987
CONTENTS
Sperling-Evolution of the Papilio machaon species group in western Canada (Lepidoptera:
Papilionidae) 198
EVOLUTION OF THE PAPILIO MACHAON SPECIES GROUP IN WESTERN CANADA
(LEPIDOPTERA: PAPILIONIDAE)
Felix A.H. Sperling'
Dept, of Entomology
University of Alberta
Edmonton, Alberta, T6G 2E3
Canada
Quaestiones Entomologicae
23: 198-315 1987
ABSTRACT
Three species of the Papilio machaon group live in western Canada: P. machaon, P.
zelicaon and P. polyxenes. Most specimens of these three species are distinguished on the basis
of morphological, electrophoretic or ecological characters. However, all three species
hybridize along zones of parapatry, as well as in restricted areas within regions of sympatry.
All populations of the species group whose larvae eat Artemisia dracunculus are considered
subspecies of P. machaon. Also P. machaon pikei (type locality: Dunvegan, Alberta ) is
described as a new subspecies from the Peace River region of northern Alberta and British
Columbia. P. m. avinoffi and P. kahli are considered different expressions of hybridization
between P. machaon and P. polyxenes, while P. nitra is treated as a genetically integrated
morph within P. zelicaon.
Three separate principal components analyses (PCAs), on 10 electrophoretic loci, 11 wing
and body color characters, and the combined data set, respectively, gave very similar relative
distributions of individuals and populations. Hybrid populations had intermediate mean
character values, but much broader ranges of variation than those of the parental species.
Enzyme genotypes were tested for conformance to Hardy-Weinberg proportions, with the same
loci showing major interruptions in gene flow in some regions, but not in areas where hybrid
swarms had formed. Discriminant function analyses of specimens reared on different
foodplants supported the conclusions based on PCAs, and gave better species separations in
some regions.
The P. machaon group includes a wide variety of populations associated with different
foodplant, habitat and climate conditions. The genetic versatility of the group leads either to
ecological divergence between populations, or to localized genetic merging. Large interspecific
hybrid populations have formed in central and southern Alberta (P. zelicaon X machaon,) and
in central Manitoba (P. polyxenes X machaon,) , and show varying amounts of ecological
isolation from the parental species. On the other hand, P. m. pikei is a grassland race which
was probably derived early in the Holocene from arctic) alpine populations of P. machaon.
Hybrid swarm formation predated habitat alteration by European settlers in central Alberta,
but may have been a more recent and agriculturally related phenomenon in central Manitoba.
Both P. zelicaon and P. machaon also show introgression of a black wing morph from P.
polyxenes. This morph has moved far beyond the range of P. polyxenes, and such specimens
show electrophoretic and ecological characteristics identical to those of the populations of P.
zelicaon and P. machaon in which they are now found.
Present address: Section of Ecology and Systematics, Corson Hall, Cornell University, Ithaca, New York 14853-2701,
U.S. A.
200
Sperling
In general, allopatric differentiation and peripheral race formation appear to account for
most of the systematic structure of the P. machaon group, though gene exchange between
species is also an important factor. The presence of a widespread but uneven pattern of gene
flow among the species of the P. machaon group necessitates a loose application of current
species concepts, and causes considerable uncertainty in phylogenetic reconstructions.
Comparison of features of the P. machaon species group with those of potential sister
groups in the genus Papilio indicates that color pattern of the ancestral stock of the former
was probably like that of P. machaon itself, and that the larvae were probably
umbellifer-feeders. The P. machaon group contains P. alexanor, which has problematical,
though basal, affinities to the remaining four lineages (1, P. machaon + hospiton; 2, P.
zelicaon; 3, P. polyxenes + joanae + brevicauda; and 4, P. indraj. The Holarctic P. machaon
lineage is older than the other three, which are Nearctic in distribution. Relationships of the
latter three lineages to one another are unclear, and their divergence is represented as a
trichotomy. Calculation of Nei’s genetic distance suggests that P. machaon, P. zelicaon and P.
polyxenes arose in Pliocene time.
North America was invaded via Beringia by the ancestral stock of the P. machaon group
and, during Pleistocene time, by P. machaon itself. Environmental changes, particularly
glaciation, caused several periods of range expansion and retraction. Differentiation in
isolation led to shifts in ecological features, such as larval foodplants, as well as
morphological features and electromorph frequencies. Subsequent contact led to genetic
introgression and hybridization, depending in part on the extent to which isolates had diverged
ecologically. These events are reflected in western Canada in the genetic structure and
relationships of: three species; five subspecies of P. machaon; two combinations of hybrid
swarms; and evidence of introgression, particularly of genes for black wing morphs.
RESUME
Trois especes du groupe Papilio machaon sont retrouvees dans I’Ouest Canadien: P. machaon, P. zelicaon and P.
polyxenes. La plus part des specimens de ces trois especes se distinguent sur la base de characteres morphologiques,
electrophoretiques et ecologiques. Toutefois, des individus hybrides de ces trois especes sont retrouves aux abords des
zones parapatriques, ainsi que dans des endroits restreints d I’interieur des regions sympatriques.
Toutes les populations de I’espece groupe Artemisia dracunculus dont les larves se nourissent, sont considerees
comme sous-especes de P. machaon. Une nouvelle espece, P. machaon pikei (localite type: Dunvegan, Alberta ) est decrite
pour la region de Peace River, au nord de I’Alberta et de la Colombie-Britanique. P. m. avinoffi et P. kahli sont
consideres comme differentes expressions de I’hybridation de P. machaon et P. polyxenes, tandis que P. nitra est consider e
comme un morphotype genetiquement integre d P. zelicaon.
Trois differentes Analyses en Composantes Principales (ACPs) effectuees respectivement sur 10 loci
electrophoretiques, 11 caracteres bases sur la couleur des ailes et du corps, et sur un groupe combine de ces caracteres,
ont produit des distributions relatives tres similaires pour les populations et les individus. Les populations hybrides
demontrent des caracteres moyens de valeurs intermediates, mais possedent des variations bien superieures d celle des
especes parentales.
Les enzymes genotypiques son testes pour conformite aux proportions de Hardy-Weiberg, utilisant les meme loci
demontrant une interruption majeure du courant d echange genetique de quelques regions, mais non dans les regions ou
les essaims hybrides se sont formes. Une analyse en fonctions discriminantes utilisant les specimens ecimens eleves sur
differentes source de nouriture, supportent les conclusions basees sur les A.C.P., et demontre pour quelques regions une
meilleure separation des especes.
Le groupe P. machaon inclut une grande variete de populations associees d differentes sources alimentaires vegetales,
habitats et conditions climatiques. La versatility genetique du groupe conduit soit d une divergeance ecologique, soit d
des combinaisons genetiques localisees. De larges populations hybrides se sont formees dans le centre et le sud de
T Alberta (P. zelicaon X machaon) et dans le Maitoba central (P. polyxenes X machaon). Ces populations demontrent
differents degres d’isolation ecologique de leurs especes parentales. Toutefois, P.m. pikei considere comme une espece des
prairies, a probablement evolue au debut de I’Holocene, d’une population arctique/ alpine de P. machaon. La formation
d'essaims hybrides dans T Alberta centrale, precede la periode d’alteration de Thabitat induite par I’etablissement des
Papilio machaon species group
201
colons Europeens; mais est probablement plus recente et reliee h I’agriculture dans le Manitoba central. P. zelicaon ainsi
que P. machaon demontrent /’integration d’un morphotype h aile noire provenant de iespece P. polyxenes. Ce type s’est
reparti bien au delh de I’aire de distribution de P. polyxenes, et de tels specimens demontrent des characters
electrophoretiques et ecologiques identiques it ceux des populations de P. zelicaon et P. machaon dans lesquelles ils sont
maintenant retrouves.
De facon generale, la differenciation allopatrique et la formation de races periferiques semblent etre responsable
d’une grande portion de la structure phylogenetique du groupe P. machaon, bien que Pechange genetique interspecifique
soit aussi un facteur important. La presence d’un courant d’echanges genetiques largement rependu, mais irregulie, entre
les especes du groupe P. machaon necessite une utilisation au sens large du concept d’especes, et de ce fait induit de
considerables incertitudes lors de la reconstitution phylogenetique.
La comparaison des traits caracteristiques du groupe-espece P. machaon a ceux de potentiels groupes-soeurs dans le
genre Papilio, indique le patron de couleurs de la souche ancestrale etait probablement comme celui de P, machaon
lui-meme, et que les larves se nourissaient probablement de plantes umbelliferes. L’espece P. alexanor, qui fait parti du
groupe P. machaon, a des rapports fondamentaux mais problematiques aux quatres autres lignees (1, P. machaon +
hospiton; 2, P. zelicaon; 3, P. polyxenes + joanae + brevicauda; and 4, P. indraj. Parmi les quatres lignees, la lignee
Holarctique P. machaon est plus ancienne que les trois autres qui sont de distribution Nearctique. Les rapports entre les
trois lignees nearctiques sont ambigus et ainsi la divergence est decrite en trichotomie. Le calcul de distance de Nei
semble indiquer que P. machaon, P. zelicaon et P. polyxenes proviennent de Page Pliocene.
L’amerique du nord a ete envahie via le Beringia par la souche ancestrale du groupe P. machaon et durant Page
Pleistocene par P. machaon lui-meme. Des changements d’environnement en particulier la glaciation, ont cause plusieurs
periodes d’expansion et de retraction de distribution. Les differences d’isolement ont mene aux changements de
caracteres ecologiques tels que plantes-hotes des larves autant que caracteres morphologiques et frequences
d’electomorphs. Le contact subsequent a mene h Pintrogression et it P hybridisation, selon le degre de divergence
ecologique des isolees. Ces evennements se sont reflechis dans Pouest du Canada par la structure genetique et les
rapports entre trois especies; cinq sous-especes de P. machaon, deux combinaisons de deux essaims hybrides; et Pevidence
d’introgression, en particulier les genes pour les morphs aux ailes noires.
TABLE OF CONTENTS
Introduction 201
Materials and Methods 206
Classification and Recognition of taxa 218
Morphometric and Electrophoretic Characters 224
Ecological Characteristics 258
Evolutionary Hypotheses 275
Acknowledgements 291
Literature Cited 291
Appendices: Tables 12-14 301
Index 312
INTRODUCTION
The genus Papilio traditionally includes 30 to 40 species groups, of which the Papilio
machaon L. group is one. Adults of the group are predominantly black and yellow (Plates 1-3),
and are relatively large (forewing length: 3.5 to 5.0 cm). The P. machaon group is principally
defined by larval characters, especially a color pattern of black segmental bands and by use of
hostplants in the families Umbellifefae or Compositae.
Opinions about the number of species included within the P. machaon group range from
four (Eller, 1939) to between 10 and 20 species (e.g., Remington, 1968a). Eight species are
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202
Sperling
recognized in this study, and three of them are dealt with in detail. There is much less
difference of opinion about the limits of the group itself, with most recent authors following
Monroe (1961).
Most of the uncertainty about the number of species which should be recognized within the
P. machaon group is caused by discordant character variation. For example, P. machaon and P.
hospiton Gene remain distinct and easy to distinguish where they contact each other on Corsica
and Sardinia, but P. machaon has races in North Africa and the Arabian peninsula which are
much more similar to P. hospiton (Clarke and Larsen, 1986). Wing and body color pattern,
and wing shape, have been the main morphometric characters used to distinguish among P.
machaon group species. However, for identifications of P. machaon , P. zelicaon Lucas and P.
polyxenes Fabricius, no one character is completely dependable. Most diagnoses list several
characters, and there has been no effort published previously to quantify rigorously these
differences using multivariate techniques. The presence of several color polymorphisms within
populations has also been a consistent source of taxonomic confusion. The most notable
example is the adult morph with black wings, whose features are probably determined by
homologous alleles in different species.
Other kinds of taxonomic characters have shown only limited utility. Differences in male
genitalia, which are taxonomically valuable in many groups in the Papilionidae, are of minor
importance in the P. machaon group. In North America adults of only one species, Papilio
indra Reakirt, can be consistently distinguished on this basis. Use of chromosomal characters
(Maeki and Remington, 1960; Maeki, 1976; Clarke et al. , 1977) involves substantial technical
difficulties. Although gel electrophoresis has been useful for elucidating systematic
relationships in many taxa, there have been no reports about variation of enzyme alleles in the
P. machaon group.
The P. machaon species group has a holarctic distribution. It occurs throughout North
America and in higher elevations as far south as Peru and Venezuela. It also ranges across
Eurasia, southward to South China and North Africa. The group is represented in virtually any
vegetation zone in this region. Different ecological races occupy habitats as varied as arctic
tundra, high altitude steppe, Saharan desert oases, temperate coastal forest, vegetable gardens
and citrus orchards. However, the western United States and Canada is the only part of the
range of the P. machaon group with extensive sympatry between species. All other regions
support only a single species, or a contact zone between two species which is maintained in part
by habitat segregation.
Studies of populations of the P. machaon group in sympatry are few, based principally on
samples from geographically distant localities. Comparisons among many samples show either
marked phenetic or ecological differences or a confusing interplay of character variation.
Clearly, intensive surveys are needed of population variation in regions of sympatry or
parapatry between major species of the P. machaon group. As well, there has been little explicit
recognition of natural hybridization among species in the group. Since there is evidence of
hybridization among the three species included in this study, and virtually all studies on hybrid
zones in other taxa involve only two species, work on the P. machaon group is of significance to
the general study of hybrid phenomena.
Historical Perspective
The historical context of work on the machaon group has influenced many of the resulting
taxonomic decisions. In particular, the pattern of exploration of new regions and changing
Papilio machaon species group
203
motivations for research have left a diverse legacy of names for taxa. These names represent a
number of major stages in the development of systematics as a separate discipline.
The recognized starting point for modern taxonomy, Linnaeus’ Systema Naturae , Tenth
Edition (1758), contained the description of P. machaon itself, the type species for the genus
Papilio and, by extension to the Papilioniodea, it is the type species for all butterflies. New
names such as P. polyxenes (Fabricius, 1775) and P. asterius (Stoll, 1782) were also published
in that early period of endeavor to provide a full description of nature and what were perceived
as God’s works.
A steady stream of taxonomic descriptions followed, in step with the economic conquest of
the remaining parts of North America and Eurasia. These provided a sense of the primacy of
order, and the security and power of knowledge, for the growing class of people who engaged in
this uniquely western activity. In addition, the practice of figuring the name of the author
prominently behind the name of a taxon ensured that considerable effort and money was
expended in the race to acquire this form of immortality. Papilio zelicaon was described by
Lucas in 1852 under such circumstances, edging out Boisduval’s (1852) P. zolicaon by a matter
of a few months (Dos Passos, 1962). Obviously new species were exhausted relatively quickly,
while the plethora of geographic races in the species of the P. machaon group provided an
excuse for new names that has not been exhausted to this day. Subspecies names became
fashionable around the turn of the last century and many of the older names were subsumed
under the very oldest names. The most recent treatment of Papilio machaon throughout
Eurasia (Seyer, 1974, 1976a, 1976b, 1977) recognizes 36 subspecies in that region alone and
synonomizes many more names.
The settlement of western North America in the latter part of the 1800’s was integrally
associated with many new names. P. machaon aliaska was contributed by Scudder (1869),
based on material collected by an American lieutenant in a “Russo-American Telegraph
Expedition” to Alaska. W.H. Edwards added a number of other names in the same sort of
environment. He became wealthy through investments in the expanding railway industry of this
period, and his financial security allowed him to play an important role in North American
butterfly taxonomy. To the Papilio machaon group he contributed the names P. bairdii (1866),
P. oregonius (1876) and P. nitra (1884), among others. Several of these taxa are treated in this
study.
The discovery and classification of new taxa saw a watershed of sorts in the Victorian era.
The abundance of newly collected specimens led to a new understanding of their inexhaustible
variation and yet also to a philosophical rift among taxonomists. Darwin’s publication of The
Origin of Species (1859) provided a starting point for this process. By 1883 W.H. Edwards was
arguing that criticism of his species designations in the P. machaon group was tantamount to a
refusal of the teachings of Darwin, which was in turn equivalent to failing to admit the truths of
Copernicus. Yet Edwards’ names are an example of far greater emphasis on phenetic
homogeneity and covariance than on hybridization information. This is particularly noticeable
in his treatment of the P. machaon populations in the western United States, upon which he
bestowed four specific epithets despite knowing that the forms the names referred to were
probably all part of the same extended gene pool.
Hagen’s (1882) report on the P. machaon group was published during the same period as
Edwards’ work, but showed the opposite tendency to that of Edwards. He looked for and found
specimens with character states and combinations that were intermediate between those
attributed to all the previously described forms. He concluded that all the North American
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204
Sperling
species of the P. machaon group (excluding P. indra ) should be considered as local or climatic
varieties of P. machaon.
Edwards’ views prevailed among butterfly taxonomists, in part because David Bruce found
that larvae of P. bairdii fed on Artemisia dracunculus Linneaus (Edwards, 1893 and 1895).
This foodplant is a member of the Compositae, rather than the Umbelliferae or Rutaceae that
are used by larvae of P. zelicaon. Also Bruce found that, in Colorado, P. bairdii adults were
polymorphic for a yellow wing form which was much more like that of P. oregonius and P.
zelicaon than the mostly black form it had previously been known for. However, Edwards’
(1895) taxonomic response was to describe the yellow form from Colorado as yet another new
species, P. brucei. This conflict between an increasing understanding of evolutionary
phenomena such as polymorphism and the need for comprehensible, consistent classifications
has continued to provide friction today.
Lord Walter Rothschild dealt with this conflict in a more balanced fashion. He was the
epitome of acquisitiveness in the dying days of Victorian thought (Rothschild, 1983), and
played a positive role in the systematics of the P. machaon group. His curator, Karl Jordan,
contributed substantially to putting taxonomic practice in better accordance with evolutionary
principles such as geographic differentiation (Mayr, 1955). Rothschild and Jordan consistently
and accurately applied the concept of geographic races to their formal recognition of
subspecies, and by this contributed to a lasting reduction in the number of taxonomic names
used in the P. machaon group. Interestingly enough, they saw some specimens from west of
Calgary from the same P. machaon X zelicaon hybrid populations which piqued my own
interest in the group. They continued to use relatively traditional assignments, but remarked on
the close resemblance of the black individuals from this area to the black forms of P. bairdii
(Rothschild and Jordan, 1906).
The 1930’s saw a fresh burst of new names proposed for taxa within the P. machaon group.
Several of these were described from western Canada and Alaska. A. H. Clark (1932)
contributed P. machaon hudsonianus from the boreal zone of northern Manitoba and Ontario,
and P. machaon petersi from Alaska. Chermock and Chermock (1937) described P. machaon
race avinoffi and P. nitra form kahli , both from the Riding Mountains of central Manitoba.
McDunnough (1939) described Papilio machaon dodi from the prairies of southern Alberta
and Saskatchewan. A number of new taxa were also described for the rest of North America
during this period, while Eller (1936) produced a major monograph on the races of Papilio
machaon in Eurasia. Eller (1939) followed this with a shorter treatment of the P. machaon
group worldwide, in which he, like Hagen (1882), classified all North American taxa except P.
indra as subspecies of P. machaon.
The names P. machaon avinoffi and P. m. petersi are now generally accepted as synonyms
of P. m. hudsonianus and P. m. aliaska , respectively. P. m. dodi is accepted as a valid taxon,
but is variously placed as a subspecies of P. machaon , P. bairdii , or P. oregonius. P. nitra form
kahli was elevated without explanation to species status by Wilson (1961) and has generally
continued to be used in that manner. Eller’s work was rejected in North America, and cited as
an example of poorly informed taxonomy (Remington, 1968a).
During the late 1950’s, an understanding of the Papilio machaon group was placed on a
rather different footing. The technique of mating Papilio by hand was described in detail by
Clarke (1952), and Clarke collaborated in obtaining numerous hybrids in the following years,
many of them within the P. machaon group (Clarke and Knudsen, 1953; Clarke and Sheppard,
1953, 1955a, 1955b, 1956a; Clarke et al., 1977). Hand pairing became a commonly used
Papilio machaon species group
205
technique in hybridizing even distantly related species within the genus Papilio. The papers
which Clarke co-authored with Sheppard have been the best studies to date on the genetics of
the adult and larval color patterns of various taxa within the P. machaon group. They extended
their experiments to other species of Papilio and produced a number of classic works, including
studies on the African mimetic complex of P. dardanus. (The understanding of the interactions
of genes which was gained from these hybridizing experiments was applied to the prevention of
rhesus haemolytic disease in newborn humans, and Clarke was eventually knighted for his
work.)
In the United States, Remington also conducted numerous hybridization and rearing
experiments on the P. machaon group. His first report (1956) on this work was concerned with
a collecting trip made to the Riding Mountains to obtain P. nitra form kahli, but he did not
fully publish any of this research. His last report (1968a) included a description of a new
species closely allied to P. zelicaon. The separation of Remington’s P. gothica (1968a) was
based on the hybrids it produced, as well as slight color pattern and ecological differences from
P. zelicaon. Sibling species were in fashion in the evolutionary biology and systematics of the
time, since they confirmed the primacy of genetic considerations in species definitions.
However, Remington’s P. gothica was soon criticized for a variety of reasons (Clarke and
Sheppard, 1970; Shapiro, 1975; Emmel and Shields, 1980) and the only remnant of his concept
survived in the form of a subspecific division of P. zelicaon.
The name of Remington’s (1968a) taxon was later changed to P. zelicaon nitra on the basis
of hybridization and rearing studies by Fisher (1977). These showed that Edwards’ (1884) P.
nitra was just a dark form of P. zelicaon , produced by a single dominant allele. Fisher theorized
that the black form had arisen through the introgression of genes from P. polyxenes, like
Remington’s (1956, 1958) earlier thoughts on the origin of P. nitra form kahli through
hybridization between P. machaon and P. polyxenes.
One of Remington’s students, S. Ae, also carried out numerous hybridizations of Papilio
species. He continued this work for more than two decades in Japan, publishing numerous
progress reports and culminating in a major paper on Papilio phylogeny (Ae, 1979). He showed
that even relatively distant interspecific crosses could produce adults, and many crosses
between species within the P. machaon group had a reasonable degree of FI viability. Ae’s
work is the tip of a veritable iceberg of Papilio hybridization studies, carried on by numerous
enthusiasts, usually amateur, who rarely if ever publish. Similar situations can be found in
saturniid and killifish circles, where the considerable effort to do such work seems to be
maintained by a joy derived from the creation of new kinds.
During the last 15 years a substantial number of publications have appeared about the
ecology of various members of the P. machaon group. These include studies of oviposition
behavior ( e.g ., Wiklund, 1981), larval growth ( e.g ., Scriber and Feeny, 1979), diapause
dynamics (e.g., Sims, 1980) and pupal color determination (e.g., Smith, 1978), among others.
Although aimed at an ecological audience, they demonstrate a diversity of potential adaptive
mechanisms. A consequence to systematic studies of the P. machaon group is that they show
how numerous ecological races could have arisen, even within the last century (Shapiro and
Masuda, 1980), as variations derived from a single basic gene pool.
There is considerable potential for future research on the Papilio machaon group. Clarke
and Sheppard (1955b) stated that “It is clear that the Machaon-group provides some of the
most suitable material ever investigated in animals for studying the process of speciation in
detail, taking into account genetic, ecological and behaviour differences as well as time”. I
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206
Sperling
agree wholeheartedly with this view, in part because the wide variation present within this
group of beautiful butterflies makes genetic investigations relatively tractable.
However, study of the P. machaon group is also interesting because the variety of hybrid
interactions poses conceptual problems in understanding the nature and origin of species. Such
hybrid interactions “contrast two views of the species: as a set of populations delimited by
genetic barriers to gene exchange; and as a set of populations maintained in a particular stable
equilibrium by selection” (Barton and Hewitt, 1985). The appropriateness of study of the P.
machaon group to an examination of species concepts is illustrated by the fact that Hagen
(1882) and Edwards (1883) expressed similar opposing views a century ago.
MATERIALS AND METHODS
Acquisition of Specimens
I examined about 1200 specimens from my own collection, and about 2000 from the
collections of the individuals and institutions listed in Table 1. The majority of material from
my own collection, including the holotype and a series of paratypes of P. machaon pikei, have
been deposited at the Canadian National Collection, Ottawa. Voucher specimens have been
deposited at the University of Alberta Strickland Museum, as well as locality listings for all
specimens examined in the course of this study.
I tried to obtain as many specimens as possible from each of a number of localities located in
a rough grid pattern with intervals of 150-200 km across Alberta and northern British
Columbia. I collected 20 or more adults from most of these localities, though I had to return to
several of them a number of times in different years to do so. These localities comprise much of
the geographic survey portion of my study, and are compared to more widely spaced localities
in the remainder of western Canada and adjacent United States. I sampled most of the Alberta
and northern British Columbia localities several times throughout the summer. Some
population samples include adults obtained from larvae collected within about five km of a
hilltop locality, as well as specimens from the hilltop itself. They have been distinguished from
the wild-collected adults in the sections dealing with foodplant associations. Wherever possible,
material taken by other collectors at or near a major locality was included in the morphometric
portion of my study.
During the summers of 1980 through 1984 I collected about 650 adults of the Papilio
machaon group, mostly on hilltops and prominent edges of river banks in Alberta and northern
British Columbia. For some samples, I obtained pupae through the mail from other collectors.
These included all the samples used in electrophoretic analysis for P. polyxenes, as well as most
of the material from the P. polyxenes X machaon hybrid zone and most of the P. zelicaon
specimens from southern British Columbia. I also collected about 1800 larvae during
1980-1984, and reared about 450 of these to the adult stage on the same foodplant from which
the larvae has been collected. All larval records are listed in Table 14.
Pressed samples of all new foodplant records were identified by Dr. J. G. Packer,
Department of Botany, University of Alberta. Records attributed to individuals other than me
were identified by those individuals.
Papilio machaon species group
207
Table 1 . Sources of specimens examined of the P. machaon species group
Names of Curators of institutional collections are listed at ends of entries.
Acorn, J.H., Dept, of Entomology, University of Alberta, Edmonton, Alberta, T6G 2E3
Canada
Alberta Provincial Museum, Natural History Dept., 12845-102 Ave., Edmonton, Alberta, T5N
OM6 Canada (A.T. Finnamore)
Allyn Museum of Entomology, Sarasota, Florida, 3261 1 1 U.S.A. (L.D. Miller)
Bird, C.D., Box 165, Mirror, Alberta, TOB 3C0 Canada
British Columbia Provincial Museum, Parliament Buildings, Victoria, British Columbia, V8V
1X4 Canada (R.A. Cannings)
British Museum (Natural History), Cromwell Road, London, SW7 5BD, England (R.I.
Vane-Wright)
Canadian National Collections of Insects, Arachnids and Nematodes, Biosystematics Research
Institute, Ottawa, Ontario, K1 A 0C6 Canada (J.D. Lafontaine)
Guppy, C.S., 4120 St. Georges Ave., North Vancouver, British Columbia, V7N 1W8 Canada
Hilchie, G.J., Dept, of Entomology, University of Alberta, Edmonton, Alberta, T6G 2E3
Canada
Hooper, D.F., Somme, Saskatchewan, SOE 1N0 Canada
Hooper, R.R., Box 205, Fort Qu’Appelle, Saskatchewan, SOG ISO Canada
Klassen, P., Box 212, Elm Creek, Manitoba, ROG 0N0 Canada
Kohler, S.J., Forest Insect and Disease Section, Montana Department of Natural Resources
and Conservation, 2275 Spurgin Road, Missoula, Montana, 59809 U.S.A.
Kondla, N.G., 22 Brock Place, Lethbridge, Alberta, T1K 4C7 Canada
Krivda, W., 319 Crossley Ave., The Pas, Manitoba, R9A 1B7 Canada
Pike, E.M., 141 0-4th Ave. N.W., Drummheller, Alberta, T0H 1L0 Canada
Royal Ontario Museum, Dept, of Entomology, 100 Queens Park, Toronto, Ontario, M5S 2C6
Canada (R. Jaagumagi)
Saskatchewan Museum of Natural History, Wascana Park, Regina, Saskatchewan, S4P 3V7
Canada (R.R. Hooper)
Shepard, J.H., Sproule Creek Road, RR#2, Nelson, British Columbia, V1L 5P5 Canada
Shigematsu, S., 2314-22nd St. S., Lethbridge, Alberta, T1K 2K2 Canada
Sperling, F.A.H., Box 3 1 , Bragg Creek, Alberta, TOL OKO Canada
University of Alberta, Entomology Dept., Strickland Museum, Edmonton, Alberta, T6G 2E3
Canada (G.E. Ball & D. Shpeley)
University of British Columbia, Zoology Dept., Spencer Entomological Museum, Vancouver,
British Columbia, V6T 2A9 Canada (S. Cannings)
University of Calgary, Biology Dept., Calgary, T2N 1N4 Canada (G. Pritchard)
University of Manitoba, Entomology Dept., Winnipeg, Manitoba R3T 2N2 Canada (R.E.
Roughley)
Waterton National Park, Interpretation Administration, Waterton, Alberta, T0K 2M0 Canada
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Table 2. Descriptions of morphometric characters of adults and larvae of the P. machaon
group, used in multivariate analyses
Numbering of character states is the same as that used in multivariate analyses.
A. Extent of yellow scaling in cell Cu2, in anal margin of dorsal hindwing (Fig. 2)
1. All except a few yellow scales are restricted to area from inner margin of median
dark band to less than halfway to divergence of veins Cu2 and Cul. ( e.g ., Fig. 3)
2. Virtually all yellow scales restricted to area between median band and divergence of
Cul and Cu2.
3. Yellow scales extend past junction of Cul and Cu2, but less than halfway between
Cul-Cu2 junction and wing base.
4. Yellow scales extend from median band to more than 3/4 of way to wing base, (e.g.,
Fig. 2)
B. Shape of pupil in anal eyespot of dorsal hindwing. (Fig. 1)
1 . Thin line at lower edge of blue region, connected to wing margin, (e.g.. Fig. la)
2. Club shaped, in that thinnest portion near margin is less than half the width of
thickest portion closer to center of eyespot. (e.g., Fig. lb, 2, 3)
3. Oblong spot at lower edge of red area, not connected to margin, (e.g., Fig. lc)
4. Round spot centerd in red area, generally with less than two times as much red above
pupil as below it. (Fig. Id)
C. Extent of black scales between blue and red portions of anal eyespot of dorsal forewing.
(Fig. 1-3)
1 . Black line extending along less than 1/4 of blue-red boundary, (e.g., Fig. la )
2. Black line separating blue from red along between 1 /4 and 3/4 of the width of anal
eye. ( eg. Fig. lc )
3. Black line separating more than 3/4 of blue from red (boundary line may be wide or
narrow). (e.g.. Fig. lb, Id, 2, 3)
D. Color of hairs on tegula. (Fig. 4 and 6)
1 . Virtually all hairs on tegula yellow, (e.g., Fig. 4a-b and 6a-e)
2. Less than half as much yellow on tegula as in typical yellow morph adults but more
than about 15%. (e.g., Fig. 4c)
3. Virtually all tegula hairs black, (e.g., Fig. 4d)
E. Extent of yellow scales in basal half of disc of central forewing. (Fig. 5)
1. Yellow scales in more than 50% of area, (e.g., Fig. 5a)
2. Thick streaks or a more general flush to less than 50% of total area, (e.g.. Fig. 5b)
3. Few thin streaks or a light sprinkling of yellow scales, (e.g., Fig. 5c)
4. Virtually no yellow scales in basal half of disc of ventral forewing, (e.g., Fig. 5d-e)
F. Extent of yellow scales of postmedian yellow band in apical cell of ventral forewing.
(Fig. 5)
1. Postmedian spot large, occupying more area than bordering black scales, (e.g., Fig.
5a-c)
2. Definite patch of diffuse yellow, but occupying less area than black scales, (e.g., Fig.
5d)
3. Virtually no postmedian yellow scales in cell, (e.g., Fig. 5e)
(continued on next page)
Papilio machaon species group
209
Table 2 (continued)
G. Number of cells with orange patch in postmedian area of ventral hind wing. (Fig. 3)
1 . No cells with a distinct patch of orange cells on distal side of postmedian yellow area.
2-8. The total number of cells, plus one, which have a distinct patch of orange, up to a
total of 7 cells. Postmedian wing cells which are covered in only black scales are
assumed to have an orange patch, as in some female P. polyxenes.
H. Amount of yellow hair on metathorax below base of hindwings. (Fig. 6)
1 . Yellow hairs extend around the ventral part of metathorax. ( e.g ., Fig. 6a)
2. Substantial patches of yellow hairs on each side of metathorax which do not meet
ventrally. {e.g., Fig. 6b-d)
3. All metathoracic hairs black, or at most a very few short yellow hairs restricted to
immediate base of wing, (e.g., Fig. 6e-f)
I. Ventral abdominal line. (Fig. 6)
1. All abdominal segments with distinct ventral line of yellow hairs along sagittal plane.
(e.g., Fig. 6a)
2-9. The total number of segments, plus one, which do not have a distinct patch of yellow
scales or hairs along sagittal line. Start counting from first abdominal segment after
thorax, to a maximum of 8.
J. Lateral abdominal yellow. (Fig. 6)
1 . Broad band of yellow on each side, extending along length of abdomen (male claspers
excluded), (e.g., Fig. 6a-d)
2. Large square lateral spots on some or all abdominal segments, with narrow divisions
between spots, (e.g., Fig. 6e)
3. Small round lateral yellow spots on all or most segments, distance separating spots
generally greater than width of spots, (e.g., Fig. 6f)
K. Upper abdominal spots. (Fig. 6)
1 . All abdominal segments with a distinct pair of subdorsal yellow spots, separated from
lateral abdominal band or line of spots (character J., above), (e.g., Fig. 6e-f)
2-9. The total number of segments, plus one, which do not have at least one yellow spot
distinct from yellow line.
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210
Sperling
Figures 1 to 4. Figure 1. Anal eyespot of dorsal hind wing: A, P. m. aliaska Mi. 391, Alaska Hwy., British Columbia. B, P.
m. dodi Nacmine, Alberta. C, P. zelicaon X machaon Nacmine, Alberta. D, P. zelicaon Wintering Hills, Alberta. Figure
2. Yellow scaling of dorsal hindwing anal margin P. m. oregonius , Brewster, WA. Character states for yellow anal scales
are numbered beside figure. Figure 3. Ventral hindwing, with location of orange scales P. p. asterius, Karlsruhe, North
Dakota. Arrow shows postmedian band, which contains orange scales. Figure 4. Dorsal view of thorax, showing tegula: A,
P. zelicaon yellow morph Wintering Hills, Alberta; B, P. zelicaon X machaon black morph Bragg Creek, Alberta; C, P. p.
asterius Karlsruhe, North Dakota; D, P. p. asterius Karlsruhe, North Dakota.
Papilio machaon species group
211
Figure 5. Yellow scales on disc and apex of ventral forewing. Arrows on Figure 5b show location of disc and apical cell: A,
P. m. aliaska Pink Mountain, British Columbia; B, P. m. dodi yellow morph Nacmine, Alberta; C, P. zelicaon X machaon
black morph Bragg Creek, Alberta; D, P. zelicaon X machaon black morph Bragg Creek, Alberta; E. P. p. asterius
Karlsruhe, North Dakota.
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212
Sperling
Figure 6. Lateral view of thorax + abdomen. Wings and head have been removed, a, P. m. aliaska Pink Mountain, British
Columbia; b, P. m. dodi yellow morph Nacmine, Alberta; c, P. zelicaon yellow morph Wintering Hills, Alberta; d, P.
zelicaon X machaon yellow morph Bragg Creek, Alberta; e, P. zelicaon X machaon black morph Bragg Creek, Alberta; f,
P. p. asterius Karlsruhe, North Dakota.
Papilio machaon species group
213
Table 3. Names, sources and quantities of proteins, and conditions of gels used in
electrophoretic analyses of specimens of the P. machaon species group.
Characters Examined
Morphometric characters. — Eleven morphometric characters, defined in Table 2, and
illustrated in Figures 1 to 6, were scored for adults used in this study. Individuals were scored
against both the written descriptions of character states and a number of standard specimens.
Standard and illustrated specimens were labelled as such and are in the Strickland Museum,
University of Alberta.
The choice of characters for analysis was made for the following reasons:
1. The characters had been used previously by other workers for distinguishing between species.
Those listed by Remington (1968a), were especially useful in this regard, because of the clear
manner in which they were defined. This allowed me to relate populations in western Canada to
particular species concepts developed elsewhere in North America, and also to test diagnostic
structural characters against variation in electrophoretic characters.
2. The characters had to be fast and easy to score by eye, and hence allow accurate processing
of numbers of specimens in a relatively short time.
3. There had to be a large amount of variation in character states expressed in the major study
area, western Canada. The purpose of this criterion was to maximize the likelihood that useful
information would be recorded when a character was scored.
4. Characters were chosen which appeared to vary fairly independently of each other within
populations, to maximize the likelihood of sampling the effects of several different genes, and
hence of obtaining information of significance to gene flow between populations. I tried to
minimize the number of times I scored the presence of the gene for black color, which is known
to affect a large area of the adult wings and body (Clarke and Sheppard, 1953, 1955b and
1956a; Fisher, 1977).
I also recorded color of spots on each larva. Spot color was determined by eye and by
comparing larvae against each other. I did not use consistently a standard for comparison, but I
judge error in assignment of larvae to either a yellow or an orange group to be small. Orange
spotted larvae included both the pale orange and bright orange groups delineated by Clarke and
Sheppard (1955b). Records were also obtained by contacting lepidopterists who had reared
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214
Sperling
larvae in western Canada, and by examining published records for spot color for the whole P.
machaon group.
Electrophoretic characters. — A total of 860 adults of the Papilio machaon group were used
in electrophoretic analyses. These were frozen live and maintained at -20 C, and most were
homogenized within a few weeks of being frozen. However, bands were still readable for whole
specimens kept frozen for more than two years.
Tissue for electrophoresis was obtained by dissecting thoracic and abdominal contents from
specimens which had been allowed to thaw on ice just before use. The corpus bursa of females
was not used, since it may have contained male spermatophores. After tissue had been removed,
specimens were pinned through one side of the thorax and mounted in the manner standard for
Lepidoptera curation. Each specimen was given a unique number and labelled to allow the
morphometric and electrophoretic character states to be correctly associated.
Thoracic and abdominal tissue samples were homogenized separately, each in 0.2 ml
homogenizing solution. Homogenate not used immediately was frozen and stored at -20 C in
the centrifuge tubes. Most samples gave interpretable bands from frozen homogenate, though
esterases and IDH deteriorated the most rapidly when subjected to successive freeze-thaw
cycles.
The electrophoretic apparatus used was the same as that described by Rolseth and Gooding
(1978), in whose laboratory and under whose direction I did the preliminary work for this
study. Some changes have been made in their methods since publication. These include: 1)
making the “B” solution with 25.6 ml of 1M H3P04, 5.7 gm Tris and 0.46 ml TEMED, brought
up to 100 ml with distilled water; 2) making the homogenizing buffer with 7.0 ml H20, 1.0 ml
B, .300 gm polyvinylpyrolidine and 40 mg DL-dithiothreitol; 3) forming stacking gels with 20
slots; and 4) sealing the edges of the apparatus with parafilm rather than a plug gel. Also gels
were used at pH 8.9 and pH 8.2, using 24 ml and 60 ml of 1M HC1, respectively, in the stock
“A” solution. Tissue was electrophoresed at 40 milliamp/gel for 2.0 hours in pH 8.9 gels, and
3.5 hours in pH 8.2 gels. Electrophoresis conditions for the loci used in this study are noted in
Table 3; Rf values listed in Tables 6 and 13 refer to electrophoresis performed under these
conditions.
Except for the use of MgCl2 rather than MnCl2 as a cofactor for IDH, staining methods are
fairly standard applications of recipes available in Shaw and Prasad (1970) or Brewer (1970).
Both esterase loci showed much greater activity with a-naphthyl acetate than with /3-naphthyl
acetate or AS-D-acetate, and were not significantly inhibited by eserine. APK was usually
stained with Coomassie blue, since it seemed to be the most common protein that could be
resolved by this method in thoraces. The identity of the APK bands was ascertained in several
homozygous individuals and the single individual (a male) with an allelic variant, using the
staining technique of Gooding and Rolseth (1979).
I stained for a variety of other enzymes, using recipes of Brewer (1970), Shaw and Prasad
(1970) and Menken (1980). These included ADH (with ethanol, isopropanol and sorbitol), AO
(with benzaldehyde and heptaldehyde), GLUO, GDH, GA-3-PD, GlyDH, LDH, G6PDH,
SDH, TO and XDH. In these attempts, bands were either very faint or did not appear.
All except three of the protein loci stained were dimers, while APK and Est4 were
monomers, and ME may have been a tetramer. None of the loci included in this study, except
possibly APK, were sex linked. Heterozygotes were present in both sexes for all other loci,
including G-6-PD, which is frequently sex linked in other taxa.
Papilio machaon species group
215
The genetic inheritance of alleles was not checked through breeding experiments. However,
since most of the local populations surveyed had allelic distributions that were at
Hardy-Weinberg equilibrium proportions, environmental induction of particular alleles seems
unlikely. Also the general distribution of alleles showed an excellent correspondence with
morphometric, and ultimately taxonomic, characters.
In any event, my taxonomic conclusions are not strictly dependent on genetic interpretations
of protein banding patterns. The bands are treated as equivalent to any other taxonomic
character and are more discrete than the character states in my morphometric analysis. An
additional advantage of the electrophoretic characters I used is that they probably represent a
relatively unbiased selection of loci. The choice of loci for staining was influenced by my ability
to obtain bands consistently and my ability to interpret variation in a consistent manner, both of
which factors are independent of the morphometric characters measured.
Numerical Analyses
Principal components analysis (PCA) helps to visualize clusters in multivariate data, is
relatively assumption-free, and provides a basis for internally consistent, simple comparisons of
both individuals and populations. PCA seems to give fairly accurate representation of distances
between data points, and has been used successfully in studies of hybrid swarms (Neff and
Smith, 1979; Pimentel, 1981). Relatively few characters appear to be required to elucidate
patterns of racial variation. Thorpe (1985) showed that patterns with 90% confidence could be
obtained from as few as 8-10 characters in grass snakes in Europe. Also, it is easy to relate
samples not used in the initial analysis to those that were, by applying factor loadings.
Three major PCAs were applied to an initial group of 728 specimens, almost all of which
were from western Canada and particularly Alberta, and all I had available, in early 1984, with
complete scores for all morphometric and electrophoretic characters considered in this study.
An effort had been made prior to this to obtain a reasonably broad sampling of the different
geographic populations and morphotypes known from the region, including those described by
other authors. One PCA was performed on only the morphometric characters, one on only the
electrophoretic characters and one on the combined morphometric and electrophoretic
characters together. The same individuals were used in all three PCAs, to give a more
meaningful basis for comparison of variation in character patterns. Then the factor loadings
were applied to individuals which could only be completely scored for one of the two main
character suites.
Discriminant function analysis (DFA) was applied to subgroups where PCAs did not give
clear indications of the number of species in a region or the nature of distinctions among them.
DFA was applied to reared adults obtained from southern and south-central Alberta, where
adults from different major populations showed only slight separation into the major groups
distinguished in other regions. Discriminations were based on the different foodplants, on which
larvae were collected. DFA was also applied to the geographically separated subspecies of P.
machaon , to give an assessment of the accuracy of identification of specimens.
All work for this section of the study was carried out with the Midas statistical package
(Fox and Guries, 1976) on the Amdahl computer system at the University of Alberta. The
principal component function was used only with the “unsealed” option in this package. The 1 1
morphometric characters were scaled before PCA or DFA, to make their ranges of variation
equivalent. Scaling factors and PCA/DFA loadings are included in Tables 12 and 13.
Electrophoretic character scores were not scaled, since these were already recorded in a manner
Quaest. Ent., 1987, 23 (2)
216
Sperling
that gave equivalent weights to each character.
Electrophoretic characters were scored one allele at a time, for each butterfly. Each allele
known from my work on the P. machaon group was considered a character with three character
states and scored: 1), if the allele was absent; 2), if it was present with another allele at that
locus; and 3), if it was present in the homozygous state. Of the 42 electrophoretic characters,
only a small proportion were significant to the scores on the first few principal component axes.
The APK locus was not used in PCA, because there was only one variant in 728 individuals.
An alternative method for scoring electrophoretic characters is used in some numerical
analyses (Mickevich and Mitter, 1981; Buth, 1984), with each locus a character, and each
allele coded on the basis of its relative mobility. This method was rejected because it does not
allow distinction between an individual homozygous for a particular allele, and an individual
heterozygous for two alleles located an equal distance on either side of the homozygous allele.
This situation occurred frequently at the G-6-PD locus.
Electrophoretic data were also analysed using the Biosys-1 package of Swofford and
Selander (1981). Allele frequencies, heterozygosity indices and tests for Hardy-Weinberg
proportions were calculated. All individuals with partial electrophoretic information were used,
giving a total sample size of 860.
Hardy-Weinberg equilibrium measures provided tests of gene pool homogeneity, and
complemented the multivariate clustering techniques. First, the entire sample for a region was
tested as a whole before being divided into major groups which might be different species. If the
subsets were much closer to equilibrium after the subdivision, this was considered evidence of a
significant degree of gene flow within but not between the subgroups. Excess homozygosity
may be due to other factors as well, such as the presence of null alleles or temporal and spatial
variation within samples. I consider these possibilities unlikely in the context of this study, since
the occurrence of excess homozygosity coincided fairly well with taxonomic expectation.
Taxonomic Interpretation and Conventions
I view species concepts as a balance between practicality and meaningfulness. Strict
definitions can be established, but they may not distinguish populations of biological relevance.
Alternatively, if a species concept is particularly vague or difficult to apply in practical
situations, then its potential biological meaningfulness is of little use. Both practicality and
meaningfulness should be assessed in terms of the reason for naming species, which is the
identification of organisms in a way that allows the user of the name to efficiently communicate
information about their relationships with other organisms.
Some taxonomists, for example Blackwelder (1967), view species recognition as a sort of
learned trade which cannot be precisely characterized or defined. Here species are kinds of
primary concepts. Understanding how to interpret certain kinds of information becomes a
matter of developing a sense of similarity relations and applying it in a manner sanctioned by
experienced peers. This part of taxonomic training thus involves the transfer among individuals
of a conceptual paradigm, in the sense of Kuhn (1970).
However, the uses of systematics are primarily scientific, and there is a need to make its
operations repeatable, quantifiable and testable. Most systematists have acknowledged that
species tend to be clusters of like individuals, and many of them have focussed on this aspect.
Even workers whose main research objectives are in elucidation of evolutionary mechanisms
may begin their discussion of species by referring to them as “discontinuous arrays”
(Dobzhansky et al. , 1977:166). However, some systematists treat species as phenetic covariance
Papilio machaon species group
217
clusters, and de-emphasize reference to reasons for the existence of these clusters. Examples
include Ehrlich (1961), who identifies species as relatively arbitrary groups of organisms
delineated by overall character similarity, or Neff and Platnick (1981:12), who characterize
species as “the smallest detected samples of self-perpetuating organisms that have unique sets
of characters”. This view appears strongly influenced by the desire to make the process of
distinguishing species as tractable as possible, particularly in terms of mechanical simplicity.
Other taxonomists emphasize the process perceived to maintain distinctions between species
and unity within species, perceiving gene exchange as the characteristic that makes such a
group of organisms a biologically coherent entity. A widely accepted definition of this type is
that of Mayr (1969), who defines species as “groups of interbreeding natural organisms that
are reproductively isolated from other such groups”. However, species descriptions based on
hybrid sterility are in some ways as arbitrary as species descriptions based on phenetic
clustering, since the degree of hybrid fertility which is accepted before a specific distinction is
recognized is itself an arbitrary procedure. One way of dealing with the latter objection has
been to characterize different populations as species only if they exhibit 100% hybrid sterility
( e.g ., Key, 1982). Unfortunately, this definition is so broad that many phenetically distinctive
groups of organisms presently recognized as belonging to different species would have to be
combined if the definition were rigorously applied.
The attempt to make the process of species recognition more objective thus seems to have
led in two major directions: 1), grouping by phenetic covariance; or 2), by interbreeding data.
However, grouping on the basis of either data type alone can lead to the absurd extreme of
operationalism, in which a definition is conceived as no more than a corresponding set of
operations (Hull, 1968). The main fault of operationalism is that it emphasizes practicality in
the application of a definition, but restricts the flexibility and general usefulness of the
definition.
The history of classification of the P. machaon group provides examples of both kinds of
operationalism. W.H. Edwards’ names are an example of an overemphasis on morphotypes,
relative to hybridization information. On the other hand, Hagen’s (1882) taxonomic
conclusions show an overly strict adherence to the interbreeding criterion of his species concept;
thus, he fell into the same trap of operationalism that reduced the value of Edwards’ work.
Considering the ecological and genetic complexity of the Papilio machaon group, one should
not expect sytematic research to have simple taxonomic consequences. I use numerical methods
in a predominantly descriptive manner, to characterize the pattern of phenetic variation within
and between populations. As in most taxonomic work, there is a need to distinguish variation at
the level of local populations, geographic races and species. Geographic patterns of variation
were first examined within major character suites, such as structural and electrophoretic
characters, and then compared among suites. Finally, these patterns of variation were loosely
interpreted in terms of current species concepts, especially through inferences of gene flow and
the maintenance of identity in time and space.
Recognition of subspecies is somewhat more arbitrary. My main criterion for formally
recognizing a differentiated series of populations or an ecological race as a subspecies is that
75% or more of the specimens can be distinguished without the aid of locality labels ( cf ’., Mayr,
1969). The main reason for using subspecies names is to relate my own findings to previous
work, much of which has been couched in terms of description of new specific and subspecific
taxa.
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218
Sperling
Since subspecies are preeminently geographic divisions of species, the former should be at
least parapatric, with intermediates occurring along only a relatively narrow zone. Phenetic
homogeneity within a subspecies should be quite high compared to that within zones of
intergradation. Although most subspecies have been described on the basis of structural
characters, ecological characters are also important indicators of substantial genetic
distinctions.
The main difficulty in classifying populations and individuals of the P. machaon group from
western Canada is that in some regions groups of individuals seem like genetically distinct
entities in sympatry, while in other areas extensive hybridization occurs. In general, this
difficulty is dealt with by continuing to recognize populations as separate species only if areas
of sympatry without hybridization are much more extensive than areas in which substantial
hybridization occurs. Examples in the P. machaon group are discussed in detail in the section
on diagnosis of adults and ranking of taxa.
Populations composed predominantly of hybrid individuals were given names which reflect
such hybrid origins. Since the International Code of Zoological Nomenclature (1985) does not
provide rules for hybrid names, I follow the International Code of Botanical Nomenclature
(1983), and more general guides in taxonomy such as Schenk and McMasters (1956).
However, the convention of ordering parental names by the sexes which contributed to the
hybrid swarms is not useful for the P. machaon group. Instead, the species epithets of the
parental species are listed, with an “X” between them, so the first epithet indicates the species
most similar to the majority of hybrid specimens.
I use form names as little as possible, because they have suffered from considerable
uneveness of usage ( e.g Scott, 1981) and have a limited communication value. A better
alternative would be to concentrate on distinguishing genes and alleles, as Clarke and Sheppard
did in the 1950’s with polymorphisms in the P. machaon group.
CLASSIFICATION AND RECOGNITION OF TAXA
The taxonomic conclusions of the present study are presented before the supporting data,
because I propose several changes to the systems of names which have been applied previously
to the Papilio machaon group. To simplify presentation of data in succeeding sections, my
system is summarized in the present section. Detailed reviews of characteristics of the included
taxa follow, and are discussed in succeeding sections, which deal with evolutionary hypotheses.
Summary of Taxonomic Assignments
The following list is based on Miller and Brown (1981), and summarizes the disposition of
all scientific names applying to the P. machaon group in western Canada and Alaska. It
includes type localities, as well as names applying to populations which are found, or have at
some time been considered to have been found, in the study area.
P. machaon Linnaeus, 1758:462. Type locality(TL)-Sweden.
a. P. m. aliaska S.H. Scudder, 1869:407. TL-Nulato, Alaska.
=joannisi R. Verity, 1 907 :pl. 1 0, Fig. 17. TL-Nulato, Alaska.
=petersii A.H. Clark, 1932:8-9. TL-Kuyukok River, Alaska.
b. P. m. bairdii W.H. Edwards, 1866:200. TL-“Arizona”, restricted to Fort Whipple,
Arizona, by Brown (1975).
Papilio machaon species group
219
= brucei W.H. Edwards, 1895:239. TL-“Colorado”, restricted to Glenwood
Springs, Colorado, by Brown (1975).
c. P. m. oregonius W.H. Edwards, 1876:208. TL-near The Dalles, Oregon
(neotype locality is at Hepner, Oregon [Brown, 1975]).
d. P. m. hudsonianus A.H. Clark, 1932:6-7. TL-Kettle Rapids, Manitoba.
e. P. m. dodi J. McDunnough, 1939:216-217. TL-Red Deer River, 50 miles N. E. of
Gleichen, Alberta (probably near Dorothy [Kondla, 1981]).
f. P. m. pikei F.A.H. Sperling. NEW SUBSPECIES. TL-Dunvegan, Alberta.
P. zelicaon Lucas, 1852:136. TL-“California”.
= nitra W.H. Edwards, 1883:162-163. TL-Judith Mts., Montana.
=gothica C.L. Remington, 1968:2-5. TL-Gothic, Colorado.
= ab. mcdunnoughi J.D. Gunder, 1928:162. TL-Waterton Lakes, Alberta.
P. zelicaon X machaon NEW HYBRID MORPH
P. polyxenes Fabricius, 1775:444. TL-“America”, restricted to Cuba by
Rothschild and Jordan (1906).
a. P. p. asterius Stoll, 1782:194. TL-New York, Virginia and Carolina.
P. machaon X polyxenes NEW HYBRID MORPH
= avinoffi F.H. and R.L. Chermock, 1937:1 1-12. TL-Whirlpool River,
Riding Mts., Manitoba.
P. polyxenes X machaon NEW HYBRID MORPH
= kahli F.H. and R.L. Chermock, 1937:12-13. TL-Riding Mts., Manitoba.
Papilio machaon pikei , new subspecies
Of the five major sections of P. machaon which occur in western Canada, four were
described many years ago. The fifth occurs exclusively within the Peace River region, and
appears to have been collected once (Llewllyn Jones, 1951) before being rediscovered by E.M.
Pike and me in 1980. The Peace River race of P. machaon is ecologically distinct from P. m.
aliaska and P. m. hudsonianus , and is geographically disjunct from P. m. dodi and P. m.
oregonius. Although very similar in morphometric and electrophoretic features to the other
subspecies of P. machaon in western Canada, it is as different from each of these as they are
from each other. In order to facilitate discussion about the evolution of this race, it is described
below. All measurements are based on specimens used in the principal components analyses in
the following section.
Description. — Adult (Plate 1, e-f). Male (Plate 1, e). Mean forewing length, from base at thorax to apex, 40.8
mm (range 36.5-47.0). Dorsal hindwing with yellow scales extended over proximal portion of wing almost to base, except
in cell Cu2, latter with yellow scales confined to postmedian region. Black pupil of dorsal hindwing eyespot along lower
edge of red scales and connected to margin of wing. Pupil club-shaped or narrow line. Blue and red scales of eyespot
separated by few or no black scales. Basal half of ventral forewing disc covered by yellow scales. Postmedian area of
ventral hindwing of most specimens with distinct patch of orange scales in two or fewer cells. Thorax covered by long
yellow hairs ventral to wings. Ventral side of abdomen with yellow hairs on all segments. Broad yellow lateral band on
abdomen, extended over claspers. Subdorsal spots above lateral abdominal band in few specimens. Female (Plate 1, f).
Like male, but larger (mean forewing length = 42.6 mm, range = 39.5-45.5), and with more rounded forewing.
Immatures. Eggs, larvae and pupae very similar in all stages to those of P. m. oregonius (see Perkins, et al., 1968 for
photographs) and to P. m. dodi. Fifth instar larvae with segmental spots orange or yellow, but most individuals with
orange spots. Background color of mature larvae from flat blue-green to bright emerald green. Pupae either mottled brown
or green, but not mixture of brown and green as in some specimens of P. m. aliaska. Larval foodplant Artemisia
dracunculus, on warm, dry, eroding exposures.
Type series. Abbreviations: f = female, m = male, Ad = Artemisia dracunculus , E4# = F.A.H. Sperling
electrophoresis number, CNC = Canadian National Collection. All reared specimens have pupal shell and some have fifth
Quaest. Ent., 1987,23 (2)
220
Sperling
instar larval skin attached to card below specimen. Pupation and emergence dates are omitted in the following list. Seventy
eight paratypes have been deposited in public institutions, and 20 remain in the personal collection of E.M. Pike. Sites of
deposition are listed in brackets behind each entry.
Holotype: male. Canada, Alberta; Dunvegan (s. Fairview); June 14, 1981; F.A.H. Sperling; [on dry, grassy, south-facing
slopes above Peace River] (CNC).
Allotype: female. Canada, Alberta; Dunvegan; June 22, 1982; F.A.H. Sperling; e4# 546 (CNC).
Paratypes. 2f, 6m: Canada, B.C.; Attachie; larva coll. Aug 9 ’81; F. Sperling; on Ad\ e4# 521-526,528,529 [all emg.
1983], (CNC). 2f, lm: Attachie, British Columbia; 35 km W. Ft. St. John; larva coll. Aug. 9, 1981; on Ad\ [all emg.
1982]; F. Sperling (CNC). lm: Attachie, British Columbia; 35 km W. Ft. St. John; larva coll. July 9, 1981; on Ad\ F.
Sperling (CNC). lm: Taylor, B.C.; July 3 ’80; F. Sperling (CNC). If: larva on Ad at Taylor, B.C. on Aug. 18, 1980; [emg.
1981]; F.A.H. Sperling; e4# 627, (CNC). If: Canada, B.C.; Taylor; larva coll. July 8 ’82; F. Sperling; on Ad\ e4# 439
(CNC). 3f, 6m: Canada, B.C.; Taylor; June 21, 1982; F. Sperling; including e4# 6,7,55,56,59,124,128,395; (British
Columbia Provincial Museum: e4# 59[f],124[m]. American Museum of Natural History: e4# 6[m],55[f]. United States
National Museum: e4# 56[m], 1 28 [f] . remainder to CNC). If, 2m: Canada, B.C.; Taylor; July 8, 1982; F. Sperling; e4#
10,31,140; (CNC: e4# 31 [m]. British Museum [Natural History]: e4# 10[m],140[f]). 2f: Clayhurst Ferry, B.C.; larva on
Ad\ Aug. 17 ’80; emg. 1981; F.A.H. Sperling; including e4# 117; (CNC: e4# 117 [f] . Allyn Museum of Entomology: If).
3m: Clayhurst Ferry, British Columbia; larva coll. Aug. 9, 1981; on Ad\ emg. 1982[2] & 1983 [1]; F.A.H. Sperling;
including e4# 519, (CNC). 4f, 7m: Canada, B.C.; Clayhurst Ferry; larva coll. Aug. 16 ’82; F. Sperling; on Ad-, emg. 1983;
e4# 504,507,508,511-513,515-518,520, (Allyn Museum of Entomology: e4# 507 [m]. remainder to CNC). 5m: Alberta, 5
km NW Highland Park; June 14, 1981; F. Sperling; e4# 115,116,676,678,679, (CNC). 5m: Canada, Alberta; Highland
Park; 35 km w Fairview; June 9, 1982; F.A.H. Sperling; e4# 41,42,43,44,45, (Alberta Provincial Museum: e4# 41,43.
CNC: e4# 42,44,45). 9m: Canada, Alberta; Highland Park; 20 mi. W. Fairview; June 12, 1982; Ted Pike, (CNC). 5m:
Canada, Alberta; Highland Park; 20 mi. W. Fairview; June 13, 1982; Ted Pike, (Pike), lm: Dunvegan, Alberta; larva on
Ad-, Aug. 16, 1980; emg. 1981; F.A.H. Sperling, (CNC). If: Canada, Alberta; Dunvegan; June 14, 1981; F.A.H. Sperling,
(University of Alberta Strickland Museum). If, 2m: Alberta, Dunvegan; June 14, 1981. T. Pike, (Pike). If: Canada,
Alberta; Dunvegan; larva coll. Aug. 15, 1982; F.A.H. Sperling; on Ad\ emg. 1983; e4# 438, (CNC). 3m: Canada, Alberta;
Dunvegan; June 22, 1982; F.A.H. Sperling; e4# 142,150,547, (University of Alberta Strickland Museum: e4# 150[m].
remainder to CNC). If: Dunvegan, Alta.; 30 VI 85; coll, by E.M. Pike; (Pike), lm: EX OVA; Dunvegan, Alta.; 18 VI 85;
coll, by E.M. Pike; (Pike). 4m: 10 mi. S.E. Fairview; Alberta; 17 June 1981; coll, by E.M. Pike; (Pike). 2m: 10 mi. S.E.
Fairview; Alberta; 20 & 22 June 1981; coll, by E.M. Pike; (Pike). If, 2m: Canada, Alberta; 10 mi. S.E. Fairview; June 22,
1982; Ted Pike; (Pike). If: 10 mi. S.E. Fairview; Alta., 20 VI 85; coll, by E.M. Pike, (Pike), lm: Canada, Alberta; Peace
R. area, Camp Island; 22 mi. E. Dunvegan; F.A.H. Sperling; larva on Ad on Aug. 15, 1980; [emg. 1981], (CNC). If: larva
on Ad\ at Peace R. (town), Alberta; on Aug. 15, 1980; emg. 1981; F.A.H. Sperling; e4# 625, (CNC). 4m: Alberta, Peace
River (town); June 10, 1981; F. Sperling (CNC). 2m: Canada, Alberta; Peace River (town); June 13, 1981; F.A.H.
Sperling; e4# 114,681, (CNC). lm: Canada, Alberta; Kleskun Hills; 25 km n.e. Grande Prairie; June 19, 1982; F.A.H.
Sperling; e4# 394, (CNC). If: Canada, Alberta; Kleskun Hills; e. Grande Prairie; larva coll. Aug. 12 ’81; F.A.H.
Sperling; on Ad\ emg. 1983; e4# 437, (CNC).
Derivation of subspecific epithet. — It is a pleasure to name this subspecies after E.M. (Ted)
Pike, who has resided in the Peace River region from 1979 to 1985, and has given me much
help and encouragement during the past 15 years.
Distinguishing features. — Approximately 75% of individuals of P. m. pikei can be correctly
distinguished from those of other subspecies of P. machaon. Features which distinguish this
subspecies are discussed at greater length in the following chapters. Adults of P. m. pikei
resemble those of P. m. oregonius in general maculation and size, but most are distinguished by
the more narrow, connected eyespot. P. m. aliaska adults resemble those of P. m. pikei
markedly in maculation, but are separated by larval foodplant, preference for alpine habitat,
and smaller size (mean forewing length = 37.5 mm for males, 40.3 mm for females). Though
the range of P. m. pikei extends in isolated populations to within 25 km of P. m. aliaska , at
Hudson Hope, there is no evidence of any increased similarity of these two subspecies in the
area. P. m. hudsonianus adults are separated by preference for boreal forest habitats and a
much higher frequency of subdorsal abdominal spots. P. m. dodi is easily distinguished from P.
m. pikei by the greater amount of black scales and hairs, especially on the ventral forewing disc
and ventral side of the thorax.
P. m. pikei has the same larval foodplant as the southern subspecies of P. machaon , but
shares several morphometric and electrophoretic similarities with the northern subspecies. For
these reasons, as well as its geographic range, P. m. pikei is important in illustrating the
Papilio machaon species group
221
previously unrecognized link between these taxa.
Range. — P. m. pikei is composed of a series of populations distributed along approximately
500 km of the Peace River, in northeastern British Columbia and northwestern Alberta. It also
occurs at the Kleskun Hills badlands, northeast of Grande Prairie, Alberta. The range of P. m.
pikei may have once extended farther westward along the Peace River. A specimen in the
University of British Columbia collection, which is labelled “Findlay, B.C.”, may be from
Findlay Forks, 110 km west of Hudson Hope. However, the populations along this part of the
Peace River may now be extinct, since it was flooded to form Williston Lake in the late 1960’s.
Key to Adults of P. machaon Group in Western Canada
The following key is based on color pattern and habitat information. Electrophoretic
characters are not used.
1 Black scales on disc of dorsal hindwing (DHW) restricted to basal half
(Fig. 2); side of abdomen with broad, yellow, longitudinal band and in
some specimens with rounded spots above it (Figs. 6a-d) 2
V Black scales extended over more than half of DHW disc (Fig. 3); side of
abdomen with only series of square or rounded segmental yellow spots
(Figs. 6e-f) 9
2 (1) All of the following character states : Black pupil in anal region of DHW
connected to margin (Figs. la-b,2,3); yellow scales over most of ventral
forewing (VFW) disc (Fig. 5A); yellow hairs extended around ventral part
of metathorax (Fig. 6a); yellow scales extended over more than 80% of
male claspers (Fig. 6a).
Or no more than one of following : Anal pupil unconnected to margin but
flattened and at bottom of red area (Fig. 1C); yellow scales in VFW disc
restricted to thick yellow streaks or general flush extended over more than
quarter of disc (Fig. 5B); yellow scales extended over only 50-80% of male
claspers (Fig. 6b) P. machaon Linneaus (Plate 1) 4
2' Not as above 3
3 (2') All of following : Anal pupil round and centered in red area (Fig. ID); red
and blue areas of anal eyespot more than 3/4 separated by black scales
(Figs, lb, Id, 2, 3); disc of VFW with at most few thin streaks or light
sprinkling of yellow scales (Figs. 5c-e); metathorax with yellow hairs from
both sides not in contact ventrally (Figs. 6b-d); without distinct yellow
spots above lateral abdominal band (Figs. 6b-c); less than 50% of male
claspers covered by yellow scales (Figs. 6c-d).
Or no more than one of following : Anal pupil large, round and centered if
connected to margin (Fig. 2) or small and oval at bottom of red area if
unconnected (Fig. 1C); red and blue areas of anal eyespot separated
between 1 /4 and 3/4 of full width by black scales (Fig. 1C); disc of VFW
with thick streaks of yellow or a general flush over less than 1 /4 of the disc
(Fig. 5B); some distinct yellow spots above lateral abdominal band (Fig.
6d); yellow scales over 50-80% of male claspers (Fig. 6b)
P. zelicaon Lucas (Plate 2g-h)
y Not as above: most specimens with club-shaped pupil connected to margin;
Quaest. Ent., 1987,23 (2)
222
Sperling
4 (2)
4'
5 (4)
5'
6 (4')
6'
7 (30
also most specimens with two or more of character states intermediate
between extremes of P. machaon and P. zelicaon as defined above, rather
than combination of extreme states 7
Found near dry grasslands or eroded clay banks in hot habitats; large (FW
length usually 40 mm or more in males); forewing apices pointed or not,
with distal margin convex or concave; most specimens with yellow scaling
of DHW anal cell Cu2 extended close to or beyond divergence of veins Cul
and Cu2 (Fig. 2; character states 2-4); few specimens with abdomen with
spots above lateral band 5
Found in forested boreal regions or on alpine tundra in cool habitats;
smaller (FW length usually less than 40 mm in males); forewing apices of
most ‘Specimens rounded, with convex outer margin (Fig. 5A); yellow
scaling of DHW anal cell Cu2 in most specimens restricted to distal 1 /4
(Fig. 2: character state 1); abdomen with or without yellow spots above
lateral band 6
Found in southern and central B.C. Interior, during April to September;
anal pupil of eyespot connected to margin in most specimens, but
club-shaped rather than flat line; separation between blue and red areas of
anal eyespot various; with substantial amount of orange in two or more
cells of VHW postmedian band; most specimens with forewing apices
pointed, with concave distal margin; (Note: a few summer generation P. m.
dodi from the southern Alberta and Saskatchewan prairies key out here) .
P. machaon oregonius (Edwards) (Plate lg-h)
Found in Peace River region of northeastern B.C. and northwestern
Alberta, during June and early July; anal pupil of many specimens flat
rather than club shaped; most specimens with very little black separation
between red and blue in anal eyespot; most specimens with substantial
amounts of orange in only one or no cell of the VHW postmedian band;
forewing apices pointed or rounded
P. machaon pikei Sperling (Plate le-f)
Found in Alaska, Yukon, western Northwest Territories, and northern
British Columbia, most specimens on alpine tundra; DHW anal pupil in
form of thin line, at bottom of red area, and connected to margin (Fig. 1 A);
red and blue areas of anal eyespot with no or very little black separation
(Fig. 1 A); no spots or in few specimens one or two spots on abdomen above
lateral band P. machaon aliaska Scudder (Plate la-b)
Found in boreal forest from Alberta to northern Quebec; DHW anal pupil
in most specimens club shaped; red and blue areas of anal eyespot
separated or not by black scales; at least one yellow spot above lateral
abdominal band in most specimens
P. machaon hudsonianus Clark (Plate lc-d)
Found near dry grassland or eroding clay banks in hot prairie habitats of
southern Alberta or Saskatchewan; anal pupil of DHW club-shaped and
connected to margin; forewing apex of many specimens pointed, with
concave distal margin; hindwing tails of many specimens long, slightly
narrowed in middle and curved (Fig. 2); yellow scales in DHW anal cell
Papilio machaon species group
223
8(7')
8'
9 (10
9'
10 (9)
Cu2 extended or not beyond divergence of veins Cul and Cu2 (Fig. 2:
character states 3-4); no distinctly separated yellow spots above lateral
abdominal band P. machaon dodi McDunnough (Plate 2a-b)
Found in broad range of habitats, but most in southern zones of boreal
forest; anal pupil of DHW varied; forewing apex of most specimens
rounded, and distal margin straight or rounded; hindwing tails of medium
or short length, straight and not constricted in middle (Fig. 3); yellow
scales of distal hindwing cell Cu2 in most specimens restricted to distal
quarter; less than five yellow spots above lateral abdominal band
yellow morph hybrids 8
Found in Manitoba or eastern Saskatchewan and with one or more of the
following character states: anal pupil on DHW round and centered; disc of
VFW with at most light sprinkling or thin streaks of yellow scales; thorax
with yellow hair not in contact ventrally and no yellow hairs on ventral
midline of first two abdominal segments; male claspers covered over less
than 50% of surface by yellow scales P. machaon X polyxenes
Found in western Saskatchewan and westward, with any of the following
combinations:
A. In predominantly forested habitats and with club shaped, connected
pupil.
or B. Specimen with between two and five of the following six character
states: 1, DHW anal pupil connected to margin or unconnected, flattened
and at bottom of red area; 2, Blue and red areas of anal eyespot not
separated by black along at least 1/4 of boundary; 3, disc of VFW with
yellow scales or thick streaks over at least 1 /4 of area; 4, metathorax with
yellow hair meeting ventrally; 5, one to five distinct spots above lateral
abdominal band; 6, yellow scales over more than 50% of male claspers
P. zelicaon X machaon (Plate 2c-f)
Found in southern and central Manitoba or southeastern Saskatchewan;
postmedian band of VHW with substantial amounts of orange in at least
two cells, and all cells in most specimens; distinct yellow spots in two
subdorsal rows on at least five and in most specimens on all segments of
abdomen 10
Found in southwestern Saskatchewan and south or central Alberta;
postmedian band of VHW with substantial amounts of orange in less than
six cells (only two or three in most specimens); distinct yellow spots in
subdorsal position on abdomen usually absent on at least 2 segments 11
Anal pupil of DHW unconnected to margin or club shaped if connected;
blue and red areas of anal eyespot fully separated by band of black scales;
less than half of hairs on tegula yellow; yellow scales in apical cell of
postmedian band of VFW varied; postmedian band of VHW with orange in
all eight cells; lower half of side of abdomen with rounded yellow spot on
each abdominal segment; yellow spots in subdorsal position on abdomen
absent on no more than two segments; less than 10% of male claspers
covered by yellow scales; females with markedly reduced postmedian band
on DHW, compared to males
P. polyxenes asterius Stoll (Plate 3g-h)
Quaest. Ent., 1987, 23 (2)
Sperling
All three of following character states : anal pupil club-shaped and
connected to margin; more than 50% of hairs on tegula yellow; apical cell
of VFW postmedian band with distinct patch of yellow, but occupying less
than half of cell area;
or one or more of the following character states ; anal pupil thin line at
lower edge of red area, connected to margin; blue and red areas of anal
eyespot not completely separated by black scales; apical cell of VFW
postmedian band more than 50% covered by yellow scales; postmedian
band of VHW with no orange in at least one cell; large square spots or
broad band of yellow along lower half of abdomen; yellow spots in
subdorsal position on abdomen absent from at least three segments; more
than 10% of male claspers covered by yellow scales; females with
postmedian band on DHW same width as on males
P. polyxenes X machaon (Plate 3d)
Anal pupil of DHW round and centered in red area; blue and red areas of
anal eyespot fully separated by black scales; male claspers with less than
10% yellow scales black morph of P. zelicaon (Plate 3a)
One or more of following character states; anal pupil of DHW connected to
margin or low and oval if unconnected; blue and red areas of anal eyespot
of DHW not separated by black scales along at least 1 /4 of boundary; male
claspers with more than 10% yellow scales 12
12 (IT) Found near dry grassland or eroding clay banks in hot prairie habitats . . .
black morph of P. machaon dodi
12' Found in predominantly forested habitats
black morph of P. zelicaon X machaon (Plate 3b-c)
MORPHOMETRIC AND ELECTROPHORETIC CHARACTERS
Characters of Adults
Only a few species within the Papilio machaon group are easy to distinguish on the basis of
morphometric characters. The most divergent of these is P. alexanor Papilio, which has a
striped wing pattern and male genitalia unlike the other species of the group (Higgins, 1975),
but shares with them an apotypic (derived) larval color pattern and larval foodplant. Diagnostic
interspecific distinctions in genitalia are also found in P. indra and P. hospiton. The remaining
five species in the P. machaon group are much more similar with respect to adult characters.
The character states traditionally used by systematists to distinguish among P. machaon , P.
zelicaon and P. polyxenes are especially difficult to employ, because the variation in color
pattern in any one species is paralleled by the other species in other areas. Also, virtually no
character states stand on their own, without consideration in combination with other
characters. For this reason, I use multivariate statistical methods to provide more reliability in
clustering groups of similar individuals, both at the level of populations and species. As well,
two character suites were surveyed and compared: one, the traditionally employed color pattern
data; and two, new information about enzyme alleles.
Cluster resolution with principal components analysis. — Three principal components
analyses (PCAs) were applied to the same 728 individuals using, respectively, morphometric
data, electrophoretic data, and both data sets combined. All PCAs gave generally similar
orientations of locality samples (Figures 7 and 8).
224
10'
11 (90
ir
Papilio machaon species group
225
These samples were then compared with samples from or near the type localities of named
populations, which were scored with factor loadings derived from analysis on morphometric
data alone (Figure 9). From this comparison, it was clear that in all three principal components
analyses the first axis separated most yellow morph populations of Papilio machaon (No.
1,2,3,5,12,13,15) from Papilio zelicaon (No. 8,9, 10, 16, 17a, 18, 19), the second axis separated
Papilio polyxenes (No. 11, 17b, 20) from the previous two groups and the third axis provided a
partial separation of the P. machaon cluster. Factor loadings for all three PCAs are included in
Tables 12 and 13.
Electrophoretic characters showed a close association between (P. m. dodi Figure 7, No. 5)
and other P. machaon subspecies, while morphometric characters (Figure 8 and 9) indicated a
more intermediate position for P. m. dodi between P. zelicaon and other P. machaon
subspecies. Populations from the Alberta foothills, such as Bragg Creek, were intermediate in
both electrophoretic and morphometric characters. P. m. oregonius populations, which have not
previously been associated with P. machaon in most publications, showed a close association
with P. machaon on the basis of both character suites.
Although the second axis of each of the three PCAs served to separate P. polyxenes from
both P. machaon and P. zelicaon , the black morphs associated with populations of
predominantly yellow individuals were placed in an intermediate position between them in
those analyses which included morphometric data. Electrophoretic characters showed a much
closer association between the black and yellow morphs of most populations. The sample size
from central Manitoba (No. 14) was relatively small; nonetheless, the single yellow morph
specimen showed a close association with P. machaon for both character types. The black
samples from central Manitoba showed a somewhat closer association with P. machaon than
with P. polyxenes on the basis of electrophoretic characters, and grouped closely with P.
polyxenes in morphometric characters.
Although plotting entire population samples on the principal component axes served to
group most of these with either P. machaon , P. zelicaon or P. polyxenes , the associations of a
number of intermediate samples were uncertain. In particular, this procedure did not
distinguish between samples which were intermediate because the whole population was
intermediate, and samples which contained a mixture of individuals of more than one of the
above species. To facilitate such a distinction, the scores of all the individuals within a region or
at a locality were plotted as frequency histograms on principal component axes (Figures
12-13). Since there seemed to be regional trends with respect to the frequency of intermediate
individuals, the total sample used in the original PCAs was divided into five major regions
(Figure 10).
Both morphometric and electrophoretic characters provided a good separation of P.
machaon from P. zelicaon in southern and central British Columbia, as well as in the Peace
River region (Figure 12). The few specimens which were intermediate on the basis of either
character type grouped with P. zelicaon when both character types were considered together.
The sample from southern Alberta and Saskatchewan showed a reasonable degree of clustering
on the basis of electrophoretic but not morphometric characters, and the low frequency section
between these two clusters was shifted toward P. zelicaon , with both character suites
considered simultaneously.
With samples from the southern Alberta and Saskatchewan region considered separately, it
was clear that the frequencies of P. machaon and P. zelicaon differed markedly by locality
(Figure 1 1 and 15). Specimens collected along high river banks were likely to be more similar
Quaest. Ent., 1987, 23 (2)
226
Sperling
Plate 1 . Color pattern of P. machaon subspecies. Each specimen has dorsal and ventral views of right wings figured on
right and left sides, respectively: a, P. m. aliaska, male Mi. 391, Alaska Hwy., British Columbia, July 2, 1972; b, P. m.
aliaska, female Pink Mountain, British Columbia, Ex larva, coll. 17 Aug. 1982,. on Artemisia arctica; c, P. m.
hudsonianus , male Thompson, Manitoba, July 2, 1983; d, P. m. hudsonianus, female 25 km NE of The Pas, Manitoba,
June 14, 1980; e, P. m. pikei, male 7 km NE of Hudson Hope, British Columbia, Ex larva, coll. 20 Aug. 1984, on
Artemisia dracunculus ; f, P. m. pikei, female Dunvegan, Alberta, June 14, 1981; g, P. m. oregonius, male Kamloops,
British Columbia, Ex larva, coll. 27 Aug. 1983, on Artemisia dracunculus-, h. P. m. oregonius, female Kamloops, British
Columbia, Ex larva, coll. 27 Aug. 1983, on Artemisia dracunculus.
Papilio machaon species group
227
Quaest. Ent., 1987, 23 (2)
228
Sperling
Plate 2. Color pattern of P. machaon, P. zelicaon and hybrids Each specimen has dorsal and ventral views of right wings
figured on right and left sides, respectively. Figure h is reversed and shows left wings: a, P. m. dodi , male 1 1 mi. N of
Taber, Alberta, Ex larva, coll. 19 Aug. 1981, on Artemisia dracunculus\ b, P. m. dodi , female Drumheller, Alberta, Ex
larva, coll. 22 July 1981, on Artemisia dracunculus\ c, P. zelicaon X machaon , male Bragg Creek, Alberta, Ex larva, coll.
18 July -7 Aug. 1982, on Zizia aptera\ d, P. zelicaon X machaon , female Bragg Creek, Alberta, Ex larva, coll. 18 July -7
Aug. 1982, on Zizia aptera\ e, P. zelicaon X machaon , male Bragg Creek, Alberta, May 31, 1980; f, P. zelicaon X
machaon , male Bragg Creek, Alberta, May 31, 1980; g, P. zelicaon, male Wintering Hills, 18 km S Drumheller, Alberta.
May 24, 1982; h, P. zelicaon, female Waterton Park, Alberta. Ex larva, coll. 19 Aug. 1981, on Lomatium dissectum.
Papilio machaon species group
229
Quaest. Ent., 1987, 23 (2)
230
Sperling
Plate 3. Color pattern of dark morphs of P. machaon group species. Each specimen has dorsal and ventral views of right
wings figured on right and left sides, respectively: a, P. zelicaon, male Wintering Hills, 18 km S of Drumheller, Alberta,
May 30, 1982; b, P. zelicaon X machaon , male Bragg Creek, Alberta, Ex larva, coll. 15 July -7 Aug. 1982, on Zizia
aptera\ c, P. zelicaon X machaon , male Bragg Creek, Alberta, June 23, 1974; d, P. polyxenes X machaon , female Duck
Mountain Park, Manitoba, Ex larva, coll. 25 June 1980, on Zizia aptera\ e, P. m. bairdii , male Sunset Crater, E of
Flagstaff, Arizona, May 5, 1980; f, P. m. bairdii, female Flagstaff, Arizona, May 27, 1980; g, P. p. asterius, male
Burlington, Ontario, Ex larva, coll. Aug. 1981, on garden carrot; h, P. p. asterius, female Burlington, Ontario, Ex larva,
coll. Aug. 1 98 1 , on garden carrot.
Papilio machaon species group
231
Quaest. Ent., 1987, 23 (2)
232
Sperling
Figures 7 to 9. Mean scores of representative populations plotted on first three principal component axes. See Table 4 for
key to locations. Black circles indicate black morph adults, and empty circles indicate yellow morph adults. PC 1, PC 2
and PC 3 refer to the first, second and third principal component axes. Figure 7. 3D.PCA on electrophoretic data alone
Populations include only individuals used in the original analysis. Figure 8. 3D.PCA on morphometric data alone
Populations include only individuals used in the original analysis. Figure 9. Additional samples scored with morphometric
loadings. Populations are partly or completely composed of individuals not included in original analysis, but scored with
factor loadings from PCA on morphometric data alone. Most populations are either topotypic or from close to type
localities.
Papilio machaon species group
233
Table 4. Population samples of the P. machaon species group used in Figures 7-9.
Letter after taxon name indicates yellow color morph (Y) or black color morph (B).
(continued on next page)
Quaest. Ent., 1987, 23 (2)
234
Sperling
to P. machaon from other regions, especially in electrophoretic characters. Specimens from
prairie hilltops were more likely to belong to P. zelicaon. This situation was well illustrated by
the locality samples from the Drumheller region. The sample from the river bank just above the
town of Drumheller contained only P. m. dodi and a few intermediate specimens. There were
mostly P. zelicaon in the samples from the Hand Hills and the western part of the Wintering
Hills, which are about 15 and 12 km, respectively, from the nearest deeply cut river valleys or
ravines. A mixture of both species was at a hilltop on the eastern part of the Wintering Hills,
about 4 km from the nearest deep ravine and 6 km from the banks of the Red Deer River.
This pattern was basically the same as that in southern British Columbia, where P. m.
oregonius lives in the dry grassland habitats of the central Interior, while P. zelicaon is far
more common in forested and wetter habitats. In the Peace River region P. m. pikei also tended
to occur on the dry river banks and P. zelicaon on the hills farther away from the river.
However, an added complication is that another subspecies, P. m. aliaska, frequents the boreal
and especially the alpine regions of northern British Columbia, north of the Peace River.
In the predominantly forested regions of central Alberta, there seemed to be a different sort
of relationship between P. machaon and P. zelicaon (Figure 11 and 14). At the northern
localities (Marten Mt. to Adams Lkt.) there was a predominance of electrophoretic and
morphometric character combinations which tended to resemble P. zelicaon , as well as a
significant proportion of more intermediate individuals. However, a few individuals were
indistinguishable from northern P. machaon even with the two character types considered
together, and it was unclear whether these formed a distinctive group from the others. In the
more southerly localities (Buck Lk. to Bragg Cr.), the different phenotypes evident in the north
tended to merge even more. Populations from single localities were composed of a few
individuals indistinguishable from either P. machaon or P. zelicaon , but the majority was
intermediate. There was a single peak near the midpoint between the two extremes, which
tapered off to either side. The different phenotypes all occurred within the same habitat as well.
Though electrophoretic and morphometric characters showed a generally concordant pattern,
electrophoretic character combinations were more obviously intermediate than were
morphometric characters.
Papilio machaon species group
235
10
Figures 10 and 11. Figure 10. Western Canada, showing 5 major regions: 1, southern and central British Columbia
(s+cent. BC); 2, Peace River region (Peace R.); 3, central Alberta (cent. AB); 4, southern Alberta and Saskatchewan (s
AB + SK); 5, Manitoba and eastward (MB + east). Dots show localities from which specimens were used in the initial
PCAs. Figure 11. Central and southern Alberta, with major localities. Localities refer to those used in figures 14 and 15.
Quaest. Ent., 1987, 23 (2)
236
Sperling
s ♦ cen t . BC
Peace R.
central AB
s AB ♦ SK
M B ♦ ea st
Figures 12 and 13. Component axes of three separate PCAs, with frequency histograms of all individuals in each of five
major geographic regions (see Figure 10). Only specimens used in original PCAs are included. Darkened portions of
histograms indicate black morphs. Dashed lines indicate divisions between taxa. Mo. = morphometic characters. E4 =
electrophoretic characters. Figure 12. First component axes, by major region Figure 13. Second component axes, by major
region
Papilio machaon species group
237
35
96
(8)
16
(5)
f3.0
-0.5
E4
-3.5
14
26
21
6
6
73
(3)
5
6
74
(7)
Tolman Br.
Hand Hills
Drumhe Her
Wint. Hills - E
Wint. Hills - W
Out look
Taber
Waterton
N
6
9
(1 )
85
(2)
21
(1)
21
(3)
13
20
16
Mo. E4
15
Figures 14 and 15. Locality samples plotted on first component axes. Some samples include individuals not used in original
PCAs. Darkened parts of histograms indicate black morphs. Dashed lines indicate divisions between taxa. Mo. =
morphometric characters. E4 = electrophoretic characters. Figure 14. Central Alberta samples plotted on PC.l. Figure
15. Southern Alberta and Saskatchewan samples on PC.l.
Quaest. Ent., 1987, 23 (2)
238
Sperling
n M B + SK
Duck Mt. to
R iding Mt. Pk.
s MB + n ND
ON + WN
50%
£
■ ■■
i II
J79
(74) r—
39
(7!
i
21
►1.6 -1.2 -3.2 -2.4 0.0 +2.8
M o. E 4
16
I A. dracunculus
0 A.arctica
□ Umbeltiferae
17
Figures 16 and 17. Dashed lines indicate divisions between taxa. Mo.= morphometric characters. E4 = electrophoretic
characters. Figure 16. Area samples from Manitoba and eastward. Samples are plotted on second component axes, and
some include individuals not included in original PCAs. Black morph individuals are indicated by darkened portions of
histograms. Geographic areas: n MB + SK = northern Manitoba and northern Saskatchewan; Duck Mt. to Riding Mt.
Pk. = Duck Mountain and Riding Mountain Parks; s MB + n ND = southern Manitoba and northern North Dakota;
ON + WN = Ontario and Wisconsin. Figure 17. Adults reared from wild-collected larvae Histogram key: darkened =
larvae on Artemisia dracunculus-, cross-hatched = on Artemisia arctica, clear = on umbelliferous larval hostplants.
Individuals are plotted on first component axis of PCA on combined character set and include only those used in original
PCA. Regions refer to those indicated in Figure 10.
Papilio machaon species group
239
Figures 18 and 19. Reared adults plotted on discriminant axes, including only those used in original discriminant analyses.
Numbers indicate more than one data point with the same coordinates. Circles indicate maximum diameters of the three
original groups. Dashed lines indicate locations of populations from opposite figure. Figure 18. 2D.DFA plot of reared
samples: southern Alberta. On A. dracunculus ( + ) N = 1 1 9; on umbellifers (triangles) N = 22. Figure 1 9. 2D.DFA plot of
reared samples: central Alberta. Only umbellifer-reared individuals shown, N = 45.
Quaest. Ent., 1987,23 (2)
240
Sperling
251
20-
10 -
0 -
20“l
| on Umbelli ferae
0 on A. d r ac u n c u I u s
□ larval foodplant
unknown
20
io -
0-f=h
- 5.0
central AB
D i sc r. Axis 1
2.4
♦5.0
21
Figures 20 and 21. All adults from central and southern Alberta (AB), and southern Saskatchewan (SK), plotted on first
discriminant axis. Reared individuals are indicated as subsets. Figure 20. All southern Alberta and Saskatchewan
specimens, plotted on DFA.l N = 497, including N= 1 19 reared from A. dracunculus and N = 22 reared from umbellifers.
Figure 21. All central Alberta specimens, plotted on DFA.l N=481, including N = 7 reared from A. dracunculus and
N = 45 reared from umbellifers.
The most P. machaon-Yike individuals from the southern foothills of Alberta had
morphometric character combinations more similar to P. m. hudsonianus than to P. m. dodi. In
fact, no specimens were collected on the southern prairies which were as P. m. hudsonianus or
P. m. aliaska- like as a few individuals taken at Bragg Creek and Buck Lake, localities
geographically close to P. m. dodi populations. This suggests that the hybrid populations are at
least partly a product of hybridization between P. zelicaon and the more northerly P. machaon
races, rather than with P. m. dodi. On the other hand, most of the specimens from these two
localities tended to be very similar to P. m. dodi in morphometric characters, while most were
more intermediate between P. zelicaon and P. machaon in electrophoretic characters. This
suggests that the very similar wing and body color pattern combinations which occur in both P.
m. dodi and the hybrid swarms may have arisen in different ways.
In Alberta, the morph with black wings occurred together with the more common yellow
members of both P. machaon and P. zelicaon. The former specimens had a range of
electrophoretic character combinations matching the remainder of the population with which
they occurred. This applied to individuals collected with other P. zelicaon specimens on prairie
hilltops, the intermediate hybrid populations of the Alberta foothills, and the P. m. dodi
collected along dry river banks. On this basis it appears as though this morph has become a
Papilio machaon species group
241
regular part of all of these populations. Although the color pattern of the black morph in many
respects resembles that of P. p. asterius, there was no good electrophoretic evidence of
hybridization with P. polyxenes in these populations.
The situation in Manitoba was far less clear (Figure 16), though there probably are hybrid
swarm populations in this region as well. These hybrid populations appear to be the result of
interactions between P. machaon and P. polyxenes , rather than between P. machaon and P.
zelicaon as in central Alberta. The dramatic effect of the gene for the black wing morph made
it more difficult to demonstrate phenotypic intermediacy in morphometric characters, and I
was able to subject only a small number of individuals from central Manitoba to electrophoretic
analysis. However, populations scored on the second PC axis tended to be intermediate in
electrophoretic characters. Also a sizable proportion of black morph individuals in central
Manitoba tended to take on character states found in P. machaon. For example, most had a
club-shaped anal pupil and many had more yellow on the tegulae and apical forewing cell than
in P. p. asterius from southern Ontario or the United States.
Adult characters versus larval foodplants. — Separation of P. machaon from P. zelicaon
and P. polyxenes on the basis of electrophoretic and morphometric characters was supported by
a comparison of larval foodplants. Scores of adults reared from larvae collected on Artemisia
were plotted against those of individuals from various species of Umbelliferae, on the first PC
axis derived from both the electrophoretic and the morphometric characters combined (Figure
17). In the Peace River region, individuals reared on Artemisia arctica from alpine habitats
grouped with those reared on Artemisia dracunculus from dry, grassy river banks. The P.
machaon from southern Alberta and Saskatchewan, reared on A. dracunculus , were also
separated from P. zelicaon on this basis, although their more similar morphometric characters
resulted in a somewhat closer grouping. The single P. machaon-likc individual reared from
central Manitoba was obtained on an umbellifer.
The specimens reared on A. dracunculus from central Alberta were collected at Nevis
Junction, on a northward extension of prairie habitat along the Red Deer River. These adults
resembled those of P. m. dodi from further south along the river and were undoubtedly just an
outlying population of this race. They also, however, resembled some specimens collected on
umbellifers farther to the west. Larvae of the hybrid populations from central Alberta feed on
umbellifers, and in this respect are similar to P. zelicaon. Reared material showed the same
wide range of phenotypes as the wild- collected adults.
Since PCAs on the total sample from western Canada provided only a partial separation
between P. machaon and P. zelicaon from central and southern Alberta on the basis of
morphometric characters, I attempted to improve the separation with discriminant factor
analysis (DFA) of foodplant groups. Three foodplant groups were defined. The first included
all specimens reared from A. dracunculus in either southern or central Alberta. The second
included all material from umbellifers in the southern Alberta region, all of which were from
Angelica , Lomatium or Heracleum in the Waterton Park and Crowsnest Pass area. The third
group included all the adults reared from umbellifers in the central Alberta region. The
morphometric characters used in this analysis were the same as those used in the PCAs, except
that only 10 characters were used because one character (tegula color) showed no variation in
the groups defined above. The discriminant axis loadings are included in Table 12.
As with the PCAs on electrophoretic characters, the DFA gave a fairly good separation of P.
machaon and P. zelicaon in southern Alberta (Figures 18-19). However, the umbellifer feeders
from central Alberta did not separate very well from either of the other two groups. With all
Quaest. Ent., 1987, 23 (2)
242
Sperling
Peace R. region ~ °.
o m ^
I IZL r I ■ —
P. machaon
P. machaon oregonius
R zelicaon
al pi ne - P. m. alia ska
>. K
-O cs
>
• - o
P. zelicaon
parkland - P. m.pi kei
central AB
P, machaon
P . zelicaon
P. machaon
P. zelicaon
on A. dracunculus
H *
on U m be 1 1 i ferae
s of H w y. 1 A
P. machaon
P . zelicaon
P. machaon
P. zelicaon
P. mach aon
P. polyxenes
Thompson
central Mani toba
Wisconsi n + Ontario
Figure 22. Hardy-Weinberg equilibrium tests on subpopulations. Six loci are shown as squares within bar for each
population. Significance levels (key at center left) refer to deviation from equilibrium. Names for some populations refer to
population divisions based on artificial criteria, such as arbitrary points on discriminant axes.
Papilio machaon species group
243
wild collected adults from these regions scored on the first discriminant axis (Figures 20-21), a
distribution appeared of character combinations similar to that obtained from reared material.
This indicated that the reared material probably included a representative sample of the
foodplants which larvae of these populations feed on in nature.
Tests for Hardy-Weinberg equilibrium. — Chi-squared tests for deviation from
Hardy-Weinberg (HW) proportions of enzyme genotypes were used as an indication of whether
or not gene flow occured relatively freely within populations. A sample showing a significant
excess of homozygous genotypes at a particular locus then suggests some sort of behavioral or
genetic incompatibility between the different alleles. In this study, I assume that homozygous
excesses indicate likelihood of positive assortative mating or higher heterozygote mortalities,
which in turn suggest the presence of more than one species. I also assume that the different
enzyme banding patterns are inherited, and for that reason I refer to “loci” rather than
“electrophoretic characters” in this section.
Figure 22 includes all six loci which showed a difference from HW proportions at the 5%
level in at least one of the five major geographic regions. Since numerous tests for HW
equlibrium were performed, at least some of these are likely to show spurious differences.
HW tests on the six loci showed that all five of the major regions had at least one locus with
proportions of genotypes different from equilibrium at the 1% level. This suggested more than
one species in each region. However, the pattern of deviation from equilibrium and the loci that
deviated were different in each of the region.
In southern and central British Columbia, highly significant excesses were identified of
homozygous genotypes at the Est4 and G-6-PD loci. These excesses disappeared when the total
regional sample was divided into two groups on the basis of the clusters formed on the first PC
axis of the analysis on both electrophoretic and morphometric characters, and the tests were
rerun. This corroborates the hypothesis that two species are in that region. The P. m. oregonius
subsample showed an excess of heterozygotes of the Prot2 locus which was significant at the 3.3
% level. No biological explanation is offered for this.
The total sample from the Peace River region showed large deviations from equilibrium in
five of the six loci, due to homozygote excesses. When it was subdivided in the same way as for
southern British Columbia, most of these excesses were eliminated in the P. machaon and P.
zelicaon subsamples. A further subdivision of P. machaon on the basis of habitat eliminated the
slight excesses in the Est5 and IDH loci, and supported the division of P. machaon in this
region into two ecological races, P. m. alias ka and P. m. pikei. However, a small homozygote
excess showed up in P. m. aliaska in the Protl locus. The biological significance of this, if any,
is unknown, though a reasonably large allele frequency difference at this locus between the two
races should be noted.
The central Alberta regional sample was very different from the previous two, despite the
fact that the total sample had fairly high frequencies of both of the pairs of alleles at the Est4
and G-6-PD loci which normally distinguish P. machaon and P. zelicaon. The total sample
contained genotypes not much different from equilibrium at any of the loci, except Est4. When
P. machaon was separated from P. zelicaon in the same way as for the Peace River and
southern British Columbia regions, the highly significant homozygote excess was retained in
the P. zelicaon zelicaon subsample. A division of the total sample at 4.0, rather than 3.0, on the
same PCA axis showed a slightly significant excess of heterozygotes at the IDH locus. Clearly,
the second division furnished subsamples more consistent with the hypothesis that there are two
species in that region. However, it divided the total central Alberta sample down the middle of
Quaest. Ent., 1987, 23 (2)
Table 5. Morphometric character state distributions for population samples of the P. machaon species group
244
Sperling
<0 c
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oreaonius pikei dodi X machaon polyxenes polyxen<
Papilio machaon species group
245
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Quaest. Ent., 1987, 23 (2)
Table 6. Allele frequencies, by region, taxon and hybrid assemblage of the P. machaon species group.
Sperling
246
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Papilio machaon species group
W c
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Quaest. Ent., 1987,23 (2)
Heterozygosity:0.233 0.155 0.188 0.203 0.218 0.196 0.219 0.246 0.076 0.162 0.158 0.078
248
Sperling
a somewhat bell-shaped curve of character combinations (Figure 12), and may simply reflect
the large contribution of the Est4 locus to the scores on that axis.
With division of the sample from central Alberta region into three subregions (a northern,
middle and southern one), both the northern and southern subregions appeared not to be
significantly different from equilibrium at the Est4 locus either, despite the higher proportion
of alleles characteristic of P. zelicaon in the northern subregion (Table 6). This was especially
interesting in the sample for the southern subregion, since it was composed of specimens from
only a single locality at Bragg Creek. The deviation from equilibrium in the Bragg Creek
sample at the IDH locus was caused by an excess of heterozygotes. The sample from the middle
subregion continued to show an excess of homozygotes at Est4. With specimens reared on
different foodplants considered separately, it was clear that a slightly less significant excess
remained at that locus. These adults were obtained from larvae collected on Heracleum plants
(Umbelliferae).
The highly significant homozygote excess was retained even with the largest sample from a
single locality in that subregion, Buck Lake, considered separately (not shown on Figure 22).
This locality is, to my knowledge, about 80 km from the nearest stand of Artemisia
dracunculus. These results suggest that P. machaon and P. zelicaon have merged their gene
pools in a large part of the central Alberta region, although not all loci are at equilibrium in the
middle part of that region.
The regional sample from southern Alberta and Saskatchewan showed significant
homozygote excesses at three loci. A division of the sample on the basis of scores on the first
PCA axis of the analysis on both morphometric and electrophoretic characters eliminated the
excess at the Est4 locus, but only reduced it at the IDH and G-6-PD loci. With the sample
divided on the basis of scores on the first axis of the discriminant analysis of reared specimens,
the homozygote excesses were retained at the Est4 and IDH loci. Division of the sample from
this region into two species is supported by the fact that the homozygote excesses were partially
reduced even when the sample was subdivided on the basis of just morphometric characters.
However, this subdivision is not as good as that effected in the southern British Columbia
and Peace River regions. One reason may be a higher probability of incorrect species
assignment, because of the greater morphometric similarity between P. machaon and P.
zelicaon from this region. A second explanation is that, even though the species retain a
separate genetic identity, there is a biologically significant amount of gene introgression
between the species. The rate of introgression may differ between loci, as for example between
Est4 and G-6-PD. This suggestion is supported by the fact that these two loci have very similar
differences in allele frequency between P. machaon and P. zelicaon in the southern British
Columbia and Peace River regions, and yet G-6-PD seems to have reached equilibrium before
Est4 in the middle part of the hybrid zone in central Alberta.
In the region that included samples from Manitoba, Wisconsin and Ontario, there was also
evidence for two species, with some hybridization between them. Here the main alleles
distinguishing between P. polyxenes and P. machaon were G-6-PD and Prot2, rather than
G-6-PD and Est4, as between P. zelicaon and P. machaon. The total regional sample had
several significant deviations from equilibrium. When it was divided on the basis of scores on
the second PCA axis using both morphometric and electrophoretic characters, which essentially
separated black morph from yellow morph individuals, then a highly significant deviation
remained at the G-6-PD locus in the sample comprised of black morph specimens. These
deviations were eliminated when the region was divided geographically into three subregions.
Papilio machaon species group
249
The main difference from the previous subdivision was that one yellow morph and nine black
morph specimens were placed in a group by themselves. Under both schemes there was a
significant excess of homozygotes in the Est4 locus of the northern Manitoba sample, because
of presence at Thompson of two individuals homozygous for the “B” allele (most common in P.
zelicaon ), compared with only four heterozygous individuals and 33 individuals homozygous for
the “A” allele (most common in P. machaon and P. polyxenes ). I can offer no convincing
biological explanation for this situation, since Thompson is many hundreds of kilometers from
the nearest localities where P. zelicaon specimens have been found. Instead, I suspect, it may be
due to sampling error. Clearly, more work is needed to ascertain species relationships in
Manitoba.
Diagnosis of Adults and Ranking of Taxa
To tabulate character variation quantitatively, 13 major populations were defined. The
arrangement of these groups was based on both electrophoretic and morphometric characters,
and the groups resemble those described in the previous sections involving multivariate analyses
and Hardy-Weinberg equilibria. Table 6, for electrophoretic characters, includes some
specimens for which not all loci were scored, but for which it was possible to be certain of their
identification by reference to their morphometric characters. Table 5, for morphometric
characters, includes data only for those specimens used in the PCAs.
Papilio machaon. — In general, P. machaon adults from western Canada were distinguished
by yellow hair on the ventral part of the thorax and abdomen, yellow scales covering most of the
forewing disc on the ventral side, and the anal pupil connected to the wing margin, whether
club shaped or a thin line. This result verifies the utility of the color pattern characters of adults
used by others to identify this species ( e.g ., Edwards, 1883; Dornfeld, 1980). In electrophoretic
characters, P. machaon individuals were distinguished by the A allele at Est4, the C allele at
G-6-PD and a relatively high proportion of D alleles at IDH.
P. m. dodi specimens were more difficult to identify in the absence of electrophoretic
information, but could best be distinguished from the P. zelicaon populations sympatric with
them by the club shaped, connected anal pupil. I have examined the types of P. m. dodi in the
Canadian National Collection, and they definitely belong to the race of P. machaon whose
larvae feed on Artemisia dracunculus on the prairies of southern Alberta and Saskatchewan.
Subspecies of P. machaon in western Canada were best separated from each other by
locality and habitat, though there were major changes in the frequency of particular states of
several characters, including IDH, ODH, Est4, Protl and Prot2. Though almost all specimens
of P. m. oregonius and P. m. dodi from western Canada are distinguished by both adult and
larval color pattern, these two subspecies grade into each other in western Montana and
southern Idaho. Since the zone of intergradation is narrow, relative to the phenetically more
homogeneous ranges of the subspecies, I recognize the populations on either side of the
continental divide in Montana and northward as separate subspecies.
I am uncertain of the extent and location of the intergradation between these subspecies
south of Montana. In Utah and Colorado, the black adult wing morph becomes more common
(Emmel, 1975) and the name P. machaon bairdii should be applied. The name P. brucei has
been applied to yellow morph adults in polymorphic populations within the range of P. m.
bairdii{ Figure 42). Its type locality is from the northern part of the major clinal shift to yellow
forms, and its use in a subspecific sense is probably not of much value. I follow the practice of
Fisher (1980) and Miller and Brown (1981) in treating the name as a junior synomym of P. m.
Quaest. Ent., 1987,23 (2)
250
Sperling
bairdii.
The previous subdivision of the southern subspecies of P. machaon as separate species is
probably a consequence of W.H. Edwards’ relatively typological species concept, and the
natural tendency of many workers to view the black morph adults as fundamentally different
from the yellow morph adults. However, black morph adults of P. machaon occur in low
frequencies as far north as Drumheller, Alberta, where they are electrophoretically identical to
the yellow morph adults. Hence, I feel that the inclusion of P. bairdii in P. machaon is an
inescapable consequence of the application of the biological species concept to geographic
dines. Fisher (1980) has also recognised the specific unity of all the Artemisia
dracunculus- feeding populations in the western United States.
Papilio zelicaon and hybrids. — Most P. zelicaon individuals from western Canada are
recognized by the black hair on the ventral part of the thorax and abdomen, black or almost
black ventral forewing disc and the rounded, centered anal pupil. As in P. machaon , these
characters match those previously used in traditional taxonomic treatments. Important
diagnostic electrophoretic characters included the B allele in both the Est4 and G-6-PD loci.
I do not believe that formal subspecific divisions are justified in P. zelicaon. The species is
composed of innumerable slightly differentiated populations with adult features that grade into
each other. Local foodplant and climatic adaptations of most populations are usually far more
pronounced than are the relatively minor differences in morphometric characteristics. I believe
that the recent practice, of referring to the populations that Remington (1968a) named P.
gothica as P. zelicaon nitra, is unwarranted. Yellow morph adults are more common than the
black form even at the type locality of P. nitra , and I find the eastern and western yellow
morphs of P. zelicaon to be impossible to separate with any degree of consistency.
The presumed type of P. zelicaon Lucas was examined for me in considerable detail by G.E.
Ball in 1980, on a trip he made to the Paris Museum. Using Ball’s description and comparative
material, as well as photographs of the specimen (taken by J.J. Menier), features of P. zelicaon
were checked against Remington’s (1968a) diagnosis of P. gothica. The specimen is closer to
Remington’s conception of P. gothica than his conception of P. zelicaon. This is not surprising,
since as Shapiro (1975) and Emmel and Shields (1980) pointed out, P. zelicaon from the type
locality in central California has undergone basic ecological changes since its description in
1852, while the remaining populations at higher altitudes in central California are still very
similar in appearance to topotypic P. gothica *
In some regions, particularly central Alberta, a high proportion of individuals exhibited
intermediate character states, or character combinations which placed them in an intermediate
position between P. zelicaon and P. machaon. These were considered to be hybrids (individuals
of mixed ancestry), and such individuals formed the majority of some populations. Since these
populations included individuals with phenotypes occupying the complete range between the
typical parental forms, many individuals were difficult to identify as hybrids. Hybrid
populations were also highly variable in composition, and were only identified as such when
they showed a unimodal distribution of phenotypes, of which the peak was clearly intermediate
between the parental species.
The P. zelicaon X machaon hybrid swarms in the Cypress Hills have been much less
completely documented than those in central Alberta. I designate these populations as hybrid
swarms mainly because most individuals look very similar to the hybrid material collected in
the southern part of central Alberta. As well, they are intermediate in wing and body pattern
between the P. machaon and P. zelicaon specimens collected in the prairie habitats surrounding
Papilio machaon species group
251
the Cypress Hills.
In regions where hybridization between P. zelicaon and P. machaon is rare, possibly there
are structural isolating mechanisms between the species, in addition to behavioral ones. This
was suggested by the only natural interspecific mating which I have observed in such regions of
sympatry. The mating took place at Taylor, at a site where P. m. pikei adults are common, and
involved a fresh P. zelicaon female and slightly worn P. machaon male. They remained in
copula for at least 1 1 hours before they separated. Such an abnormally long mating (Clarke
and , 1956b) may result from disturbance of being netted, but seems more likely to be due to
some sort of prezygotic mating disfunction.
Papilio polyxenes and hybrids. — Most specimens of P. polyxenes were easily distinguished
from those of P. zelicaon , P. machaon and their hybrids by the much greater amount of black
scales on the hindwing, covering more than half of the hindwing disc, and yellow spots rather
than a broad yellow band on the sides of the abdomen. Separation was also based on the K
allele of G-6-PD and the A allele of Prot2.
A small proportion of P. polyxenes- like individuals were noted in P. zelicaon , P. machaon
and their hybrids in western Canada. These, however, had the same electrophoretic alleles as
the yellow morph individuals with which they were found, and were also distinguished from P.
polyxenes by the greater amount of yellow on the tegula and apex of the forewing, as well as
the lesser amount of orange on the postmedian band of the ventral hindwing. The black morph
specimens of P. zelicaon from Alberta prairies were identical in appearance to a series which I
have seen from the type locality of P. nitra in Montana, and so there is no reason to expect
these individuals to comprise a separate species outside of western Canada.
Although the morphometric differences between P. zelicaon and P. polyxenes in western
Canada suggest a greater ease of species identification than Fisher (1980) reported in
Colorado, I expect that I would have found similar difficulties if I had been able to obtain a
larger sample from localities where these two species are in closer contact in southern
Saskatchewan.
Since the interactions of P. polyxenes with P. machaon and P. zelicaon in western Canada
are not well understood, I rely on the opinions of authors who are familiar with the three species
in the western United States (e.g., Ferris and Emmel, 1982; Fisher, 1977 and 1980), and who
have consistently reported that P. polyxenes maintains a distinct genetic identity from both P.
zelicaon and P. machaon throughout most of their region of potential interaction. As well,
although electrophoretic characters indicate some intermediacy in central Manitoba, samples
of P. polyxenes from Ontario and Wisconsin are as different from P. zelicaon and P. machaon
as these two species are from each other.
Most of the specimens of P. polyxenes from southern Manitoba are indistinguishable in
appearance from P. p. asterius from Ontario and the eastern United States. The remainder
show signs of introgression with P. machaon. Specimens exhibiting substantial introgression are
designated as P. polyxenes X machaon hybrids. The identification of such natural hybrid
specimens is supported by comparisons with those obtained by artificial hybridization. In
particular, many of the adults collected in central Manitoba appear very similar or identical to
the hybrids obtained by other workers (see particularly Clarke and , 1953, 1955a; Ae, 1961,
1964; Remington, 1958, 1968a). The same applies to hybrid specimens of P. zelicaon and P.
machaon from central Alberta. These studies clearly indicate the genetic basis of these
characters, and for this reason I have used several of these characters in the morphometric
portion of this study. I consider the similarity between the experimentally produced and
Quaest. Ent., 1987,23 (2)
252
Sperling
wild-collected specimens to be adequate evidence for the hybrid origin of the collected material.
Two taxonomic descriptions refer to adult forms which are due to hybridization between P.
polyxenes and P. machaon. These are P. kahli and P. m. avinoffi, both of which are referred to
in this study either as black or yellow wing morph adults of P. polyxenes X machaon , or as P.
polyxenes X machaon and P. machaon X polyxenes , respectively. My use of these names is
based on photographs I have seen of the holotypes. I have also seen several paratypes, but these
differ slightly from one another, as well as from the holotype. At least one of the female
paratypes of P. kahli in the Canadian National Collection seems to me to be identical to typical
P. p. asterius specimens. My opinion was apparently shared by J.D. McDunnough, who
indicated his opinion on a folded slip of paper attached to the specimen pin.
Specimens which fit the description of P. m. avinoffi were obtained by Remington (1958,
1968a), when he crossed two comparatively yellowish individuals of the black adult morph from
central Manitoba, and got some yellow morph as well as black morph offspring. The avinoffi
form tends to grade into more typical P. m. hudsonianus and so identification of specimens is
arbitrary.
The systematic relationship of P. polyxenes and P. machaon in central Manitoba clearly
needs more investigation than I have provided in the present study. The recognition of central
Manitoba populations as interspecific hybrid populations, rather than as intermediates between
subspecies, allows the retention of established taxonomic practice, pending a more thorough
study of these two taxa in this region, as well as elsewhere in their ranges.
Ranking and accuracy of identification. — The distribution of morphometric and
electrophoretic character states showed, in several ways, that more than one species of the P.
machaon group was present in western Canada. First, multivariate analysis of either of these
two character suites indicated three major clusters of individuals in western Canada, and two
major clusters in each of four of the five regions in western Canada. Second, the proportions of
enzyme genotypes suggested interruptions to gene flow which corresponded to the breaks
between clusters in most of western Canada. Third, the morphometric and electrophoretic
character distributions showed good correspondence with each other, as well as with ecological
features such as preferred habitat and larval foodplant. This character concordance applied to
areas where there appeared to be a large amount of interspecific hybridization, as well as those
in which species appeared to interbreed very little. The characters of wing and body color
pattern, which had been used by taxonomists in the past to distinguish among species, proved
useful under critical examination. A few electrophoretic loci were also diagnostic for species,
and so gave additional information about inter- and intra-population relationships.
However, evidence of hybridization between each of the three species showed that
recognition of some populations as species, and others as interspecific hybrid swarms or
subspecies, was partially arbitrary. This was resolved by an arrangement reflecting the fact
that species hybridize only rarely over most of their sympatric range, and that also involved a
minimum of change in existing taxonomic arrangements. Since previous taxonomic
arrangements were not based on electrophoretic characters, consideration of these allowed an
independent test of the biological significance of these arrangements.
The electrophoretic characters also allowed a more direct comparison with the degree of
genetic similarity between species of other, unrelated, taxa. This comparison was obtained by
calculating Nei’s (1972) Genetic Identity (/) for all combinations of each of the 13
geographically separated populations of the P. machaon group which showed little or no
internal interruption in gene flow. Nei’s Genetic Identity is the most commonly used of several
Mean Genetic Identity is below diagonal and mean Genetic Distance is above diagonal.
Papilio machaon species group
253
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254
Sperling
standardized genetic similarity coefficients, and / has been determined for a wide variety of
taxa.
Of the 13 major populations distinguished in this study, all pairs listed as separate species
had I values less than or close to 0.85 (Table 7). These pairs included those populations from
the three regions in which P. machaon and P. zelicaon occur sympatrically. Thorpe (1982)
showed that when two populations have an I value of less than 0.85, the probability is very high
that they are distinct species. Thus, despite difficulties in separating individuals of some
populations on the basis of morphometric characters, as well as the presence of several hybrid
swarms, genetic similarity coefficients based on elctrophoretic characters suggest that at least
the main clusters were different enough to rank as separate species.
/ values can also be used to make intraspecific pairwise comparisons. About 80% of
conspecific I values are above 0.95 (Thorpe, 1982). In the present study, all comparisons of
populations within P. machaon and P. zelicaon were close to or above 0.95.
The interval between 0.85 and 0.95 is occupied by a few values from interspecific pairs and
a much larger proportion of intraspecific pairs. In the P. machaon group, those comparisons
involving hybrid swarms and one of the parental species generally show / values between 0.85
and 0.95. Several of these, however, resemble one parental species more than the other and
show I values above 0.95 when compared with the more similar species. For example, the
northern part of the P. machaon X zelicaon swarm is closer to P. zelicaon while the southern
part (Bragg Creek) is more similar to P. machaon. This result could have been expected on the
basis of morphometric character similarities. However, the central Manitoba population is
much more like P. machaon than P. polyxenes, a result in contrast to that which might be
expected on the basis of morphometric similarities (for rough comparisons see 3D.PCA scores
in Figures 7-9).
Without information about locality, habitat or electrophoretic alleles, I estimate that I am
able to correctly identify 95% of all specimens from western Canada as members of one of the
groups listed in the key in the previous chapter. My accuracy is probably higher for
distinguishing P. machaon from P. zelicaon in the absence of a large hybrid swarm. P. zelicaon
X machaon and P. polyxenes X machaon hybrids, as well as P. m. dodi, are more difficult to
distinguish from each other and I estimate that I can correctly identify about three quarters of
all such specimens with only morphometric information.
Diagnosis of specimens on the basis of morphometric characters in the key was found to be
fairly reliable when compared to scores obtained from PCA factor loadings. Several characters
used in the keys were not used in the original multivariate character analyses, generally
because they were difficult to score consistently. Both the key and PCA factors produce
arbitrary divisions which are not particularly meaningful in hybrid populations.
Since the five subspecies of P. machaon are allopatric, or parapatric and separated by
habitat in western Canada (see next section), it is possible to obtain a more precise estimate of
accuracy of identification. Using habitat and geographic range to define groups, I performed a
discriminant function analysis on the five subspecies, using the morphometric and
electrophoretic characters which were employed in the multivariate analyses in previous
sections. Since sample sizes were small, only the 27 variables which showed more than 10%
variation in frequency between groups were used. The results are contained in Table 8.
This analysis indicates that a high frequency of correct identification can be achieved for
these taxa if both major character suites are used. The lowest accuracy, 76% for P. m.
hudsonianus, is still fairly high. If only the 1 1 morphometric characters listed in Table 2 are
Papilio machaon species group
255
Table 8. Frequency of correct identification of subspecies of Papilio machaon.
Values based on discriminant function analysis (DFA) of 27 morphometric and electrophoretic
characters. Classification percent shows frequency of correct identification ( e.g ., 82.9% for P.
m. aliaska) and incorrect identification (e.g., 9.8 and 7.2 % of P. m. aliaska were misclassified
as P. m. hudsonianus and P. m. pikei, respectively).
% Classification with DFA
used in a new DFA, rather than the 27 morphometric and electrophoretic characters used to
obtain the results in Table 9, then the lowest accuracy is 62%, again for P. m. hudsonianus.
However, if forewing length and tail length are added to these 1 1 characters, and a third DFA
is performed, then the accuracy of correct identification of P. m. hudsonianus rises to 70%, and
the lowest is 68%, for P. m. pikei. I estimate that my personal lowest accuracy of identification
of these five P. machaon subspecies is 75% if characters such as wing shape and color are
considered, which are difficult to quantify for computer work.
It is difficult to obtain a precise assessment of the relative systematic utility of
morphometric and electrophoretic characters in the context of the present study. The
morphometric characters were chosen on the basis of their variability within western Canada,
and also as a means of comparison to systematic descriptions and diagnoses. Electrophoretic
characters were selected much more randomly, since any protein that showed consistent, simple
banding patterns was used. As well, only three loci showed more than 50% allele frequency
differences between populations, and it is possible that results were affected by sampling error.
Furthermore, the coding scheme for morphometric characters was somewhat different from
that used for electrophoretic characters in the principal components analyses. A more strictly
analogous scheme would have reduced the number of electrophoretic characters from 42 to 10,
a number more comparable to the 1 1 morphometric characters used. Despite these factors, it is
clear that electrophoretic analysis is of considerable systematic utility (cf Wake, 1981). The
large degree of correspondence of the two types of characters in the context of the present study
is a demonstration of the potential usefulness of electrophoretic analysis in systematic research
on species complexes.
Larval Color Pattern
Larvae of P. machaon , P. zelicaon and P. polyxenes do not show consistent interspecific
differences in color pattern, though intraspecific variability is marked. Most fifth instar larvae
of these three species are predominantly green, with a prominent black band extending around
each segment, and six colored spots on most segments. Within populations, the background
green color varies from pale bluish-green to bright emerald green, and the black bands vary
considerably in width. Color of the segmental spots varies from lemon yellow to orange-red.
Quaest. Ent., 1987, 23 (2)
256
Sperling
Figures 23 and 24. Frequencies of spot color in larvae of the P. machaon group in western Canada. Dark areas of
histograms indicate orange or red spots, and light areas indicate yellow spots. Figure 23. Spot colors of larvae collected on
composites. Histograms with broken borders indicate small sample sizes. Figure 24. Spot colors of larvae collected on
umbellifers.
CORRIGENDA-<2wa£s//0/i£s Entomologicae , Volume 22(4)
Griffiths, G.C.D. (1986, 22: 253-260).-Relative Abundance of the Root Maggots Delia radicum
(L.) and D.floralis (Fallen) (Diptera: Anthomyiidae) as Pests of Canola in Alberta.
page
256
Fig. 2
Substitute following figure for the one on page 256.
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per day
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per day
Papilio machaon species group
257
The larvae of P. alexanor have a color pattern which is very similar to that of the above
three species, despite a very different adult wing pattern. P. indra and P. hospiton each have a
larval color pattern which is divergent from that of the other species in the P. machaon group,
but which is more similar to that of other P. machaon group members than to other Papilio.
Color of the segmental spots on mature larvae is consistent on individuals, but many larval
populations are composed of discrete color classes with few intermediates. Clarke and Knudsen
(1953) were the first to study the genetic mechanism controlling spot color, by crossing
yellow-spotted P. polyxenes larvae with orange-spotted P. machaon larvae, to produce
orange-spotted hybrids. Clarke and Sheppard (1955b, 1956a) later showed that the hybrid
larvae had spots which were a paler orange than those of P. machaon , and although yellow is
recessive to orange, the degree of dominance of the orange allelomorph varies with the
subspecies of P. machaon , They suggested that inheritance was controlled by more than one
allele which could produce orange spots, or through modifiers at another locus. Clarke et al.
(1977) established that the main locus controlling larval spot color was not linked to the locus
controlling black and yellow wing morphs in adults.
P. machaon , P. zelicaon , P. polyxenes and P. indra are to some extent all polymorphic for
larval spot color. Thus, this polymorphism probably predates the most recent common ancestor
of these species. Nonetheless, differences in spot color have been used to support some
taxonomic distinctions. For example, Remington (1968a) considered the fact that he had found
and reared only yellow-spotted larvae of P. gothica as evidence contributing to his decision to
name it as a separate species. His sample was comprised of few independent observations and
provided little support for his decision, considering that most populations of P. zelicaon were
known to produce both yellow and orange spotted larvae (Clarke and Sheppard, 1970). All 34
late-instar larvae which I found at Gothic had yellow spots, supporting Remington’s
observation of apparent allelic homogeneity at that locality. It would be interesting to
determine if larvae of P. zelicaon in central Colorado generally have yellow spots and if their
proportion decreases in populations farther away from P. polyxenes.
In western Canada, many samples from single localities contained a mixture of both
yellow-spotted and orange-spotted larvae, but there were also a number of interesting frequency
shifts between different taxa and between different regions (Figures 23-24).
The largest differences in frequency of spot color occurred within P. machaon. All larvae
collected on Artemisia dracunculus in southern and central British Columbia had yellow spots,
while in southern Alberta and Saskatchewan 99.6% had orange spots. Thus P. m. dodi and P.
m. oregonius may have undergone a complete allele substitution over much of their range in
western Canada. I do not know what the predominant spot color is where these races contact
each other in the western United States.
In the Peace River region 28% of the larvae of P. m. pikei had yellow spots. If spot color is
controlled by a single gene with orange dominant over yellow, then the Peace River A.
dracunculus- feeding populations have a 50:50 ratio of these two alleles, making them exactly
intermediate between P. m. dodi and P. m. oregonius.
There were far more yellow-spotted larvae in P. m. aliaska than in P. m. pikei , since 95% of
the larvae collected on A. arctica had yellow spots. The single larva of P. m. hudsonianus which
was scored (from a photograph by G. Anweiler) also had yellow spots. Thus P. machaon is
clearly polymorphic for spot color in North America, but different ecological and geographic
races have major frequency differences in spot color.
Quaest. Ent., 1987, 23 (2)
258
Sperling
Samples of larvae collected on umbellifers in almost all localities from Alberta and British
Columbia were polymorphic for spot color. Hence geographically separate populations of P.
zelicaon and its hybrids may have quite different frequencies of spot color, as in P. machaon.
However, the frequency shifts seem to be somewhat more clinal. Also, P. zelicaon larvae
consistently had different spot color frequencies from P. machaon where these species have low
hybridization rates. In Interior British Columbia and the Peace River region, P. zelicaon larvae
had more orange spots, while in southern Alberta they had more yellow spots.
Hybrid populations did not show much difference from parent species. In central Manitoba,
most of the larvae had yellow spots, while P. polyxenes larvae generally have yellow spots
farther to the southeast, and the only known larval P. m. hudsonianus also had yellow spots. In
central Alberta the northern populations have mostly orange spots and in this respect merge
into the P. zelicaon populations farther to the north and west. This trend is mirrored in the
adult morphometric and electrophoretic characters of these populations.
The shift toward predominantly yellow spots in the P. zelicaon X machaon hybrid swarm
west of Calgary is more abrupt. Also, it is interesting to note that the only larva found on
Heracleum at Bragg Creek had orange spots, while 42 of 44 on Zizia had yellow spots.
Heracleum is a much more common foodplant for P. zelicaon populations immediately to the
west, and larvae of these populations may have developed from eggs laid by a typical P.
zelicaon that strayed in from the west. The fact that P. m. dodi larvae almost always have
orange spots, even at the outer edges of the range of this subspecies, distinguishes the latter
from the southern hybrid populations. This supports the contention that hybrid populations are
the result of hybridization of P. zelicaon with races similar to P. m. hudsonianus , rather than to
P. m. dodi.
ECOLOGICAL CHARACTERISTICS
Geographic Distribution and Habitat
The Papilio machaon group has a generally Holarctic distribution. Of the eight species
recognized, four are restricted to North America, P. alexanor and P. hospiton occur only in
western Eurasia, and P. machaon spans both continents. P. polyxenes is found mainly in North
America, but is the only species that also occurs in South America.
Species of the P. machaon group which have broad ranges also show a considerable diversity
of habitat use. For example, different populations of P. machaon occur in habitats varying from
cool temperate wetlands (Wiklund, 1974; Dempster and Hall, 1980) to hot Saharan deserts
(Larsen, 1980), and in Nepal two distinct races are separated only by a continuous cloud belt
(Dierl, 1976). Such varied habitat use may occur with relatively little evidence of regional
morphological differentiation, as in mountain- versus prairie-adapted populations of P.
zelicaon.
The three species of the P. machaon group living in western Canada interact in a complex
pattern of geographic overlap, replacement along contact zones, and varying frequencies of
hybridization (Table 9). P. machaon has five subspecies in this area, and is the only species
which is represented by more than one subspecies. The subspecies of P. machaon are all either
allopatric with each other, or parapatric but with very limited opportunities for gene flow
(Figures 25 and 26). P. m. hudsonianus is rare in northern Alberta and northwestern
Saskatchewan ( e.g ., Bird et al. , 1982), and there is little opportunity for contact with either P.
m. aliaska or P. m. pikei. P. m. dodi and P. m. oregonius are separated from the northern
subspecies and, in Canada, from each other.
Papilio machaon species group
259
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-central Manitoba AS AAAAAAAP2 approx. 97%
P. p. asterius A PI A A P2 A A A A P2 P3 100%
260
Sperling
Figures 25 and 26. Empty symbols indicate unverified published records. Figure 25. Distribution of P. m. aliaska. Figure
26. Distribution of P. machaon in western Canada.
Papilio machaon species group
261
• zelicaon - yellow a zelicaon - black
polyxenes
• zelicaon X machaon - yellow machaon X polyxenes - yellow
▲ zelicaon X machaon -black ■ polyxenes X machaon - black
Figures 27 and 28. Figure 27. Distribution of P. zelicaon and P. polyxenes asterius in western Canada. Empty symbols
indicate unverified published records. Figure 28. Distribution of interspecific hybrids in western Canada.
Quaest. Ent., 1987, 23 (2)
262
Sperling
D P machaon
S hybrids
| Rzelicaon or polyxenes
P z e l i c a o n
P. polyxenes
Figure 29. Frequency of interspecific hybrids in western Canada.
Papilio machaon occurs in most of the available major vegetation zones in western Canada
(Figures 30 and 31), as shown by relating long term records for temperature and precipitation
(Canadian Climate Normals, 1951-1980. [1982a and 1982b]) for weather stations close to
localities at which specimens have been collected. These show a clear pattern of differing
habitat use among the subspecies of P. machaon , as well as between P. machaon and the other
two species (Figure 32 and 33).
P. m. dodi , P. m. oregonius and P. m. pikei are most common in patchy populations in dry
valley bottoms and slopes of river banks or badlands. They are replaced by P. zelicaon at higher
altitudes and in moister habitats ( e.g ., McDunnough, 1927), though males of the two species
are occasionally collected together on hilltops immediately surrounding dry valleys. P. m.
aliaska is replaced by P. zelicaon in forested areas south of the Peace River, although P.
zelicaon is not a resident in alpine habitats and only a few individuals fly to the tops of the
lower mountains. According to Freeman (1972), the northern populations of P. machaon are
also absent from areas with acidic granitic formations.
In contrast to the situation within P. machaon , populations of P. zelicaon are relatively
continuous, with no evidence of any major disjunctions within the species (Figure 27). P.
zelicaon occurs in broad sympatry with P. machaon , with a frequency of less than 5% of
natural hybrids in much of western Canada (Figure 29), but large hybrid populations occur in
south and central Alberta.
The Cypress Hills of southeastern Alberta and southwestern Saskatchewan contain one
major hybrid population (Figures 28 and 29). P. zelicaon is found on the partially wooded and
prairie hills surrounding the plateau, sometimes together with P. m. dodi , but both species
merge into a hybrid population in the more heavily wooded central areas. The “Cypress Hills
Papilio machaon species group
263
Old World Swallowtail” (misidentified as P. m. dodi ) of Hooper (1973), most likely refers to
this hybrid swarm material.
A much larger series of hybrid populations is in central Alberta - probably the result of
genetic swamping of a P. m. hudsonianus- like population which once existed in this region
(Figure 42 and 44). P. zelicaon abruptly replaces the hybrid swarm populations west of the
easternmost slope of the Rockies in Alberta, as well as south of the Crowsnest Pass. Near
Lesser Slave Lake, at the northern edge of the central Alberta hybrid swarm, hybrid specimens
form 20 to 40% of the total population at any one locality. This frequency increases toward the
south and reaches a maximum west of Calgary, where specimens assigned as hybrids comprise
more than 90% of the total populations (Figure 29). I have noticed no difference in habitat
between individuals which are the most P. zelicaon-Yike, and those which are the most P.
machaon-Wkt (“nr. hudsonianus’’'’ in Figure 26). Almost all the localities at which hybrid
populations are found occur between 1000 and 2000 m elevation in central Alberta, while
localities recorded for P. m. dodi are below 1 100 m (Figure 11). Hybrid forms are less common
farther south and east of Calgary, probably because the foothills and mixed forest habitat they
occupy is greatly reduced in extent. A few hybrid specimens from Bragg Creek and Buck Lake
seem likely to have been derived in part from P. m. dodi. These adults have the long tails and
pointed forewings which usually distinguish P. m. dodi from both P. m. hudsonianus and P.
zelicaon.
P. polyxenes occurs in Manitoba, where the species fills part of the gap between boreal P.
m. hudsonianus and prairie P. m. dodi and P. zelicaon (Figures 26 and 27). P. polyxenes X
machaon populations from central Manitoba occupy a habitat very similar to that of P.
zelicaon X machaon populations from central Alberta (Figures 28 and 33).
The hybrid populations in central Manitoba are for the most part isolated from the main
range of P. m. hudsonianus and may be in the process of being swamped by P. polyxenes in the
area, along with the remnants of P. m. hudsonianus. Most typical P. m. hudsonianus adults
were collected in Riding Mountain Park in the 1930’s and 1940’s, at which time they appeared
to form about half of the catch of local collectors. By 1955 P. m. hudsonianus was already quite
uncommon (Remington, 1956), and in the mid 1970’s it was certainly very rare (Heron and
Robinson, 1976). Intermediate black morph adults also may be becoming less common, since
they were at least as common as more typical P. polyxenes in the 1930’s to 1950’s, but have
formed a lower proportion of the total catch in the last two decades. As well, specimens closer
to the typical appearance of P. p. asterius are more common in the farmland, which surrounds
Riding Mountain Park completely and Duck Mountain Park on three sides. Hybrid specimens
are very rare north of Duck Mountain Park (Figure 16). The changing status of the hybrid
populations in central Manitoba is a major reason for retaining the established taxonomic
practice of recognizing these taxa as separate species.
P. p. asterius is very uncommon in southern Saskatchewan, and so there is little contact with
P. zelicaon in Canada. In Colorado, P. polyxenes occurs at lower altitudes than P. zelicaon ,
though these two species meet and occasionally hybridize along a broad zone of contact
(Remington, 1968b; Fisher, 1980; Scott, 1981). In Missouri, P. polyxenes completely
surrounds the range of P. joanae, and apparently these two species are reproductively isolated,
in part by habitat preferences (Heitzman, 1973). Since P. polyxenes and P. joanae are
distinguished by very few morphological characteristics, this contention of isolation is clearly in
need of confirmation. Since the problem has not been investigated in the present study, I follow
current practice (Opler and Krizek, 1984), and refer to P. joanae as a separate species.
Quaest. Ent., 1987, 23 (2)
Table 10. Flight periods in western Canada
264
Sperling
x +
X X
X X
X X
+ 4
X +
X 4
X 4
X
4 +
+ X
+ 4 X 4
X 4 4 X
XI 4 4
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Papilio machaon species group
265
■ extensive alpine tundra
0 low arctic tundra
□ forest
E3 aspen parkland
B grassland
Figures 30 and 31. Figure 30. Major vegetation zones in western Canada Figure 31. Mean annual temperature and
precipitation of major vegetation zones Canadian Climate Normals, 1951-1980 (1982a and 1982b). 1, Grassland in
southern British Columbia. 2, Grassland in southern Alberta and Saskatchewan. 3, Aspen parkland of Alberta to
Manitoba. 4a, Boreal forest of northern British Columbia to Quebec. 4b, Forest of south and central British Columbia. 5a,
Arctic tundra of Northwest Terr, to Quebec. 5b, Alpine tundra of Alberta, British Columbia and Yukon Territory.
Quaest. Ent., 1987, 23 (2)
266
Sperling
Figures 32 and 33. Figure 32. Occurrence of P. machaon subspecies vs. mean annual temperature and precipitation: 1, P.
m. oregonius in southern British Columbia. 2, P. m. dodi in southern Alberta and Saskatchewan. 3, P. m. pikei. 4a, P. m.
hudsonianus in Alberta to Manitoba. 4b, P. m. hudsonianus in Ontario and Quebec. 5, P. m. aliaska. Figure 33.
Occurrence of P. zelicaon and P. polyxenes vs. mean annual temperature and precipitation 6, P. zelicaon in Alberta and
British Columbia; 7, P. zelcaon X machaon in central Alberta; 8, P. polyxenes X machaon in central Manitoba; 9, P. p.
asterius in southern Manitoba; 10, P. p. asterius in eastern Canada.
Papilio machaon species group
267
Degree Days (above 10 °C ) 34
Figure 34. Relationship between voltinism and degree days. Bars show range of degree days (above +10 C) to which P.
machaon group taxa are exposed in western Canada. Data are taken from weather stations near collection localities
(Canadian Climate Normals, 1951-1980. [1982c]). Dashed line indicates approximate number of degree days above which
populations have a partial second brood.
Phenology
Variation in flight period and voltinism in the Papilio machaon group is primarily
geographic and within species. P. machaon , P. zelicaon, and P. polyxenes each include a
variety of different populations which range from strict univoltinism to multivoltinism, and
have a flight period for adults which ranges from a few weeks per year to an almost or
completely continuous emergence (Blau, 1981a, 1981c; Dornfeld, 1980; Emmel and Emmel,
1973; Fisher, 1980; Wiklund, 1973; Wiltshire, 1958). The remaining species in the P. machaon
group also have fairly flexible phenologies. Some populations of P. indra and P. hospiton are
partially bivoltine (Fisher, 1980; Kettlewell, 1955). These two species and P. alexanor may
have a very extended emergence period, depending on climatic conditions, and some pupae of P.
indra and P. alexanor have remained in diapause for several years (Fisher, 1980; Nakamura
and Ae, 1973). Since most species in the P. machaon group exhibit labile phenological
responses to different habitats, the genetic potential to adjust in these ways is probably
plesiotypic (ancestral) within the species group.
Quaest. Ent., 1987, 23 (2)
268
Sperling
Phenological variation in the P. machaon group species is less pronounced in western
Canada (Table 10), perhaps due to a more limited range of habitats. P. machaon and P.
zelicaon are strictly univoltine in the northern regions of western Canada, and have a large
second generation in the warmer southern regions (Figure 34). Partial bivoltinism is more
widespread in southern P. machaon than in P. zelicaon , probably because the P. machaon
populations occupy warmer habitats. P. polyxenes is at least partly bivoltine through most of its
range in western Canada, but also shows a marked decrease northward in the size of the second
generation, even though it occupies the relatively warm agricultural areas. The main flight
period tends to occur slightly later in the year at higher altitudes and latitudes for all three
species.
Although P. zelicaon adults tend to emerge slightly earlier in the season than those of P.
machaon where these species are sympatric, the amount of overlap in flight period is still very
large and cannot account for any interruption in gene flow between the species. A substantial
amount of overlap in flight periods is also true of P. machaon and P. polyxenes in Manitoba.
Voltinism of P. machaon group populations is related to growing temperatures in western
Canada. Figure 34 shows the approximate range of degree days above 10 C (Canadian Climate
Normals, 1951-1980. [1982c]) to which these different populations are exposed. Populations
which have some adults emerging in a second brood occur in areas receiving approximately 900
or more degree days per year. This contrasts with the situation in P. glaucus , in which the
potential for multivoltinism appears in populations from areas receiving more than 1200 degree
days per year (Scriber, 1982).
Despite similarities among related species with respect to phenologies, many artificial
interspecific hybrids in Papilio show unusual characteristics of adult emergence. These can be
relatively pronounced even when both species are within the same species group (Clarke et al.,
1972 and Oliver, 1969), as well as when they are more distantly related (Shimada, 1979).
Natural hybrids of the P. machaon group in western Canada usually fly at the same time as the
main flight of the parental species. However, several interesting exceptions are noted. In the
grasslands of both the Peace River region and southern Alberta, where hybrids are rare, the
latest record for any P. machaon group individual is for a hybrid specimen.
In central Manitoba, P. p. asterius has a partial second brood, while P. m. hudsonianus is
univoltine. Some black morph individuals of P. polyxenes X machaon have been collected
during the second brood flight period, but yellow morph hybrid individuals are only known
from the first brood. This provides evidence, independent of similarities in color pattern, that
yellow morph hybrids represent only the most P. m. hudsonianus-Wke, proportion of the central
Manitoba hybrid populations.
In central Alberta, the most P. machaon-Yike and the most P. zelicaon-Wke, individuals
generally fly in about the same proportions through most of the flight period. However, at
Bragg Creek P. zelicaon-\ike individuals occur at low frequency throughout the flight period,
but include the only two specimens collected as late as mid August (Table 10). The most likely
explanation for the occurrence of these individuals is that they have dispersed in from the more
typical P. zelicaon populations in the mountainous Kananaskis area immediately to the west.
Since most of the P. zelicaon adults west of Bragg Creek fly at least a month later than hybrid
populations at Bragg Creek, there is a partial temporal isolation of these hybrid populations
from the parental species.
Although phenological variation within species of the P. machaon group grades clinally
from one region to another and is generally not useful for making taxonomic distinctions, P. m.
Papilio machaon species group
269
pikei shows some distinctive features. It differs from southern A. dracunculus- feeding P.
machaon subspecies in that P. m. pikei is strictly univoltine, while some proportion of these
southern populations emerges as a second generation. These differences are maintained when
larvae of these subspecies are reared together in the laboratory. Also, adults of P. m. pikei
emerge a relatively long time after the onset of warm temperatures, compared to P. m. dodi, P.
m. oregonius and P. m. aliaska. This difference in emergence time is especially noticeable
considering that P. m. dodi adults are out early in the flight periods of such species as Oeneis
uhleri Reakirt and Papilio glaucus L. in southern Alberta, but P. m. pikei adults do not fly
until after the main flight of these species in the Peace River region. A possible explanation for
this is that the short growing season in the Peace River region provides strong selection against
bivoltinism, while larval foodplants are more abundant or palatable later in the season.
Larval Food Plants
Larvae of the P. machaon group feed on a variety of species of Umbelliferae, Rutaceae and
Compositae (Berenbaum, 1981; Emmel, 1975; Emmel and Shields, 1980; Higgins and Riley,
1970; Wiklund, 1974). Umbellifers are the most commonly recorded foodplants, though larvae
of most species can feed opportunistically on rutaceous plants, and some populations of P.
machaon have switched to composites. Within a restricted area, however, most larvae of a
species are found only on one or a few species of foodplant.
The ability to feed on rutaceous plants is widespread in the Papilioninae, and particularly in
the genus Papilio (Ehrlich and Raven, 1964; Richard and Guedees, 1983; Miller, 1986).
Umbellifer-feeding species are restricted to Papilio, and are concentrated in the P. machaon
group. Miller (1986) postulated that Rutaceae-feeding and Umbelliferae-feeding traits each
arose at least three times in Papilio. On the other hand, the composite-feeding habits of P.
machaon larvae are unique within Papilio. For the P. machaon group, I consider the
composite-feeding habit to be the most derived trait, with umbellifer-feeding ancestral. The
common ancestor probably evolved from Rutaceae-feeding stock, and Rutaceae-feeding habits
have occurred as opportunistic reversals in several species of the P. machaon group.
Of the latter, larvae of P. machaon feed on the greatest range of foodplants. Records for
Eurasian larvae of this species are mainly from umbellifers, less commonly from rutaceous
plants, and only accidentally from plants of other families. An important exception occurs in
northwestern Afghanistan, where Muting (1972) found mature larvae of P. machaon centralis
Staudinger on “Wermutstrauchern” {Artemisia absinthium L.: Compositae). Larvae of North
American subspecies of P. machaon appear to feed almost exclusively on composites, and
particularly on plants of Artemisia (Figure 35 and 36). This contrasts with P. machaon
populations from northeastern Siberia, which often occur in similar habitats, but for which
Kurentzov (1970) mentions only various umbellifers as larval foodplants. Details about
occurrence of larvae on foodplants are provided by Sperling (1986).
The only two records for wild-collected larvae of P. m. aliaska from North America are
from composites (Table 14). Several other foodplant records for P. m. aliaska refer to
ovipositions observed on A. arctica plants. A few larvae of P. m. aliaska may feed on
umbellifers in nature (Kimmich, 1979), but no larvae have as yet been collected on umbellifers.
A single record of a wild-collected larva is known for P. m. hudsonianus (Table 14). I have
seen a photograph by G. Anweiler of a freshly molted fifth instar larva resting on a leaf of
Petasites palmatus (Compositae), with feeding signs on the side of the leaf. The plant
represented a different variety of the same species on which Bryant found P. m. aliaska larvae
Quaest. Ent., 1987,23 (2)
270
Sperling
Figures 35 and 36. Locations of larvae collected on composites: A = Artemisia arctica Less; black dots= Artemisia
dracunculus L.; P = Petasites palmatus (Ait.) Gray. Figure 35. P. machaoti larval records - Alaska and Yukon
Territory. Figure 36. P. machaon larval records - Western Canada.
Papilio machaon species group
271
at Aklavik (Leussler and Bryant, 1935). Anweiler also observed oviposition by P. m.
hudsonianus, which was reported by Hooper (1973) as being on black snakeroot
(Umbelliferae), but the record could just as well refer to Petasites palmatus (R. Hooper, in litt.
1981).
Many larvae of the P. machaon group have been found on plants of Zizia aptera
(Umbelliferae) in central Manitoba. Though most of these developed into black morph adults
(P. p. asterius or P. polyxenes X machaon hybrids), about 2% of them produced yellow adults
similar to P. m. hudsonianus (in litt., J. Troubridge). However, the yellow adults differed from
typical P. m. hudsonianus in showing a basally darkened, avinoffi- like wing pattern that
suggested they were hybrids with P. polyxenes. I have also reared adults very similar to P. m.
hudsonianus from P. zelicaon X machaon hybrid populations at several localities on the east
slope of the Rockies in Alberta. The foodplants included Z. aptera at Bragg Creek, and H.
lanatum at Buck Lake and Nordegg.
Although the southern subspecies of P. machaon have previously been considered to be
specifically distinct from P. m. aliaska and P. m. hudsonianus, it is clear that the differences in
larval foodplant between these taxa are relatively small. Larvae of P. m. pikei, P. m. oregonius,
P. m. dodi, and P. m. bairdii are all restricted to a single species of Compositae, Artemisia
dracunculus (Table 14). Many larvae of P. machaon collected on A. dracunculus feed on
umbellifers if they are transferred to them (Edwards, 1893, 1898; Emmel and Emmel, 1963; J.
Troubridge, in litt. 1981; personal observation, 1982; but contrast Newcomer, 1964). However,
mortality of these larvae is high on most umbellifer species. Larvae of P. zelicaon and P.
polyxenes, for their part, do not feed on A. dracunculus.
P. zelicaon larvae feed on rutaceous plants as well as on a wide variety of umbellifers in the
United States, and Angelica species seem to be especially favored ( e.g ., Emmel and Shields,
1980). In contrast, P. indra larvae are found on a narrower range of umbellifers, particularly
Lomatium species (Emmel, 1975). Although largely separated by habitat, some P. zelicaon
larvae feed on the same species of foodplants as those used by P. indra larvae.
In California, P. zelicaon larvae feed frequently on the introduced weedy umbellifer,
Foeniculum vulgare Mill., and in some localities feed on Citrus (Rutaceae) orchards (Shapiro
and Masuda, 1980). The foodplant shift to recently introduced umbelliferous and rutaceous
plants has allowed P. zelicaon to produce several broods a year on these host plants, rather than
the single brood that is normally possible on native umbellifers (Emmel and Shields, 1980;
Sims, 1980 and 1983). Remington (1968a) considered larval foodplant preferences as evidence
for the specific distinctness of his P. gothica. However, the foodplant preferences he listed have
been disputed by Emmel and Shields (1980). I have confirmed the findings of the latter authors
by obtaining larvae from plants of Angelica ampla. One larva was reared to the adult stage.
In western Canada, only umbelliferous foodplants are known for larvae of P. zelicaon
(Table 14, Figures 37 and 38). Heracleum plants are used commonly, though Angelica plants
tend to be used more frequently at localities where these plants are more numerous. Although
plants of Artemisia arctica and Heracleum lanatum grow together near treeline at many sites
in the Peace River region, this does not seem to promote hybridization between P. zelicaon and
P. m. aliaska.
Larvae of P. zelicaon X machaon populations in central Alberta also feed on umbellifers. At
Bragg Creek, these populations show some segregation from P. zelicaon populations to the west
and south, and the larvae feed mainly on plants of Zizia. In the northern part of central
Alberta, the hybrid populations merge into typical P. zelicaon, and the larvae feed on
Quaest. Ent., 1987,23 (2)
272
Sperling
Figures 37 and 38. Locations of larvae collected on umbellifers. Figure 37. Larvae collected on plants of Heracleum
lanatum Michx. Figure 38. Larvae collected on other umbellifers. Aa = Angelica arguta Nutt.; Ad = Angelica dawsoni
S.Wats.; Ag = Angelica genuflexa Nutt.; A1 = Angelica lucida L.; Co = Cicuta occidentalis Greene; Ld = Lomatium
dissectum (Nutt.) Mathias & Constance; Ln = Lomatium nudicale (Pursh) Coult. & Rose; Lt = Lomatium triternatum
(Pursh) Coult. & Rose; Oc = Osmorhiza chilensis Hook. & Arn.; Os = Oenanthe sarmentosa Presl.; S = Sium suave
Walt.; Z = Zizia aptera (A. Gray) Fern; gc = garden carrot ( Daucus carota ) L.; gd = garden dill ( Anethum graveolens)
L.; ge = garden celery ( Apium graveoens) L.; gn = garden parsnip ( Pastinaca sativa) L.; gp = garden parsley
(Petroselinum crispum) (Mill.)Mansf.
Papilio machaon species group
273
Heracleum plants, a more common foodplant of P. zelicaon.
I have obtained black morph adults from two species of Umbelliferae: one, with typical P.
zelicaon on Angelica arguta at Waterton Park, Alberta; others, on Zizia, and were part of the
hybrid population at Bragg Creek, Alberta. In both groups the black and yellow morph
specimens were produced in similar proportions to those of wild-collected adults. Hooper
(1973) also reported both black and yellow morphs being produced from the same
umbelliferous foodplant: garden dill ( Anethum graveolens) at Eston, Saskatchewan. As well, I
have reared one black morph adult from larvae of P. m. dodi , collected on Artemisia
dracunculus at Taber, Alberta. These observations support the electrophoretic evidence, which
indicates that the black adult morph is an integrated part of several taxonomically different P.
machaon group populations in western Canada.
Larvae of P. p. asterius are found on a broad range of umbellifers and even a few rutaceous
species (Berenbaum, 1981). Many of these are either introduced or common in cultivated areas,
and so the fact that P. p. asterius is a common butterfly in much of eastern North America
may be a recent phenomenon, aided by human agricultural patterns in the last two hundred
years or so (Feeny et al., 1985). Larvae of P. p. asterius may occasionally be found on the same
species which support P. joanae and P. brevicauda and so these ecological distinctions between
the two species are not major (Berenbaum, 1978; Jackson, 1982; Opler and Krizek, 1984). In
desert areas of the southwestern United States, P. p. coloro Wright larvae feed mainly on
plants of Thamnosa species (Rutaceae) (Ferris and Emmel, 1982). In this region P. polyxenes
is in part sympatric with P. m. bairdii (larvae of which feed on Artemisia dracunculus ), and P.
indra (larvae of which feed on umbellifers).
As with P. zelicaon , the larvae of P. p. asterius feed only on umbellifers in western Canada.
However, the larvae of P. p. asterius frequently use introduced and cultivated foodplant
species, while this is more infrequent for P. zelicaon larvae. I have examined adult series reared
from two introduced foodplant species in Manitoba and Saskatchewan (Table 14). Both of
these series contain specimens ranging from typical P. p. asterius to at least one which was
more typical of the P. polyxenes X machaon hybrid populations which are common in forested
areas.
In the zone of interaction between P. p. asterius and P. m. hudsonianus in central
Manitoba, the native umbellifer, Zizia aptera , is the primary larval foodplant (Figure 38). Z.
aptera plants are more characteristic of open meadows than forests, and so are not a major
factor in the partial habitat separation between hybrid populations of P. polyxenes and P.
polyxenes X machaon. I have not been able to confirm Tyler’s (1975) report that cow parsnip
(//. lanatum[l ]) may be a larval foodplant for these populations.
Quaest. Ent., 1987, 23 (2)
Table 11. Character state distributions within P. machaon group taxa
274
Sperling
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Q
X CL
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Q) —
3 -M -H TO •
a— — > o
a) o 3 cm
L. 3 3 (0 —
u. < < J 2
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00 Z
a a
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< <
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o o
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O 5
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Papilio machaon species group
275
Figure 39. Reconstructed phylogeny of P. machaon group. Location of numbers shows hypothesized first appearance of
derived states. Letter and number codes are the same as on Tables 2 and 11. Dots on lines represent spread of modifier
gene for black wing morph. Widely spaced dots indicate low proportion of black morph individuals or incompletely linked
gene combinations. Abreviations: c.AB = central Alberta; c.MB = central Manitoba; 1, Bl, Cl, El; 2, Al, B4, E4, H2,
14; 3, 10c; 4, no autapotypies among characters analyzed; 5, D3, F3, G4, H3, J3, Kl.
EVOLUTIONARY HYPOTHESES
Origin and Early Differentiation of the P machaon Group
Outgroup relationships of the P. machaon group are uncertain. Monroe (1961) did not
resolve its affinities to other species groups of Papilio, but associated it with the P. xuthus
group, and suggested that these two groups had affinities with the P. demoleus and P. anactus
species groups. Ae (1979) suggested that the P. machaon group was about as closely related to
the P. xuthus group as it was to the P. paris group, which Monroe did not include in his
reconstructed phylogeny. Ae also showed that the affinities of the P. machaon and the P.
demoleus groups were probably still more distant. Hancock (1983) ranked the P. machaon
group as a distinct genus with only ancestral relationships to most of the remainder of the
Papilionini. He also suggested that P. alexanor represented a lineage predating a split between
the P. machaon group and more than half of the species groups in the Papilionini.
Quaest. Ent., 1987, 23 (2)
276
Sperling
V*' <£>' O- V <<
Figure 40. Changes in geographic range and larval hostplants over time: 1, P. hospitotv, 2, Palearctic P. machaon ssp.; 3, P.
m. aliaska; 4, A. dracunculus- feeding ssp. of P. machaon\ 5, P. m. hudsonianus ; 6, P. zelicaon\ 7, P. indra\ 8, P.
polyxenes; 9, P.joanae (not shown in Late Wisconsin); 10, P. brevicauda.
Papilio machaon species group
277
Since adults of P. alexanor have a very different, and probably plesiotypic, color pattern and
male genitalia, that species must have diverged early from the remainder of the P. machaon
group. In fact, its inclusion in the P. machaon group on the basis of larval characters is so
clearly contradicted by adult characters, that I consider it with some caution in my discussion
of the phylogeny of the P. machaon group.
Because relationships of the P. machaon group are uncertain, it is difficult to determine
which of its character states are plesiotypic. Seyer (1982) polarized character states by
considering genetically dominant traits to have been more recently derived than traits which
are more recessive. Since the allele for the black wing morph of P. p. asterius is dominant, it
was considered derived relative to the yellow morph. On this basis Seyer concluded that P.
zelicaon was phylogenetically older than P. machaon , P. hospiton and P. polyxenes.
Despite the uncertainty involved in such an undertaking, I offer a hypothesis for the
character states of the most recent common ancestor of the P. machaon group, excluding P.
alexanor. My identification of character states as plesiotypic is mainly determined by similarity
of these states to those occurring frequently in different possible outgroups. On this basis, the
ancestral species was probably similar to present day P. machaon though differing in some
respects (Table 11), which indicate affinities with either P. hospiton or P. p. americus. Since
none of the possible outgroups occur in the New World, this ancestral species probably lived in
the Palearctic region, though it must have dispersed to North America early in the development
of the P. machaon group.
The ancestral species certainly lived before the Pleistocene, considering the amount of
differentiation within the group, though I doubt that the present species in the P. machaon
group (excluding P. alexanor ) began to diverge from each other very long before the beginning
of the Pleistocene (Figures 39 and 40). Nei’s (1972) genetic distance (D) can be a rough
indicator of the time of divergence of two lineages, with D increasing by 1.0 every 15-20
million years (Thorpe, 1982:153). Applying this ratio to a value of 0.2 for interspecific
comparisons within the P. machaon group (Table 7), a divergence time of 4-5 million years is
obtained for the three species now occurring in western Canada. For the subspecies within P.
machaon , divergence times of 0.1 to 1.0 million years are indicated. Although these estimates
are imprecise, they nonetheless support the contention that the main lineages of the P. machaon
group diverged before the Pleistocene, while most evolution within lineages took place during
the Pleistocene.
The species which appeared immediately prior to the Pleistocene probably gave rise to four
major lineages in the P. machaon group, in addition to P. alexanor. These lineages include
what are now: 1), P. machaon and P. hospiton ; 2), P. zelicaon; 3), P. polyxenes , P. joanae and
P. brevicauda ; and 4), P. indra. The oldest of these four lineages is probably the one that gave
rise to P. machaon (Figure 39 and 40). Both P. hospiton and P. machaon exhibit very few of
the apotypic character states of the remaining lineages (Table 11). As well, it is more
parsimonious to hypothesize that the P. machaon lineage evolved in situ in Eurasia, and is not
the product of a return dispersal from North America. However, the common ancestor of the
remaining lineages probably fragmented soon after colonizing North America. Both
electrophoretic characters (Table 7) and hybridization in the laboratory (Ae, 1979) indicate
that P. machaon , P. zelicaon and P. polyxenes are approximately equidistant from each other.
Also, natural hybridization occurs between each pair combination of these three species, as well
as between P. machaon and P. hospiton.
Quaest. Ent., 1987, 23 (2)
278
Sperling
Relationships of the three lineages which originated in North America are unclear. P.
zelicaon possesses few if any autapotypic characters, and could conceivably represent the
earliest of the three clades to diverge. However, P. zelicaon shows little internal differentiation.
Although the color pattern of adults of P. p. asterius is probably apotypic, P. polyxenes
contains other races and forms (particularly in P. p. americus) which appear to be more
plesiotypic. P. indra , on the other hand, appears to share some apotypic character states with
some subspecies of P. polyxenes (Table 11), but shows considerable internal differentiation,
and has distinctive adult genitalia and adult and larval color patterns. Though this degree of
differentiation may indicate an early divergence time relative to the other species, it may also
be a reflection of a different sort of selection regime. In fact, it is conceivable that P. indra is so
different only because some factor such as distinctive hilltopping behavior (Shapiro et al.,
1981) or male genitalia may have allowed it to avoid hybridization and introgression with other
species, even though the lineage may be no older than the other three. Since it seems plausible
that all three lineages could have diverged at the same time (in West Coast, American
southwest, and eastern refugia), I have left this portion of the reconstructed phylogeny as a
trichotomy. Further investigations on the species in the American southwest would be
important to understanding phylogenetic relationships within the group.
Two main factors contribute to the obscure phylogenetic relationships within the P.
machaon group. Species with extensive variation and large geographic ranges may
simultaneously bud off two or more peripheral populations which are substantially different
from each other. Reticulation due to interspecific hybridization is also likely to have been a
significant factor in the evolution of the group. For example, the black wing morphs of P.
zelicaon and P. machaon probably result from introgression of genes from P. polyxenes , while
hybrid populations of two of the three potential combinations are described in this study. Both
reticulation and multiple events of peripheral isolation in variable species are likely to produce
discordant character distributions, with resultant difficulties in reconstruction of phylogenetic
relationships.
Pleistocene Divergences Within Major Lineages
The first dispersal of the Papilio machaon group into North America almost certainly took
place across the Beringian region between eastern Siberia and Alaska. Land connections
through Beringia were intermittent during the Tertiary, and these also formed an intermittent
but important biotic dispersal corridor during the Pleistocene (Matthews, 1979).
Large scale glacial advances and retreats that occurred throughout the Pleistocene must
have been an important factor in differentiation of new species and races. These glaciations,
combined with dramatically altered climates, moved many vegetation associations far south of
their present ranges and caused formation of some vegetation associations which have no
modern analogs (Matthews, 1982). Glaciations would have displaced populations of the P.
machaon group a number of times, isolating those in Beringia, and probably fragmenting
populations which survived south of the ice in North America.
Distributions of vegetation associations during and after the late Wisconsinan have been
fairly well documented. I assume that the habitat associations of most P. machaon group taxa
have not changed greatly since this time, and thus infer which regions were occupied by these
taxa during late Wisconsinan time. Locations of these hypothetical refugial areas are shown in
Figure 41, based in part on Scudder (1979:159).
Papilio machaon species group
279
Figure 41. Locations of late Wisconsinan refugia. Continuous lines indicate ice masses. Broken lines show refugia
hypothesized for P. machaon group taxa in the United States and Canada: 1, P. m. aliaska\ 2, P. m. oregonius\ 3, P. m.
hudsonianus\ 3a, P. machaon populations similar to P. m. hudsonianus , remnants present in P. zelicaon X machaon
hybrid populations in central Alberta; 4, P. zelicaon ; 5, P. m. bairdii and P. p. coloro\ 6, P. p. asterius and P. joanae(p.)\ 7,
P. brevicauda.
Quaest. Ent., 1987, 23 (2)
280
Sperling
Figure 42. Distribution of P. machaon in North America. Arrows show hypothesized Holocene dispersal routes.
Papilio machaon species group
281
Figure 43. Distribution of P. zelicaon and P. polyxenes. Arrows show hypothesized Holocene dispersal routes in North
America.
Quaest. Ent., 1987,23 (2)
282
Sperling
The P. machaon lineage. — The P. machaon lineage now contains only one species other
than P. machaon itself. This is P. hospiton, which must have become isolated on Sardinia and
Corsica very early in the history of the lineage. Rumbacher and Seyer (1979) estimated that
this happened during the Mindel glaciation, the second of the four major glacial periods
recognized in Europe. The rest of the P. machaon lineage occupied most of the Palearctic
region, apparently giving little opportunity for populations to develop into anything other than
distinct regional races.
The Pleistocene glaciations were probably especially important to acquisition of
composite-feeding habits by larvae of P. machaon (Figure 40). North America was occupied
early in the history of the species group, and so P. machaon may have existed for a time in
parapatry with the southern species. This kind of replacement may have been similar to that of
P. zelicaon and P. polyxenes , which are presently parapatric over a considerable distance in the
western United States. Considering degree of differentiation in the Artemisia- feeding
subspecies, invasion of western United States by populations with this adaptation must have
occurred before the Sangamon Interglacial.
The foodplant shift may have occurred in Beringia, when local populations began to feed on
Artemisia arctica, a species which is still the main foodplant of Nearctic P. machaon in
Beringia, and is much more abundant than any umbellifer species in the region. The shift would
have been aided by several similarities in secondary plant compounds between plants of the
genus Artemisia and those belonging to the Umbelliferae (Dethier 1941; Berenbaum, 1983).
The plant assemblage of the arctic steppe tundra in Beringia (Matthews, 1982) was probably
very similar to that presently occupying the dry, cold climate of northern Afghanistan, the only
region in Eurasia in which P. machaon has been recorded on Artemisia. The later switch of P.
machaon to Artemisia dracunculus in North America may have occurred in Beringia or
farther to the south, during a warming trend when Beringian populations came into contact
with southern dryland habitats. In any event, the switch to A. dracunculus is associated with
occupation by P. machaon of large regions in sympatry with P. zelicaon , P. indra, and, in part,
P. polyxenes (Figures 40, 42 and 43).
A major factor in differentiation of P. machaon in North America would have been contact
with P. polyxenes. Introgression or even more widespread hybridization just after contact with
P. polyxenes is probably the reason why P. m. bairdii acquired the genes for the black morph
adult (Figure 39). Support for this suggestion may be derived from the marked similarity
between artificial hybrids of P. machaon and P. polyxenes , and the naturally occurring black
morph of P. m. bairdii. Though the black morph of P. machaon may also be found some
distance from the nearest population of P. p. asterius, it is restricted to a region where either
past contact or allele movement through introgression can account for its occurrence.
During late Wisconsinan time, around 18000 years before present, P. machaon must have
survived in North America in several different refugia (Figure 41). P. m. aliaska would have
been restricted to the northernmost refugium, dispersing southward along the Rocky
Mountains of northern British Columbia when glacial ice melted. Movements of P. m.
hudsonianus are postulated with less certainty. The subspecies now ranges to Quebec, but is
very uncommon west of Saskatchewan. This suggests that it diverged from P. m. aliaska before
the late Wisconsinan, and occupied the boreal region south of the ice during the last major
glaciation (Figure 42).
The stock that P. m. bairdii is derived from probably survived in the remnants of desert
habitats in the American southwest, and it seems likely that P. m. oregonius is derived from a
Papilio machaon species group
283
separate population, suggested by the fact that P. m. oregonius has no black form like the one
predominant in P. m. bairdii. Although most reconstructions of vegetational history show only
conifer forest grading into tundra at the edge of the glaciers in the northwestern United States,
I suggest that a probable Wisconsinan refugium for P. m. oregonius was along the eroding
banks of the Columbia River, between Washington and Oregon. Though very close to the ice,
the steep north bank of the river must have had a much drier climate than the surrounding
region, much like that presently characteristic of the Peace River region. It is even possible that
there were dry-tundra adapted P. machaon populations along the southern edge of the ice in
Washington during the Wisconsinan glaciation, giving rise to P. m. oregonius separately from
P. m. bairdii.
P. m. dodi is even more problematical, possibly surviving on eroding river banks in the drier
areas of the northern Great Plains. However, much of the region was covered by conifer forest,
and if P. m. hudsonianus also survived south of the ice, it would probably have had some
contact with P. m. dodi , and yet the latter has a distinctively darker wing and body coloration.
A more likely alternative is that P. m. dodi did not exist as a distinctive population during late
Wisconsinan time. The dark adult coloration could have arisen through hybridization of P. m.
oregonius and P. m. bairdii when these two populations contacted each other during the
Holocene (Figure 39). A second explanation is introgression from P. zelicaon in founding
populations of P. m. dodi , but since there is no evidence for it in electrophoretic characters, this
is less likely.
P. m. pikei probably had a separate origin from the A. dracunculus- feeding subspecies
which range into the western United States. Although almost all of the butterfly species living
with it in the Peace River grasslands are clearly derived from conspecific populations in
southern Alberta, (E.M. Pike and F.A.H. Sperling, unpublished), P. m. pikei is less like P. m.
dodi than any other subspecies of P. machaon in western Canada. P. m. pikei shows a much
greater phenetic similarity to populations of P. m. oregonius living in southern British
Columbia. However, it is unlikely that P. m. pikei has been derived from P. m. oregonius since
evidence available from pollen cores ( e.g ., Hanson, 1949, 1950, 1955; Valentine et al ., 1980;
White and Mathewes, 1982) does not suggest that the grassland vegetation of interior British
Columbia extended across the Rocky Mountains during the Holocene. Thus it is unlikely that
P. m. oregonius was able to reach the Peace River area.
I believe it likely that P. m. pikei differentiated during the Holocene from P. m. aliaska. If
larger size and more pointed wings are considered as recent adaptations convergent with other
ecologically similar races of P. machaon then the greatest morphometric similarity is with P. m.
aliaska, or perhaps P. m. hudsonianus. P. m. pikei has an unusual combination of
electrophoretic character states, perhaps due to sampling error or genetic drift in the original
colonists, but is slightly more similar to P. m. aliaska than to any other P. machaon subspecies
(Table 7). The most westerly populations of P. m. pikei presently live about 25 km from the
nearest alpine populations of P. m. aliaska.
The most likely time of divergence for P. m. pikei is between 8000 and 1 1000 years B.P. The
ice-free section of the foothills east of the Rockies had a periglacial climate at about 11,250
B.P., which was dominated by Artemisia and grasses (Schweger et al., 1981). However,
between 9000 and 6000 B.P. the climate was much hotter and drier than at present, allowing a
major expansion of grassland habitats. The relatively rapid shift from dry tundra to hot
grassland may have been an important factor in the differentiation of P. m. pikei on the Peace
River grasslands.
Quaest. Ent., 1987, 23 (2)
284
Sperling
P. zelicaon and hybrids. — P. zelicaon represents a lineage similar in age to the P. machaon
and P. polyxenes lineages, and yet shows much less tendency toward the development of
geographic races. The reason for this may be that its range has not been fragmented much by
Pleistocene glaciations and habitat changes. P. zelicaon probably occupied a large proportion of
the western United States even during the late Wisconsinan maximum (Figure 41), and its
range may have bisected that of the A. dracunculus-fQQding populations of P. machaon
During the post-Wisconsinan climatic amelioration, P. zelicaon would have expanded its
range into western Canada from two separate directions (Figure 43). One dispersal route was
along the foothills and edge of the Great Plains, and brought genes for the black morph to the
prairies and southern foothills of Alberta. The other disperal wave occupied all of British
Columbia and spread into Alberta through low mountain passes. It colonized the Peace River
region and the northern part of central Alberta.
The two-pronged dispersal of P. zelicaon into Alberta seems to have effectively isolated a
pre-existing population of P. machaon in the foothills of central Alberta. Since most remnants
of this population presently live in forested areas south of Cadomin, the population probably
was not associated with the alpine refugium discussed by Pike (1980). This P. machaon
population came in direct contact with P. zelicaon on all sides and may have had a relatively
low population density, much like P. m. hudsonianus populations in northern Saskatchewan
and Manitoba. A significant number of individuals must have begun to hybridize with those of
invading P. zelicaon and eventually formed hybrid populations along the ecotone between
montane and boreal forest in central Alberta.
In central Alberta, formation of hybrid populations may have occurred gradually during
several thousand years. However, the process appears to have stabilized before the region was
affected by agricultural disturbances about 100 years ago. I have seen several specimens
collected by F.H. Wolley Dod (1901, 1908) around the turn of the century at the “Head of Pine
Creek”, near Bragg Creek, and these are identical to the hybrid swarm specimens which I have
collected in the same area during the last decade.
The Cypress Hills P. machaon X zelicaon hybrids are very similar to many of the hybrid
specimens from the southern part of the central Alberta hybrid region, and yet do not show any
of the more extreme P. m. hudsonianus- like characters present in central Alberta. Continuing
hybridization with P. m. dodi is a possibility in the Cypress Hills, though it would be difficult to
demonstrate using the characters employed in this study. However, P. m. dodi and P. zelicaon
appear to hybridize very little in prairie areas of southern Alberta, much like P. m. oregonius
and P. zelicaon in southern British Columbia. Another explanation for the absence of P. m.
hudsonianus-Yike specimens may be that the genome of the hybrid swarm is composed of a
higher proportion of P. zelicaon genes than in central Alberta. Allele distributions in Pinus
contorta Loudon suggest that the Cypress Hills was a forest refugium during the late
Wisconsinan glaciation (Wheeler and Guries, 1982), and so there may have been a P. machaon
population on the Cypress Hills during this time.
P. polyxenes and hybrids. — P. p. asterius has a range approximately as extensive as that of
P. zelicaon (Figure 43) and shows a similar amount of phenotypic and ecological variation.
However, P. polyxenes includes several other subspecies and two other species have arisen from
the same lineage (Figure 39). The additional P. polyxenes subspecies range from the American
southwest to northern South America and tend to have a more primitive phenotype expressed in
the adults and larvae. They are probably phylogenetically older than the related species in the
P. polyxenes lineage.
Papilio machaon species group
285
In the P. polyxenes lineage is P. joanae, which appears to be a taxon with only slight (and
dubiously significant) differences from P. polyxenes. Also included is P. brevicauda, a species
restricted to the seashore rim in maritime Canada, which probably survived the late
Wisconsinan on the exposed ocean shelves in this region (Figure 41, Matthews, 1979). Adults
of both of these species have a wing pattern very similar to P. p. asterius and must have
achieved reproductive isolation from the latter in the late Pleistocene at the earliest. The gene
for the black adult wing morph probably originated in the early Pleistocene, but underwent
significant modification during the early history of the P. polyxenes lineage, after the
divergence of the southern subspecies (Figure 39).
P. p. asterius probably survived the late Wisconsinan in the ecotone between woodland and
grassland in the southern part of the eastern and central United States (Figure 41). During
post-glacial times this race would have extended its range northward to southern Canada
(Figure 43). However, P. p. asterius may have had a smaller range and a lower population
density before North America was settled by Europeans during the past three centuries (Feeny
et al. , 1985). It probably reached Nova Scotia only about 60 years ago (Ferguson, 1954), and
still seems to be expanding its range in agricultural regions in central Manitoba and
Saskatchewan. P. p. asterius must have contacted P. zelicaon much earlier in the Holocene or
even the late Pleistocene, for the black morph on the eastern edge of P. zelicaon to have spread
several hundred kilometers beyond the range of P. polyxenes.
It is doubtful that P. p. asterius had any significant amount of contact with P. m.
hudsonianus during the late Wisconsinan glaciation, even if both survived in refugia within a
few hundred kilometers of each other. They are presently allopatric over most of their range,
though hybridization has been extensive where they contact each other in central Manitoba.
This hybridization shows signs of not yet having reached an equilibrium, since P. m.
hudsonianus and hybrid forms have become less common in Riding Mountain Park during the
past 50 years.
Speciation Mechanisms
Race formation in the P. machaon group seems to occur fairly quickly, with ecologically and
even phenetically distinctive populations differentiating in a matter of a few thousand years or
even a few hundred years under exceptional circumstances. Recent race formation seems to
have taken place both at the edge of and in the middle of the range of widespread taxa, when
slightly different new larval foodplant resources became available and were opportunistically
colonized by individuals from the adjacent population. The most obvious examples include the
populations of Californian P. zelicaon whose larvae feed on introduced foodplants (Shapiro and
Masuda, 1980), and the Peace River race of P. machaon
The formation of species seems to have taken much longer than the formation of ecological
races, and was probably the result of the adaptation of geographically isolated populations to
successively more different habitats. Speciation in the P. machaon group probably takes place
over hundreds of thousands of years. The low species diversity of the P. machaon group in the
Palearctic region, despite formation of many geographic races, suggests that speciation can not
occur unless there is an extended period of geographic allopatry. However, even if two
populations have been separated for enough time to acquire an independent evolutionary
identity, hybrid populations may still form between separate species.
The maintenance of separate genetic identity is probably dependent on the degree and kind
of ecological differences between two populations when they meet. Genetic mechanisms for
Quaest. Ent., 1987,23 (2)
286
Sperling
diapause determination are examples of the importance of ecological adaptations in the P.
machaon group. Hybrids between P. zelicaon and P. machaon (see phenology section) or P.
polyxenes (Oliver, 1969) may emerge at a time which is not only different from both of the
parental species but is also likely to be inappropriate to the local habitat conditions.
Considering the many interrelated factors that are associated with phenology, such as larval
foodplant availability, speciation and race formation probably involve substantial
reorganization of polygenic balances ( sensu Carson, 1981).
In the P. machaon group, a variety of ecological factors are associated with the coexistence
of more than one species in the same geographic region. A difference in larval foodplant is
probably the most obvious of these factors. The greatest local species diversity anywhere within
the range of the P. machaon group occurs in Arizona and southeastern California. Here larvae
of P. indra feed on umbellifers, those of P. m. bairdii feed on composites, and P. p. coloro
larvae have switched back to a more ancestral foodplant group for Papilio, the Rutaceae.
However, larval resource partitioning does not necessarily imply competition for these resources
(as suggested, for example, by Miller and Brown, 1983). Though Emmel and Emmel (1969)
and Blau (1980) have indicated that larval resources may sometimes limit population sizes in
the P. machaon group, direct larval resource competition between species has not yet been
demonstrated.
Natural Hybridization
Hybridization between closely related species is a well known event in both plants and
animals. The phenomenon is, by definition, in conflict with the biological species concept. Most
animal taxonomists deal with this by describing hybridization as interspecific only if hybrids
are rare in comparison with the parental forms. However, for some species pairs, hybridization
is relatively common and yet the parental species maintain their integrity. The species in such a
taxonomically difficult group are termed semispecies by some authors, while the group itself
may be termed a superspecies (Mayr, 1963). The term semispecies is appropriate for the
species in the P. machaon group, since these are more reproductively isolated than geographic
subspecies and yet hybridize relatively freely in comparison to most other species.
Under some conditions, hybrids are especially common. Such conditions include habitat
disturbance, of which the most common source is the clearing of forests by man. However,
increased rates of hybridization generally take place in very restricted geographic areas. If
hybridization occurs along a narrow line of contact between parapatric species, then such an
area is referred to as a hybrid zone. Hybrid individuals may comprise small or large proportions
of populations in hybrid zones, and may also be present in varying frequencies within larger
areas of overlap between parental species. The species of the P. machaon group in western
Canada show a low but persistent rate of hybridization in most areas where they come into
contact, and in some areas have formed populations in which hybrid individuals are
numerically dominant (Figure 44).
Hybrid zones. — The zone of hybridization between two species may vary in width from a
few hundred meters to more than a hundred kilometers, but is much narrower than the total
range of the parental species (Barton and Hewitt, 1981 and 1985). Most hybrid zones are much
longer than they are wide, and some span an entire continent. In the P. machaon group, the
best examples of hybrid zones are along the periphery of the range of P. polyxenes In the
American West this species replaces P. zelicaon along major ecotones. It involves a
replacement over a few dozen kilometers of P. polyxenes by P. zelicaon in wetter habitats and
Papilio machaon species group
287
Figure 44. Hybridization of P. machaon group species in western Canada during recent time. Circles indicate species.
Relative thickness of lines is proportional to rates of hybridization.
higher altitudes. A similar pattern also applies to the transition from P. polyxenes to P. m.
hudsonianus in cooler habitats in Manitoba, though the width of the real zone of intergradation
between the two species remains uncertain. In both comparisons, the zones are relatively wide
compared to other animals, probably because of the good dispersal powers of Papilio butterflies
(Shields, 1967).
The pattern of variation across a hybrid zone may not be a simple cline, but rather a mosaic
of populations (Harrison, 1986). This is clearly true of the P. machaon group, and appears to
be closely associated with an interdigitation of habitats across major ecotones.
The influence of environmental factors on the location of major hybrid populations is
supported by the coincidence of the P. zelicaon X machaon populations in central Alberta with
ecological boundaries and hybrid zones in other species. For example, the southern limit of the
hybrid populations in southern Alberta occurs very near to the southernmost limit of many
boreal elements. Black spruce ( Picea mariana [Mill.] BSP) is found in Alberta only as far south
as Bragg Creek, while three butterfly taxa characteristic of spruce bogs are also found south to
near Bragg Creek. These include Colias gigantea gigantea Strecker, Oeneis jutta chermocki
Wyatt, and Erebia disa (Thunberg). The northern limit of the P. machaon X zelicaon hybrid
populations in central Alberta also coincides generally with the location of several hybrid zones
in unrelated taxa. An example is the hybrid zone between Pinus contorta and P. banksiama
Lamb., (with a corresponding contact zone between the pine-feeding butterflies Incisalia
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Sperling
eryphon (Boisduval) and I. niphon (Huebner), - Reist, 1979). I am not aware of any animal
taxa which form self reproducing hybrid populations within Alberta. However, in plants there
is a wide ranging hybrid swarm in the genus Betula, which has a distribution very similar to
that of the P. machaon X zelicaon swarms in central Alberta (Dugle, 1966).
Some hybrid zones appear to have moved a few dozen kilometers within the last century
( e.g ., McDonnell et al., 1978). However, such movement is uncommon, and most hybrid zones
appear to remain relatively stationary over long periods of time. In fact, many hybrid zones
probably become trapped in regions of low density, such as habitat dines and natural barriers
(Barton, 1979). They may also become narrower, especially if there is strong selection against
hybrids, or they may widen and eventually result in the merging of the two parental species. In
the P. machaon group, evidence suggests that some hybrid zones may presently be undergoing
change, probably because they are very recent, having been influenced by human settlement
patterns. The range expansion of P. polyxenes into Nova Scotia and Manitoba is probably
related to deforestation by man, and so it seems likely that any interaction of this species with
P. brevicauda, in the east, and P. m. hudsonianus , in the west, is less than a century old. The
decreasing proportion of P. machaon in Riding Mountain Park in Manitoba is thus probably a
result of genetic swamping by recently arrived P. polyxenes in agricultural areas.
Most hybrid zones for both plants and animals, appear to be the result of secondary contact
between formerly allopatric species {e.g., Remington, 1968b; Barton and Hewitt, 1985).
However, a few hybrid zones may be the result of in situ differentiation on either end of a sharp
environmental gradient (Endler, 1977). Most authors do not believe that they can distinguish
between these two situations, though Thorpe (1984) states that a phylogenetic analysis at the
population level makes such distinctions possible. In the P. machaon group it seems most likely
to me that most, if not all, of the hybrid zones can most parsimoniously be explained as the
result of post-Pleistocene range expansions. However, if P. m. hudsonianus spent the late
Wisconsinan south of the continental ice sheet, then there may have been a pattern of contact
between P. m. hudsonianus and P. p. asterius which was similar to the present one. The contact
zone between P. zelicaon and P. m. aliaska is certainly the result of secondary contact.
Hybrid zones which show substantial gene flow are generally no longer considered to
represent interspecific hybridization, but rather zones of contact between different races of a
single species. Examples include subspecies within both P. machaon and P. polyxenes in the
western United States. However, the degree of gene flow has only been indirectly interpreted
from morphological and ecological character gradients and could bear rechecking against
enzyme allele distributions. In particular, it should be interesting to compare the rate of gene
flow between P. p. asterius and P. p. color o in New Mexico with that between P. p. color o and
P. zelicaon in southern California. Enzyme data for western Canada show a significant
interruption in gene flow between P. machaon and P. zelicaon in most regions in Alberta and
British Columbia.
Though both electrophoretic and morphometric character distributions can indicate gene
flow across hybrid zones, there may be some differences between these character types. This is
true of the P. machaon group hybrid populations in both central Alberta and Manitoba, where
electrophoretic character combinations showed greater intermediacy than did morphometric
characters. Harrison (1986) reported a similar situation in a hybrid zone in crickets, and
suggested that there were fewer barriers to the introgression of allozyme alleles than
morphometric characters. A possible reason for this is that the inheritance of morphometric
characters is more canalized, with a greater degree of linkage between genes and a resultingly
Papilio machaon species group
289
greater resistance to the movement of such traits.
The evolutionary importance of interspecific hybridization is not clear, though various
authors have suggested that gene introgression provides an important source of allelic variation
for action of natural selection. However, most studies of gene flow in hybrid zones show only
limited intrusions of alleles into neighboring species (Barton and Hewitt, 1985). The black
morph in the P. machaon group generally follows this pattern as well, though it has moved
several hundred kilometers into the range of P. zelicaon and has displaced the yellow allele in
the southern part of the range of P. machaon.
The selective advantage of alleles which produce the black adult morph is unknown. Since
many populations are polymorphic with respect to this allele, it is probably not important as a
visual mechanism for mate recognition, as Hafernik (1982) reported for an analogous wing
pattern in Junonia (Nymphalidae). It may give hilltopping males an advantage in maintaining
a position at the very peak of hills (Scott, 1983 and personal observations). However, Miller
(1977) suggested that the allele is lethal when homozygous and in combination with the P.
zelicaon genome. Perhaps the distribution of the allele is the result of an equilibrium between
positive and negative selection, much as hybrid zones themselves may be a balanced conflict
between genes that widen zones by reducing incompatibilities and genes that narrow zones by
producing reproductive isolation (Barton and Hewitt, 1981).
Hybrid populations. — Some populations in interspecific hybrid zones are characterized by
negative or neutral selection on hybrid individuals in the contact zone. However, hybrid
populations are characterized by positive selection for interspecific hybrids in restricted areas,
even though the parental species retain their integrity over most of their area of contact. Hybrid
populations are composed predominantly of hybrid forms, and variation within most such
populations spans the full range of phenotypes between the parental forms. In a fully integrated
hybrid population, individuals phenotypically similar to parental forms simply represent the
phenotypic extremes within the population.
Hybrid populations are reasonably common in plants, with many referred to as hybrid
swarms, but are very unusual in animals (Mayr, 1963; Grant, 1971). Most animal examples are
of birds, amphibians and fish (Moore, 1977), and the studies of Sibley (1954) on towhees and
those of Blair (1941) and others on toads are still among the best documented. Examples of
hybrid populations not clearly associated with narrow zones are much less common in insects.
The present study provides a clear example of hybrid populations between broadly
sympatric species. In the P. machaon group the best examples of hybrid swarms are the P.
machaon X zelicaon populations in central Alberta, particularly the one at Bragg Creek. The
Bragg Creek population is composed of a highly varied but unimodal population made up
almost completely of hybrid forms, and probably has no significant internal impediment to gene
flow. The intermediate nature of the central Alberta populations is indicated by both the
morphometric and electrophoretic character distributions.
Many hybrid populations are associated with environmental disturbance of some sort,
resulting in a kind of hybrid habitat, in which forms intermediate between the parental species
can flourish. Habitat disturbance by man in recent times, has provided several opportunities to
observe the formation of new hybrid swarms over a period of only a few decades ( e.g ., Gillespie,
1985). An interesting aspect of some hybrid populations is that they separate again into
parental forms within about 20 years (Jones, 1973; Corbin et al., 1979). Considering the
ephemerality and dependence on habitat disturbance of many hybrid swarms, the examples of
the P. machaon group from central Alberta are fairly unusual. They occur in areas with
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290
Sperling
relatively little or no habitat disturbance, especially compared to the regions dominated by
agriculture where P. machaon and P. zelicaon coexist with only a small amount of
hybridization. As well, material collected around 1900 suggests that the hybrid populations
were already in existence when central Alberta was first being settled.
Although artificial hybrids within the P. machaon group have invariably been obtained
between individuals from geographically distant populations, and many showed substantial
infertility even when they were from interspecific populations clearly connected by character
dines, some backcrosses to either of the parental species have produced viable adults (Clarke
and , 1953; Ae, 1966; Clarke et al., 1977). In a few crosses, adults have even been obtained
from an F2 hybrid cross of P. polyxenes and P. machaon (Ae, 1964). These experiments
indicate that introgression and the formation of hybrid swarms are at least possible, though
unlikely.
However, at least part of the inviability of hybrids is due to environmental adaptations, such
as diapause characteristics (Oliver, 1969). Since the three species in western Canada are very
flexible in their adaptation to different environmental factors, it is reasonable to expect that
species coming together in a particular region tend to be more similar than would populations
from more distant regions. Unless some fundamentally different ecological adaptation has
occurred, adjacent populations seem likely to meet and hybridize on a continuing basis, until
reproductive isolation occurs. Thus the reason for formation of P. machaon X zelicaon hybrid
swarms in central Alberta is probably related to similarities in the habitat preferences of local
races before contact occurred through range expansions. The two species coexist where P.
machaon larvae feed on Artemisia and where adult contact is reduced through the occupation
of different habitats. Where P. machaon occupies a habitat more similar to that of P. zelicaon ,
as does P. m. hudsonianus, the two species have tended to merge, with the hybrid populations
feeding on plesiotypically palatable umbellifers. This situation is similar to that described by
Mayr (1963) for Passer domesticus (Linneaus) and P. hispaniolensis Temminck in Europe.
Many plants form hybrid swarms, and the frequency with which such events occur may be
related to ecological characteristics of particular taxa (Raven, 1976). Some plant species tend
to maximize the saturation density ( K ) of their populations and are separated by ecological and
other extrinsic factors, but have only slightly developed internal barriers to hybridization. They
hybridize with related species to form new recombinants, which allow populations to adapt to
changing environments. On the other hand, species whose populations maximize their rate of
increase (r), such as annual herbs, tend to hybridize much less frequently with each other.
Since they are characterized by rapid dipersal and growth in new areas, as well as a high
commitment of basic resources to reproduction, barriers to hybridization are much more
important to these species. This correspondence between maximization of saturation density
and tendency toward hybridization in plants does not seem to apply to the P. machaon group.
These butterflies would, if anything, be considered as maximizing their rate of increase, since
they feed in the larval stage on successional plants and are dependent on rapid colonization and
foodplant exploitation. However, it would be interesting to investigate the P. machaon group to
determine if introgression enhances adaptation to new ecological conditions.
The phylogenetic significance of hybrid swarm formation is uncertain. It may be rare
enough in animals so that it has had little influence on evolutionary patterns. However, it may
be that such breakdowns in reproductive barriers contribute to the formation of new
populations under conditions in which one of the parental species would have been eliminated
by habitat destruction. In this way part of the threatened gene pool is saved, albeit in a greatly
Papilio machaon species group
291
altered combination. Formation of hybrid populations may also cause a major disorganization
of the polygenic balances of the parental species, leading eventually to speciation through a
major new balanced genetic system (Carson, 1981).
New species that may have arisen from interspecific hybrid swarms would be impossible to
detect by morphological features if the hybridization occurred between a pair of sibling species.
On the other hand, if the new species is the product of hybridization between species A and C,
and there exists a species B which is more closely related to A than C is, then the hybrid origin
of the new species would be indicated by its discordant character distribution. Unfortunately,
as Mayr (1963) pointed out, such a character distribution could also easily be due to the
character convergences and parallelisms which one would expect in closely related species with
a very similar basic gene pool. For these reasons, the number of animal taxa which have had a
hybrid origin has almost certainly been underestimated, and will continue to remain so until
there has been ample opportunity to support or reject present taxonomic assignments with
independent character suites, such as enzyme alleles or mitochondrial DNA.
ACKNOWLEDGEMENTS
This paper is derived from my MSc thesis, at the University of Alberta. Its completion owes
much to the continued support and encouragement of my thesis advisor, G.E. Ball, as well as
the friendship of J.H. Acorn, G.J. Hilchie, D.R. Maddison, E.M. Pike and J.R. Spence. My
parents also contributed to this work in many ways, few of them obvious.
I thank all the individuals whose names are listed in Table 1 for allowing me to examine
specimens from their personal collections or in their care. J.C. Daniels, C.S. Guppy, G.J.
Hilchie, H.P. Kimmich, N.G. Kondla, I. Laing, E.M. Pike, K.A. Shaw and J.T. Troubridge
made special efforts to collect regionally important material for me. R.H. Gooding and B.M.
Rolseth helped with electrophoresis, and D.R. Maddison gave advice on computer work.
Taxonomic identifications for plant material were provided by J.G. Packer and A. A. Rose.
Important records and photos were made available to me by G. Anweiler, C.D. Bird, C.S.
Guppy, R.R. Hooper, H.P. Kimmich, P. Klassen, J.A. Scott, J.H. Shepard and J.T.
Troubridge. Comments on the thesis version of this paper were provided by J.F. Addicott, D.A.
Craig, J.G. Franclemont, R.H. Gooding, and especially G.E. Ball and J.R. Spence.
Improvements to the submitted manuscript were suggested by R.G. Harrison, and two
reviewers whose comments were transmitted to me by the editor. I am most grateful to all of
these individuals.
Most of this study was funded by NSERC Grant A- 1399 to G.E. Ball. Travel expenses for
the summer of 1982 were provided by a grant to me from the Boreal Institute for Northern
Studies, University of Alberta.
LITERATURE CITED
Ae, S.A. 1961. A study of interspecific hybrids in Papilio. Tyo to Ga 12:65-89.
Ae, S.A. 1964. A study of hybrids in the Papilio machaon group. Bulletin of the Japan
Entomological Academy 1:1-10.
Ae, S.A. 1966. A further study of hybrids in the Papilio machaon group. Bulletin of the Japan
Entomological Society 2:19-30.
Ae, S.A. 1979. The phylogeny of some Papilio species based on interspecific hybridization
Quaest. Ent., 1987, 23 (2)
292
Sperling
data. Systematic Entomology 4:1-16.
Barton, N.H. 1979. The dynamics of hybrid zones. Heredity 43:341-59.
Barton, N.H. and G.M. Hewitt. 1981. Hybrid zones and speciation, pp. 109-145. In , Evolution
and Speciation. (Atchley, W.R. and D.S. Woodruff, Editors). Cambridge University Press,
Cambridge. 436 pp.
Barton, N.H. and G.M. Hewitt. 1985. Analysis of hybrid zones. Annual Review of Ecology and
Systematics 16:113-148.
Berenbaum, M. 1978. Taenidia integerrima, a new foodplant record for Papilio polyxenes
(Papilionidae). Journal of the Lepidopterists’ Society 32:303-304.
Berenbaum, M. 1981. Effects of linear furanocoumarins on an adapted specialist insect
{Papilio polyxenes). Ecological Entomology 6:345-351.
Berenbaum, M. 1983. Coumarins and caterpillars: a case for coevolution. Evolution
37:163-179.
Bird, C.D., G.J. Hilchie, N.G. Kondla, W.W. Smith, E. Kuyt, J.K. Ryan, and T.W. Thormin.
1982. Butterflies of northeastern Alberta. Blue Jay 40:141-152.
Blackwelder, R.E. 1967. Taxonomy. Wiley and Sons, New York. 698 pp.
Blair, A.P. 1941. Variation, isolating mechanisms, and hybridization in certain toads. Genetics
26:398-417.
Blau, W.S. 1980. The effect of environmental disturbance on a tropical butterfly population.
Ecology 61:1005-1012.
Blau, W.S. 1981a. Life history variation in the black swallowtail butterfly. Oecologia
48:116-122.
Blau, W.S. 1981b. Notes on the natural history of Papilio polyxenes stabilis (Papilionidae) in
Costa Rica. Journal of the Lepidopterists’ Society 34:321-324.
Blau, W.S. 1981c. Latitudinal variation in the life histories of insects occupying disturbed
habitats: a case study, pp. 75-96. In , Insect life history patterns: habitat and geographic
variation (R.F. Denno and H. Dingle, Editors). Springer Verlag, New York. 225 pp.
Boisduval, J. 1852. Lepidopteres de la Californie. Annales de la Societe entomologique de
France. Series 2. 10:275-324.
Brewer, G.J. 1970. Chapter 5, Specific electrophoretic systems, pp. 62-137. In , An
Introduction to Isozyme Techniques (G.J. Brewer and C.F. Sing, Editors). Academic Press,
New York, xii + 180 pp.
Brown, F.M. 1975. The types of Papilionid butterflies named by William Henry Edwards.
Transactions of the American Entomological Society 101:1-31
Buth, D.G. 1984. The application of electrophoretic data in systematic studies. Annual Review
of Ecology and Systematics 15:501-522.
Canadian Climate Normals. 1951-1980; 1982a. Volume 2. Temperature. Environment
Canada, Ottawa. 306 pp.
Canadian Climate Normals. 1951-1980; 1982b. Volume 3. Precipitation. Environment
Canada, Ottawa. 602 pp.
Canadian Climate Normals. 1951-1980; 1982c. Volume 4. Degree days. Environment Canada,
Ottawa. 280 pp.
Carson, H.L. 1981. Speciation as a major reorganization of polygenic balances, pp. 41 1-433.
In , Mechanisms of Speciation (C. Barigossi, Editor). A.R. Liss, Inc., New York, 546 pp.
Chermock, F.H. and R.L. Chermock. 1937. Two new races of Papilios from Manitoba. Bulletin
of the Southern California Academy of Sciences 36:1 1-13.
Papilio machaon species group
293
Clark, A.H. 1932. The forms of the common old world swallowtail butterfly ( Papilio machaon)
in North America, with descriptions of two new subspecies. Proceedings of the United
States National Museum 81 (Article 1 1): pp.1-15, plates 1-8.
Clarke, C.A. 1952. Handpairing of Papilio machaon in February. Entomologist’s Record
64:98-100.
Clarke, C.A. and J.P. Knudsen. 1953. A hybrid swallowtail. Entomologist’s Record 65:76-80
+ plates 4-10.
Clarke, C.A. and T.B. Larsen. 1986. Speciation problems in the Papilio machaon group of
butterflies (Lepidoptera: Papilionidae). Systematic Entomology 11:175-181.
Clarke, C.A.; U. Mittwoch and W. Traut. 1977. Linkage and cytogenetic studies in the
swallowtail butterflies Papilio polyxenes Fab. and Papilio machaon and their hybrids.
Proceedings of the Royal Society of London. B. Biological Sciences 198:385-399.
Clarke, C.A. and P.M. Sheppard. 1953. Further observations on hybrid swallowtails.
Supplement to the Entomologist’s Record 65(9): 1-12.
Clarke, C.A. and P.M. Sheppard. 1955a. The breeding in captivity of the hybrid swallowtail
Papilio machaon gorganus Friihstorfer (F) X Papilio hospiton Gene (M). The
Entomologist 88:265-268 + plates 7-10.
Clarke, C.A. and P.M. Sheppard. 1955b. A preliminary report on the genetics of the machaon
group of swallowtail butterflies. Evolution 9:182-201.
Clarke, C.A. and P.M. Sheppard. 1956a. A further report on the genetics of the machaon
group of swallowtail butterflies. Evolution 10:66-73.
Clarke, C.A. and P.M. Sheppard. 1956b. Handpairing of butterflies. Lepidopterists’ News
10:47-53.
Clarke, C.A. and P.M. Sheppard. 1970. Is Papilio gothica (Papilionidae) a good species?
Journal of the Lepidopterists’ Society 24:229-233.
Clarke, C.A., P.M. Sheppard and A. Willig. 1972. The use of ecdysone to break a two and a
half year pupal diapuase in Papilio glaucus female X Papilio rutulus male hybrids. The
Entomologist 105:137-138.
Corbin, K.W., C.G. Sibley and A. Ferguson. 1979. Genic changes associated with the
establishment of sympatry in orioles of the genus Icterus. Evolution 33:624-633.
Darwin, C. 1859. On the Origin of Species by Means of Natural Selection (6th edition, 1972).
Murray. London.
Dempster, J.P. and M.L. Hall. 1980. An attempt at re-establishing the swallowtail butterfly at
Wicken Fen. Ecological Entomology 5:327-334.
Dethier, V.G. 1941. Chemical factors determining the choice of foodplants by Papilio larvae.
American Naturalist 75:61-73.
Dierl, W. 1976. Distribution and isolation factors in the races of Papilio machaon
(Lepidoptera: Papilionidae) in central Himalaya. Proceedings of the XV International
Congress of Entomology 1976:171-172.
Dobzhansky, T., F.J. Ayala, G.L. Stebbins and J.W. Valentine. 1977. Evolution. Freeman and
Co., San Fransico, 572 pp.
Dornfeld, E.J. 1980. The Butterflies of Oregon. Timber Press. Forest Grove, Oregon. 276 pp.
Dos Passos, C.F. 1962. The authorship of three scientific names of nearctic Rhopalocera
variously credited to Boisduval or Lucas. Journal of the Lepidopterists’ Society 16:45-46.
Dugle, J.R. 1966. A taxonomic study of western Canadian species in the genus Betula.
Canadian Jounal of Botany 44:929-1007.
Quaest. Ent., 1987, 23 (2)
294
Sperling
Edwards, W.H. 1866. Proceedings of the Entomological Society of Philadelphia 6:200.
Edwards, W.H. 1876. Description of new species of diurnal Lepidoptera, found within the
United States and British N.A. Transactions of the American Entomological Society
5:202-208.
Edwards, W.H. 1883. Comments on a paper entitled “The genus Colias from Proc. Bost.
Soc.., N.H., Vol. XXII. Papilio 3:169-185.
Edwards, W.H. 1884. Notes upon a small collection of butterflies , made in Judith Mtns.,
Montana, in 1883, by Wm. M. Courtis, M.E. Papilio 3:157-164.
Edwards, W.H. 1893. Notes on a polymorphic butterfly. The Canadian Entomologist
25:253-254.
Edwards, W.H. 1895. Notes on collecting butterflies in western Colorado, with a particular
account of certain Papilios. The Canadian Entomologist 27:229-242.
Edwards, W.H. 1898. The butterflies of North America. Third Series. Houghton, Mifflin and
Company. Boston and New York.
Ehrlich, P.R. 1961. Has the biological species concept outlived its usefulness? Systematic
Zoology 10:167-176.
Ehrlich, P.R. and P. Raven. 1964. Butterflies and plants: a study in co-evolution. Evolution
18:586-608.
Eller, K. 1936. Die Rassen von Papilio machaon L. Abhandlungen der Bayerischen Akademie
der Wissenschaften. Mathematischnaturwissenschaftliche Abteilung. Neue Folge. Heft 36,
96 pp., 16 pi.
Eller, K. 1939. Fragen und Probleme zur Zoogeographie und zur Rassen- und Artbildung in
der Papilio machaon Gruppe. Seventh International Congress of Entomology 1:74-101.
Emmel, J.F. 1975. Papilioninae, pp. 390-402. In , The Butterflies of North America.
Doubleday and Co., New York, 633 pp. + 97 plates.
Emmel, J.F. and T.C. Emmel. 1963. Larval food-plant records for six western Papilios. Journal
for Research on the Lepidoptera 1:191-193.
Emmel, J.F. and O. Shields. 1980. Larval foodplant records for Papilio zelicaon in the western
United States and further evidence for the conspecificity of P. zelicaon and P. gothica.
Journal of Research on the Lepidoptera 17:56-67.
Emmel, T.C. and J.F. Emmel. 1969. Selection and host plant overlap in two desert Papilio
butterflies. Ecology 50:158-159.
Emmel, T.C. and J.F. Emmel. 1973. The Butterflies of southern California. Los Angeles City
Museum of Natural History Science Series 26: 1-148.
Endler, J.A. 1977. Geographic Variation, Speciation and Clines. Princeton University Press.
Princeton, New Jersey. 246 pp.
Fabricius, J.C. 1775. Systema Entomologiae. Lipsiae, 832 pp.
Feeny, P., W.S. Blau and P.M. Kareiva. 1985. Larval growth and survivorship of the black
swallowtail butterfly in central New York. Ecological Monographs 55:167-187.
Ferguson, D.C. 1954. The Lepidoptera of Nova Scotia. Proceedings of the Nova Scotia
Institute of Science 23: part 3:161-375.
Ferris, C.D. and J.F. Emmel. 1982. Discussion of Papilio coloro W.G. Wright ( = Papilio
rudkini F. & R. Chermock) and Papilio polyxenes Fabricius (Papilionidae). Bulletin of the
Allyn Museum 76:1-13.
Fisher, M.S. 1977. The taxonomy and identity of Papilio nitra W.H. Edwards in Colorado
(Papilionidae). Bulletin of the Allyn Museum No. 47, pp.1-7.
Papilio machaon species group
295
Fisher, M.S. 1980. Subfamily Papilioninae. In, Butterflies of the Rocky Mountain States.
University of Oklahoma Press. Norman, Oklahoma, p. 178-193.
Freeman, T.N. 1949. Far North - Alaska to Labrador. Lepidopterists’ News 3:101.
Freeman, T.N. 1972. A correlation of some butterfly distributions with geological formations.
The Canadian Entomologist 104:443-444.
Fox, D.J. and K.E. Guire. 1976. Documentation for Midas. Statistical Research Laboratory.
University of Michigan. 203 pp.
Gillespie, G.D. 1985. Hybridization, introgression and morphometric differentiation between
mallard ( Anas platyrhynchos ) and grey duck ( Anas superciliosa ) in Otago, New Zealand.
The Auk 102:459-469.
Gooding, R.H. and B. Rolseth. 1979. Genetics of Glossina morsitans morsitans
(Diptera:Glossinidae) IV. Electrophoretic banding patterns of octanol dehydrogenase and
arginine phosphokinase. The Canadian Entomologist 1 1 1:1307-1310.
Grant, V. 1971. Plant Speciation. Columbia University Press, New York. 435 pp.
Gunder, J.D. 1928. Additional transitional forms (Lepid., Rhopalocera). Canadian
Entomologist 60:162-168, 2 plates.
Guppy, R. 1970. Further observations on “hilltopping” in Papilio zelicaon. Journal for
Research on the Lepidoptera 8:105-1 17.
Hafernik, J.E. 1982. Phenetics and ecology of hybridization in buckeye butterflies. University
of California Publications in Entomology Vol. 96, 109 pp.
Hagen, H.A. 1882. On Papilio machaon L., and its N. American representatives-; Papilio
rutulus Bvd.-; and Parnassius ; Being portion of a preliminary report on the butterflies of
Washington Territory. Papilio 2:149-164.
Hancock, D.L. 1983. Classification of the Papilionidae (Lepidoptera): a phylogenetic approach.
Smithersia No. 2:1-48.
Hansen, H.P. 1949. Postglacial forests in south central Alberta, Canada. American Journal of
Botany 36:54-65.
Hansen, H.P. 1950. Postglacial forests along the Alaska Highway in British Columbia.
Proceedings of the American Philosophical Society 94:41 1-421.
Hansen, H.P. 1955. Postglacial forests in south-central and central British Columbia.
American Jounal of Science 253:640-658.
Harrison, R.G. 1986. Pattern and process in a narrow hybrid zone. Heredity 56:337-349.
Heitzman, J.R. 1973. A new species of Papilio from the Eastern United States (Papilionidae).
Journal of Research on the Lepidoptera. 12:1-10.
Heron, R.J. and A.G. Robinson. 1976. A field survey of the butterflies and skippers of Riding
Mountain National Park. Department of Entomology, University of Manitoba, Winnipeg.
115 pp.
Higgins, L.G. 1975. The Classification of European Butterflies. Collins, London. 320 pp.
Higgins, L.G. and N.D. Riley. 1970. Butterflies of Britain and Europe. Collins, London. 381
pp.
Hooper, R.R. 1973. The Butterflies of Saskatchewan. Saskatchewan Department of Natural
Resources. Regina. 216 pp.
Hull, D.L. 1968. The operational imperative: sense and nonsense in operationalism. Systematic
Zoology 17:438-457.
International Code of Botanical Nomenclature. 1983. Bohn, Scheltema and Holkema, Utrecht,
xv + 472 pp.
Quaest. Ent., 1987, 23 (2)
296
Sperling
International Code of Zoological Nomenclature. 1985. University of California Press, Berkeley
and Los Angeles, xx + 338 pp.
Jackson, B.S. 1982. Recorded host plants and nectar sources of the short-tailed swallowtail
(Papilio brevicauda Saunders) (Lepidoptera: Papilionidae) in Newfoundland, Canada.
Atala 8:43.
Jones, J.H. 1973. Effects of thirty years hybridization on the toads Bufo americanus and Bufo
woodhousii fowleri at Bloomington, Indiana. Evolution 27:435-448.
Kettlewell, H.B.D. 1955. Natural history of P. hospiton Gene in Corsica. The Entomologist
88:280-283.
Key, K.H.L. 1982. Species, parapatry and the morabine grasshoppers. Systematic Zoology
30:425-458.
Kimmich, H.P. 1979. Life histories anonymous: Papilio machaon aliaska. Lepidopterists’
News 1979(5):9.
Kondla, N.G. 1981. Type localities of the Badlands Old World Swallowtail in Alberta. Blue
Jay 39:144.
Kuhn, T.S. 1970. The Structure of Scientific Revolutions. Second Edition. University of
Chicago Press, Chicago. 210 pp.
Kurentzov, A.I. 1970. The Butterflies of the Far East U.S.S.R.. Academy of Sciences of
U.S.S.R. (Siberian Division), Institute of Pedobiology of the Far Eastern Branch. 163 pp.,
14 pi. (Translations obtained from K. Phillip).
Larsen, T.B. 1980. The status of Papilio machaon rathjensi and its relationship to other
Arabian populations (Papilionidae). Journal of the Lepidopterists’ Society 34:365-367.
Leussler, R.A. and O. Bryant. 1935. Notes on the diurnal Lepidoptera of the Canadian arctic
collected by Owen Bryant in the summers of 1929 to 1932. Bulletin of the Brooklyn
Entomological Society 30:42-62.
Linnaeus, C. (C. von Linne). 1758. Caroli Linnaei Systema Naturae, Regnum Animale, 10th
Edition. British Museum (Natural History), London. (Reprinted 1939).
Llewllyn Jones, J.R.J. 1951. An annotated list of the macrolepidoptera of British Columbia.
Entomological Society of British Columbia, Occasional Paper No. 1, 148 pp.
Lucas, P.H. 1852. Description de nouvelles especes de Lepidopteres appartenant aux collections
entomologiques du Musee de Paris. Revue et Magasin de Zoologie Pure et Applique. Series
2. 4:128-141.
Maeki, K. 1976. On the chromosomes of Papilio machaon hippocrates C. et R. Felder. Tyo To
Ga 26:85-90.
Maeki, K. and C.L. Remington. 1960. Studies on the chomosomes of North American
Rhopalocera. 1. Papilionidae. Jounal of the Lepidopterists’ Society 13:193-203.
Matthews, J.V. Jr. 1979. Tertiary and Quaternary environments: historical background for an
analysis of the Canadian insect fauna, pp. 31-86. In, Canada and its insect fauna. (H.V.
Danks, Editor). Memoirs of The Entomological Society of Canada No. 108, 573 pp.
Matthews, J.V. Jr. 1982. East Beringia during Late Wisconsinan time:a review of the evidence,
pp. 127-150. In, Paleoecology of Beringia. (D.M. Hopkins, J.V. Matthews, Jr., C.E.
Schweger, and S.B. Young, Editors). Academic Press, New York. 489 pp.
Mayr, E. 1955. Karl Jordan’s contribution to current concepts in systematics and evolution.
Transactions of the Royal Entomological Society of London 107:45-66.
Mayr, E. 1963. Animal Species and Evolution. Belknap Press, Cambridge. 797 pp.
Mayr, E. 1969. Principles of systematic zoology. McGraw Hill, New York. 428 pp.
Papilio machaon species group
297
McDonnell, L.J., D.F. Gartside and M.J. Littlejohn. 1978. Analysis of a narrow hybrid zone
between two species of Pseudophryne (Anura:Leptodactylidae) in southeastern Australia.
Evolution 32:602-612.
McDunnough, J. 1927. The Lepidoptera of the Seton Lake region, British Columbia. The
Canadian Entomologist 59:152-162.
McDunnough, J. 1939. A new race of Papilio belonging to the machaon complex. The
Canadian Entomologist 71:216-217.
Menken, S.B.J. 1980. Allozyme polymorphism and the speciation process in small ermine
moths (Lepidoptera, Yponomeutidae). Studies in Yponomeuta 2. PhD. Thesis,
Rijkuniversiteit te Leiden. 1 19 pp.
Mickevich, M.F. and C. Mitter. 1981. Treating polymorphic characters in systematics: a
phylogenetic treatment of electrophoretic data, pp. 45-58. In , Advances in Cladistics -
Proceedings of the First Meeting of the Willi Hennig Society, (V.A. Funk and D.R. Brooks,
Editors). New York Botanical Garden, Pubs.
Miller, J.S. 1986. Phylogenetic systematics and chemical constraints on host-plant associations
in the Papilioninae (Lepidoptera: Papilionidae). PhD Thesis, Cornell University, Ithaca,
New York. 320 pp.
Miller, L.D. 1977. Appendix to The taxonomy and identity of Papilio nitra W.H. Edwards in
Colorado (Papilionidae). Bulletin of the Allyn Museum 47:7-8.
Miller, L.D. 1983. Butterfly taxonomy: a reply. Journal of Research on the Lepidoptera
20:193-198.
Miller, L.D. and F.M. Brown. 1981. A catalogue/checklist of the butterflies of America north
of Mexico. Lepidopterists’ Society Memoir No. 2, 280 pp.
Moore, W.S. 1977. An evaluation of narrow hybrid zones in vertebrates. Quarterly Review of
Biology 52:263-277.
Monroe, E.G. 1961. The classification of the Papilionidae (Lepidoptera). The Canadian
Entomologist, Supplement 17: 51 pp.
Morris, R.F. 1980. Butterflies and Moths of Newfoundland and Labrador. Agriculture Canada
Publication No. 1691, Hull, Quebec, 407 pp.
Muting, D. 1972. Ritt zum Pamir. Zoologische Eindriicke aus Nord-Afghanistan.
Entomologische Zeitschrift 82:33-44.
Nakamura, I. and S.A. Ae. 1977. Prolonged pupal diapause of Papilio alexanor. arid zone
adaptation directed by larval host plant. Annals of the Entomological Society of America
70:481-484.
Neff, N.A. and G.R. Smith. 1979. Multivariate analysis of hybrid fishes. Systematic Zoology
28:176-196.
Nei, M. 1972. Genetic distance between populations. American Naturalist 106:283-292.
Nelson, G. and N. Platnick. 1981. Systematics and Biogeography, Cladistics and Vicariance.
Columbia University Press, New York. 567 pp.
Newcomer, E.J. 1964. Life histories of Papilio indra and P. oregonius. Journal for Research on
the Lepidoptera 3:49-54.
Oliver, C.G. 1969. Experiments on the diapause dynamics of Papilio polyxenes. Journal of
Insect Physiology 15:1579-1589.
Opler, P.A. and G.O. Krizek. 1984. Butterflies East of the Great Plains. The Johns Hopkins
University Press, Baltimore. 294 pp.
Perkins, S.F., E.M. Perkins, Jr. and F.S. Shininger. 1968. Illustrated life history and notes on
Quaest. Ent., 1987, 23 (2)
298
Sperling
Papilio oregonius. Journal of the Lepidopterists’ Society 22:53-56.
Pike, E.M. 1980. Origin of tundra butterflies in Alberta. Quaestiones Entomologicae
16:555-596.
Pimentel, R.A. 1981. A comparative study of data and ordination based on a hybrid swarm of
sand verbenas ( Abronia Juss.). Systematic Zoology 30:250-267.
Raven, P.H. 1976. Systematics and plant population biology. Systematic Botany 1:284-316.
Reist, J.D. 1979. Callophrys niphon (Lycaenidae) in Alberta with notes on the identification of
C. niphon and C. eryphon. Journal of the Lepidopterists’ Society 33:248-253.
Remington, C.L. 1956. Interspecific relationships of two rare swallowtail butterflies, Papilio
nitra and Papilio hudsonianus, to other members of Papilio machaon complex. American
Philosophical Society Yearbook 1955:142-146.
Remington, C.L. 1958. Genetics of populations of Lepidoptera. Proceedings of the Tenth
International Congress of Entomology 2:787-805.
Remington, C.L. 1968a. A new sibling Papilio from the Rocky Mountains, with genetic and
biological notes (Insecta, Lepidoptera). Postilla 1 19:1-40.
Remington, C.L. 1968b. Suture-zones of hybrid interaction between recently joined biotas.
Evolutionary Biology 2:321-428.
Richard, D. and M. Guedees. 1983. The Papilionidae (Lepidoptera): coevolution with the
Angiosperms. Phyton 23:1 17-126.
Rolseth, B.M. and R.H. Gooding. 1978. Genetics of Glossina morsitans morsitans (Diptera:
Glossinidae). I. Electrophoretic banding patterns of xanthine oxidase and aldehyde oxidase.
The Canadian Entomologist 110: 1233-1239.
Rothschild, M. 1983. Dear Lord Rothschild: Birds, Butterflies and History. Hutchinson,
London, xx + 398 pp.
Rothschild, W. and K. Jordan. 1906. A revision of the American Papilios. Novitates Zoologicae
13:411-752.
Rumbacher, K. and H. Seyer. 1979. Verbreitung und Wanderwege der Papilio machaon
Untergruppen; Versuch einer Graphischen Darstellung. Entomologische Gesellschaft Basel
29:44-49.
Schenk, E.T. and J.H. McMasters. 1956. Procedure in Taxonomy, third edition. Stanford
University Press, Stanford, California. 1 19 pp.
Schweger, C., T. Habgood and M. Hickman. 1981. Late glacial -holocene climatic changes of
Alberta. The record from lake sediment studies. Proceedings of the Alberta Climatological
Association Meetings, 1981.
Scott, J.A. 1981. New Papilionoidea and Hesperioidea from North America. Papilio (new
series) 1:1-12 (privately published by the author).
Scott, J.A. 1983. Mate-locating behavior of western North American butterflies. II. New
observations and morphological adaptations. Journal of Research on the Lepidoptera
21:177-187.
Scriber, J.M. 1982. Loodplants and speciation in the Papilio glaucus group. Proceedings of the
5th International Symposium on Insect-Plant Relationships 1982:307-314.
Scriber, J.M. and P. Leeney. 1979. Growth of herbivorous caterpillars in relation to feeding
specializations and to the growth form of their foodplants. Ecology 60:829-850.
Scudder, G.G.E. 1979. Present patterns in the fauna and flora of Canada, pp. 87-179. In,
Canada and Its Insect Launa. (H.V. Danks, Editor). Memoirs of the Entomological Society
of Canada, No. 108. 573 pp.
Papilio machaon species group
299
Scudder, S.H. 1869. Report upon a collection of diurnal Lepidoptera made in Alaska by the
scientific corps of the Russo-American Telegraph Expedition under the direction of Lieut.
W.H. Dali. Proceedings of the Boston Natural History Society. 12:404-408.
Seyer, H. 1974. Versuch einer Revision der Papilio machaon- Subspezies in der Westlichen
Palaarktis. Mitteilungen der Entomologischen Gesellschaft Basel 24:64-1 17.
Seyer, H. 1976a. Versuch einer Revision der Papilio machaon- Subspezies in der Oestlichen
Palaarktis. Mitteilungen der Entomologischen Gesellschaft Basel 26:65-82.
Seyer, H. 1976b. Versuch einer Revision der Papilio machaon- Subspezies in der Oestlichen
Palaarktis. 2. Teil und Schluss. Mitteilungen der Entomologischen Gesellschaft Basel
26:97-145.
Seyer, H. 1977. 2. Nachtrag zum Versuch einer Revision der Papilio machaon-SubspQziQS in
der Palearktis und 1 . Beitrag zur Nearktis. Mitteilungen der Entomologischen Gesellschaft
Basel 27:105-115.
Seyer, H. 1982. Verwandschaft und Arealgenese der Papilio mac/mcw-Population in der
westlichen Palaarktis. PhD Dissertation, Universitat des Saarlandes. 1 15 pp.
Shapiro, A.M. 1975. Papilio "gothica” and the phenotypic plasticity of P. zelicaon
(Papilionidae). Journal of the Lepidopterists’ Society 29:79-84.
Shapiro, A.M. and K.K. Masuda. 1980. The opportunistic origin of a new citrus pest.
California Agriculture 36:4-5.
Shapiro, A.M., C.A. Palm and K.L. Weislo. 1981. The ecology and biogeography of the
butterflies of the Trinity Alps and Mount Eddy, northern California. Journal of Research on
the Lepidoptera 18:69-152.
Shaw, C.R. and R. Prasad. 1970. Starch gel electrophoresis of enzymes - A compilation of
recipes. Biochemical Genetics 4:297-320.
Shields, O. 1967. An ecological study of summit congregation behavior of butterflies on a
southern California hill. Journal of Research on the Lepidoptera 6:69-178.
Shimada, K. 1979. Disappearance of diapause in hybrid pupae between two swallowtail species,
Papilio xuthus and P. machaon. Low Temperature Science(B) 37:67-71.
Sibley, C.G. 1954. Hybridization in the red-eyed towhees of Mexico. Evolution 8:252-290.
Sims, S.R. 1980. Diapause dynamics and host plant suitability of Papilio zelicaon
(Lepidoptera: Papilionidae) American Midland Naturalist 103:375-384.
Sims, S.R. 1983. Prolonged diapause and pupal survival of Papilio zelicaon Lucas
(Lepidoptera: Papilionidae) Journal of the Lepidopterists’ Society 37:29-37.
Smith, A.G. 1978. Environmental factors influencing pupal color determination in
Lepidoptera: I. Experiments with Papilio polytes, Papilio demoleus and Papilio polyxenes.
Proceedings of the Royal Society, B 200:295-329.
Sperling, F.A.H. 1986. Evolution of the Papilio machaon species group in western Canada.
MSc thesis, University of Alberta, Edmonton, Alberta, xvii + 284 pp.
Stoll, C. 1782. In , J.G. Cramer. De Uitlandsche Kapellen. ... Volume 4. Amsterdam, Baalde,
etc. 252 + 29 pp. plates 289-400.
Swofford, D.L. and R.B. Selander. 1981. BIOSYS-1: a FORTRAN program for the
comprehensive analysis of electrophoretic data in population genetics and systematics. The
Journal of Heredity 72:28 1-283.
Thorpe, J.P. 1982. The molecular clock hypothesis: Biochemical evolution, genetic
differentiation and systematics. Annual Review of Ecology and Systematics 13:139-168.
Thorpe, J.P. 1984. Primary and secondary transition zones in speciation and population
Quaest. Ent., 1987,23 (2)
300
Sperling
differentiation: a phylogenetic analysis of range expansion. Evolution 38:233-243.
Thorpe, J.P. 1985. Character number and the multivariate analysis of simple patterns of
geographic variation: categorical or “stepped clinal” variation. Systematic Zoology
34:127-139.
Tyler, H.A. 1975. The Swallowtail Butterflies of North America. Naturgraph Publishers,
Healdsburg, California. 192 pp. + 16 plates.
Valentine, K.W.S., K.R. Fladmark and B.E. Spurling. 1980. The description, chronology and
correlation of buried soils and cultural layers in a terrace section, Peace River Valley,
British Columbia. Canadian Journal of Soil Science 60:185-198.
Verity, R. 1905-191 1. Rhopalocera Palearctica. Published by the author, Florence Italy, lxxxvi
+ 368 pp. + [ix]— xii pp. + 2 maps + 80 plates.
Wake, D.B. 1981. The application of allozyme studies to problems in the evolution of
morphology, pp. 257-270. In , Evolution Today. (G.G.E. Scudder and J.L. Reveal, Editors).
Proceedings of the Second International Congress of Systematics and Evolutionary Biology.
Hunt Institute for Botanical Documentation, Pittsburgh, iv + 486 pp.
White, J.M. and R.W. Mathewes. 1982. Holocene vegetation and climatic change in the Peace
River district, Canada. Canadian Journal of Earth Science 19:555-570.
Wheeler, N.C. and R.P. Guries. 1982. Biogeography of lodgepole pine. Canadian Journal of
Botany 60:1805-1814.
Wiklund, C. 1973. Host plant suitability and the mechanism of host selection in larvae of
Papilio machaon. Entomologica Experimentalis et Applicata 16:232-242.
Wiklund, C. 1974. The concept of oligophagy and the natural habits and host plants of Papilio
machaon in Fennoscandia. Entomologica Scandinavica 5:151-160.
Wiklund, C. 1981. Generalist vs. specialist oviposition behavior in Papilio machaon
(Lepidoptera) and functional aspects on the hierarchy of ovipostion preferences. Oikos
36:163-170.
Wilson, K.H. 1961. Family Papilionidae, pp. 30-50. In, P.R. Ehrlich and A.H. Ehrlich. How to
Know the Butterflies. W.C. Brown, Dubuque, Iowa. 262 pp.
Wiltshire, E.P. 1958. The natural history of Papilio machaon L. in Bagdad. Transactions of the
Royal Entomological Society of London 1 10:221-244.
Wolley Dod, F.H. 1901. A preliminary list of the macro-lepidoptera of Alberta, N.W.T. The
Canadian Entomologist 33:157-172.
Wolley Dod, F.H. 1908. Further notes on Alberta Lepidoptera. The Canadian Entomologist
40:181-193.
Wyatt, C. 1957. Collecting on the Mackenzie and in the eastern arctic. The Lepidopterists’
News 11:47-53
Appendices: Tables 12-14
Quaest. Ent., 1987, 23 (2)
Variance : 54.6 69.8 78.2 32.2 44.4 52.4 89.9 100.
302
Sperling
Table 13. PCA loadings for e 1 ectrophoret i c characters.
Only wild collected larvae and confirmed ovipositions under natural conditions are included. Entries are arranged
by taxon and region. All entries from a particular locality are grouped together, even though entries from major
hybrid zones produced a variety of adults. Uncredited entries refer to personal obsevations or collections.
Abbreviations: AB = Alberta, BC = British Columbia, MB = Manitoba, SK = Saskatchewan, NWT = Northwest Territories.
Papilio machaon species group
303
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Table 14 (continued)
304
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Table 14 (continued)
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Angelica genuflexa
Taxon and Locality Foodplant Date Instars Adults obtained
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Table 14 (continued)
Papilio machaon species group
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Taxon and Locality Foodplant Date Instars Adults obtained
310
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Papilio machaon species group
311
INDEX TO VOLUME 23(2)
absinthium L., Artemisia , 269
Ae, S.A., 205, 251, 267, 275, 277, 290
alexanor Papilio, Papilio , 224, 257-258,
267 , 275, 277
Anethum graveolens, 273
Angelica , 241, 271
Angelica ampla , 27 1
Angelica arguta, 273
Artemisia absinthium L., 269
Artemisia arctica , 241, 271, 282
Artemisia dracunculus Linneaus, 204,
219, 241, 248-249, 257, 271, 273, 282
asterius Stoll, Papilio, 203
Ayala, F.J.,
see Dobzhansky, T., 216
bairdii W.H. Edwards, Papilio , 203-204,
250
banksiama Lamb., Pinus, 287
Barton, N.H., 206, 286, 288-289
Berenbaum, M., 269, 273, 282
Betula , 288
Bird, C.D., 258
Blackwelder, R.E., 216
Blair, A. P.,289
Blau, W.S., , 267, 286
see a/so Feeny, P., 273, 285
Boisduval, J., 203
brevicauda Saunders, Papilio , 273, 277 ,
285, 288
Brewer, G.J., 214
Brown, F.M., , 218-219
see a/so Miller, L.D., 218, 249, 286
brucei Edwards, Papilio , 204, 249
Bryant, O.,
see Leussler, R.A., 271
Buth, D.G., 216
Carson, H.L., 286, 291
Chermock, F.H., 204
Chermock, R.L.,
see Chermock, F.H., 204
Citrus , 271
Clark, A.H., 204
Clarke, C.A., 202, 204-205, 213, 251,
257, 268, 290
Colias gigantea gigantea Strecker, 287
Compositae, 201, 204, 269
contorta Loudon, Pinus , 284, 287
Corbin, K.W., 289
dardanus Brown, Papilio, 205
Dempster, J.P., 258
Dethier, V.G., 282
Dierl, W, 258
disa (Thunberg), Erebia, 287
Dobzhansky, T., 216
domesticus (Linneaus), Passer, 290
Dornfeld, E.J., 249, 267
dracunculus Linneaus, Artemisia, 204,
219, 241, 248-249, 257, 271, 273, 282
Dugle, J.R., 288
Edwards, W.H., 203-206, 249, 271
Ehrlich, P.R., 217, 269
Eller, K, 201,204
Emmel, J.F., , 205, 249-250, 269, 271
see also Emmel, T.C., 267, 286
see also Ferris, C.D., 251, 273
Emmel, T.C., , 267, 286
see also Emmel, J.F., 271
Endler, J.A., 288
Erebia disa (Thunberg), 287
eryphon (Boisduval), Incisalia, 288
Fabricius, J.C., 203
Feeny, P„ , 273, 285
see also Scriber, J.M., 205
Ferguson, A.,
see Corbin, K.W., 289
Ferguson, D.C., 285
Ferris, C.D., 251,273
Fisher, M.S., 205, 213, 249-251, 263, 267
Fladmark, K.R.,
see Valentine, K.W.S., 283
Foeniculum vulgare Mill., 271
Fox, D.J., 215
Freeman, T.N., 262
Gartside, D.F.,
see McDonnell, L.J., 288
gigantea gigantea Strecker, Colias, 287
Gillespie, G.D., 289
glaucus L., Papilio, 268-269
Gooding, R.H.,
see also Rolseth, B.M., 214
312
Sperling
gothica Remington, Papilio , 205, 250,
257,271
Grant, V., 289
Guedees, M.,
see Richard, D., 269
Guries, R.P.,
see Wheeler, N.C., 215, 284
Habgood, T.,
see Schweger, C., 283
Hafernik, J.E., 289
Hagen, H.A., 203-204, 206, 217
Hall, M.L.,
see Dempster, J.P., 258
Hancock, D.L., 275
Hanson, H.P., 283
Harrison, R.G., 287-288
Heitzman, J.R., 263
Heracleum , 241, 248, 258, 271
Heracleum lanatum, 271
Heron, R.J., 263
Hewitt, G.M.,
see Barton, N.H., 206, 286, 288-289
Hickman, M.,
see Schweger, C., 283
Higgins, L.G., 224, 269
Hilchie, G.J.,
see Bird, C.D., 258
hispaniolensis Temminck, Passer , 290
Hooper, R.R., 263, 271, 273
hospiton Gene, Papilio , 202, 224,
257-258, 267, 277 , 282
Hull, D.L.,217
Incisalia eryphon (Boisduval), 288
Incisalia niphon (Huebner), 288
indra Reakirt, Papilio , 202, 204, 224, 257,
267, 271, 273, 277-278, 282, 286
International Code of Botanical
Nomenclature, 218
International Code of Zoological
Nomenclature, 218
Jackson, B.S., 273
joanae Heitzman, Papilio , 263, 273, 277,
285
Jones, J.H., 289
Jordan, K,
see Rothschild, W., 204, 219
Junonia , 289
jutta chermocki Wyatt, Oeneis , 287
kahli Chermock and Chermock, Papilio ,
252
Kareiva, P.M.,
see Feeny, , 273, 285
Kettlewell, H.B.D., 267
Key, K.H.L., 217
Kimmich, H.P., 269
Knudsen, J.P.,
see Clarke, C.A., 204, 257
Kondla, N.G., ,219
see also Bird, C.D., 258
Krizek, G.O.,
see Opler, P.A., 263, 273
Kuhn, T.S., 216
Kurentzov, A. I., 269
Kuyt, E.,
see Bird, C.D., 258
Larsen, T.B., , 258
see also Clarke, C.A., 202
Leussler, R.A., 271
Linnaeus, C., 203
Littlejohn, M.J.,
see McDonnell, L.J., 288
Llewllyn Jones, J.R.J., 219
Lomatium, 241, 271
Lucas, P.H., 203
machaon aliaska Scudder, Papilio ,
203-204, 219-220, 222, 234, 243,
257-258, 262, 269, 271, 282-283, 288
machaon avinoffi Chermock and
Chermock, Papilio , 204, 252
machaon bairdii Edwards, Papilio , 249,
271,273,282-283,286
machaon centralis Staudinger, Papilio ,
269
machaon dodi McDunnough, Papilio ,
204, 219-220, 222-223, 225, 234,
240-241, 249, 254, 257-258, 262-263,
269, 271, 273, 283-284
machaon hudsonianus Clark, Papilio ,
204, 219-220, 222, 240, 252, 254-255,
257-258, 263, 268-269, 271, 273,
Papilio machaon species group
313
282-285, 287-288, 290
machaon Linneaus, Papilio , 202-205,
219, 221-222, 224-225, 234, 240-241,
243, 248-252, 254-255, 257-258, 262,
267-269, 271, 277-278, 282-285,
288-290
machaon oregonius (Edwards), Papilio ,
219- 220, 222, 225, 234, 243, 249,
257-258, 262, 269, 271, 282-284
machaon petersi Clark, Papilio , 204
machaon pikei Sperling, Papilio , 206,
220- 222, 234, 243, 251, 255, 257-258,
262, 268-269, 271,283
Maeki, K., 202
Masuda, K.K.,
see Shapiro, A.M., 271, 285
Mathewes, R.W.,
see White, J.M., 283
Matthews, J.V., 278, 282, 285
Mayr, E., 204, 217, 286, 289-291
McDonnell, L.J., 288
McDunnough, J., 204, 262
Menken, S.B.J., 214
Mickevich, M.F., 216
Miller, J.S., 218, 269
Miller, L.D., 249, 286, 289
Mitter, C.,
see Mickevich, M.F., 216
Mittwoch, U.,
see Clarke, C.A., 202, 204, 257, 290
Monroe, E.G., 202, 275
Moore, W.S., 289
Muting, D., 269
Nakamura, L, 267
Neff, N. A., 215,217
Nei, M., 252, 277
niphon (Huebner), Incisalia, 288
nitra Edwards, Papilio , 203, 205, 250-251
Nymphalidae, 289
Oeneis jutta chermocki Wyatt, 287
Oeneis uhleri Reakirt, 269
Oliver, C.G., 268, 286, 290
Opler, P.A., 263, 273
oregonius W.H. Edwards, Papilio,
203-204
P. machaon X polyxenes, 219, 223
P. polyxenes X machaon , 206, 219, 224
P. zelicaonX machaon , 219, 223-224
Palm, C.A.,
see Shapiro, A.M., 278
Papilio alexanor Papilio, 224, 257-258,
267, 275, 277
Papilio asterius Stoll, 203
Papilio bairdii W.H. Edwards, 203-204,
250
Papilio brevicauda Saunders, 273, 277 ,
285,288
Papilio brucei Edwards, 204, 249
Papilio dardanus Brown, 205
Papilio glaucus L., 268-269
Papilio gothica Remington, 205, 250, 257,
271
Papilio hospiton Gene, 202, 224, 257-258,
267, 277, 282
Papilio indra Reakirt, 202, 204, 224, 257,
267, 271, 273, 277-278, 282, 286
Papilio joanae Heitzman, 263, 273, 211,
285
Papilio kahli Chermock and Chermock,
252
Papilio machaon aliaska Scudder,
203-204, 219-220, 222, 234, 243,
257-258, 262, 269, 271, 282-283, 288
Papilio machaon avinoffi Chermock and
Chermock, 204, 252
Papilio machaon bairdii Edwards, 249,
271,273,282-283,286
Papilio machaon centralis Staudinger,
269
Papilio machaon dodi McDunnough, 204,
219-220, 222-223, 225, 234, 240-241,
249, 254, 257-258, 262-263, 269, 271,
273, 283-284
Papilio machaon hudsonianus Clark, 204,
219-220, 222, 240, 252, 254-255,
257-258, 263, 268-269, 271, 273,
282-285, 287-288, 290
Papilio machaon Linneaus, 202-205, 219,
221-222, 224-225, 234, 240-241, 243,
248-252, 254-255, 257-258, 262,
267-269, 271, 277-278, 282-285,
288-290
Quaest. Ent., 1987, 23 (2)
314
Sperling
Papilio machaon oregonius (Edwards),
219- 220, 222, 225, 234, 243, 249,
257-258, 262, 269, 271, 282-284
Papilio machaon petersi Clark, 204
Papilio machaon pikei Sperling, 206,
220- 222, 234, 243, 251, 255, 257-258,
262, 268-269, 271,283
Papilio nitra Edwards, 203, 205, 250-251
Papilio oregonius W.H. Edwards,
203-204
Papilio polyxenes asterius Stoll, 223, 241,
251-252, 263, 268, 271, 273, 277-278,
282, 284-285, 288
Papilio polyxenes coloro Wright, 273
Papilio polyxenes Fabricius, 202-203,
205-206, 224-225, 241, 248-249,
251-252, 254-255, 257-258, 263,
267-268, 271, 273, 277-278, 282,
284-288, 290
Papilio zelicaon Lucas, 202-206,
221- 222, 224-225, 234, 240-241, 243,
248-251, 254-255, 257-258, 262-263,
267-268, 271, 273, 277-278, 282-286,
288-290
Papilio zelicaon nitra Edwards, 205, 250
Papilio zolicaon Boisduval, 203
Papilionidae, 202
Papilioniodea, 203
Passer domesticus (Linneaus), 290
Passer hispaniolensis Temminck, 290
Perkins, E.M. Jr.,
see Perkins, S.F., 219
Perkins,, S.F., 219
Petasites palmatus , 269
Pike, E.M., 219, 284
Pimentel, R.A., 215
Pinus banksiama Lamb., 287
Pinus contorta Loudon, 284, 287
Platnick, N.,
see Nelson, G., 217
polyxenes asterius Stoll, Papilio , 223,
241,251-252, 263,268,271,273,
277-278, 282, 284-285, 288
polyxenes coloro Wright, Papilio , 273
polyxenes Fabricius, Papilio , 202-203,
205-206, 224-225, 241, 248-249,
251-252, 254-255, 257-258, 263,
267-268, 271, 273, 277-278, 282,
284-288, 290
Prasad, R.,
see Shaw, C.R., 214
Raven, P.,
see also Ehrlich, P.R., 269
Raven, P.H., 290
Reist, J.D., 288
Remington, C.L., , 201, 204-205, 213,
250-252, 257,263,271,288
see also Maeki, K., 202
Richard, D., 269
Riley, N.D., 269
Robinson, A.G.,
see Heron, R.J., 263
Rolseth, B., 214
Rothschild, W., 204, 219
Rumbacher, K., 282
Rutaceae, 271, 273, 286
Ryan, J.K.,
see Bird, C.D., 258
Schweger, C., 283
Scott, J.A., 263
Scriber, J.M., 205, 268
Scudder, G.G.E., 203
Selander, R.B.,
see Swofford, D.L., 216
Seyer, H., , 203, 277
see also Rumbacher, K., 282
Shapiro, A.M., 205, 250, 271, 278, 285
Shaw, C.R., 214
Sheppard, P.M.,
see Clarke, C.A., 204-205, 213, 251,
257, 268, 290
Shields, O, 287
Shields, O.,
see also Emmel, J.F., 205, 250, 269,
271
Shimada, K., 268
Shininger, F.S.,
see Perkins, S.F., 219
Sibley, C.G., , 289
see also Corbin, K.W., 289
Sims,S.R., 205, 271
Smith, A.G., 205
Papilio machaon species group
315
Smith, G.R.,
see Neff, N.A., 215
Smith, W.W.,
see Bird, C.D., 258
Stebbins, G.L.,
see Dobzhansky, T., 216
Swofford, D.L., 216
Thamnosa , 273
Thormin, T.W.,
see Bird, C.D., 258
Thorpe, J.P., 215, 254, 277, 288
Traut, W.,
see Clarke, C.A., 202, 204, 257, 290
Tyler, H.A., 273
uhleri Reakirt, Oeneis , 269
Umbelliferae, 201, 248, 271
Valentine, J.W.,
see Dobzhansky, T., 216
Valentine, K.W.S., 283
vulgare Mill., Foeniculum, 271
Wake, D.B., 255
Weislo, K.L.,
see Shapiro, A.M., 278
Wheeler, N.C., 284
White, J.M., 283
Wiklund, C., 205, 258, 267, 269
Willig, A.,
see Clarke, C.A., 268
Wilson, K.H., 204
Wiltshire, E.P., 267
Wolley Dod, F.H., 284
zelicaon Lucas, Papilio , 202-206,
221-222, 224-225, 234, 240-241, 243,
248-251, 254-255, 257-258, 262-263,
267-268, 271, 273, 277-278, 282-286,
288-290
zelicaon nitra Edwards, Papilio , 205, 250
Zizia, 258, 273
Zizia apt era, 271, 273
zolicaon Boisduval, Papilio, 203
Quaest. Ent., 1987,23 (2)
Publication of Quaestiones Entomologicae was started in 1965 as part of a
memorial project for Professor E. H. Strickland, the founder of the
Department of Entomology at The University of Alberta in Edmonton in 1922.
It is intended to provide prompt relatively low-cost publication for
comprehensive accounts of entomological research of greater than average
length. However, shorter papers about insects in the Prairie Provinces of
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Copy for all types of papers should conform to the Style Manual for
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Sciences, Second Editon, 1964, except that titles of periodicals should be given
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Return Undeliverable mail to the address above. Return Postage Guaranteed,
Issued August 1987
Quaestiones
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 23
NUMBER/
SommfR
6PB1N6 1987
Publication of Quaestiones Entomologicae was started in 1965 as part of a
memorial project for Professor E. H. Strickland, the founder of the
Department of Entomology at The University of Alberta in Edmonton in 1922.
It is intended to provide prompt relatively low-cost publication for
comprehensive accounts of entomological research of greater than average
length. However, shorter papers about insects in the Prairie Provinces of
Canada are acceptable. Page charges are normally levied, the rate determined
by printer’s charges. For information about current page charges, consult the
Editor.
Copy for all types of papers should conform to the Style Manual for
Biological Journals, published by the American Institute of Biological
Sciences, Second Editon, 1964, except that titles of periodicals should be given
in full. For style of taxonomic papers, the Editor should be consulted. Two
copies of a manuscript are requested. All manuscripts will be reviewed by
referees.
Abstracts are required, one in English and one in another language,
preferably French.
Copy for illustrations must accompany the manuscript, and be of such
character as to give satisfactory reproduction at page size (less 1 / 2 inch, or 1.2
cm on plates of full page size [73/4X5 inches, or 19.7 X 13.2 cm]). Reprints
must be ordered when proofs are returned, and will be supplied at cost.
Subscription rates are the same for institutions, libraries and individuals,
$17.00* per volume of four issues, normally appearing at quarterly intervals;
single issues $5.00**. Back volumes and issues are available at the same cost.
These prices supersede those previously indicated, and are subject to change
as required by inflationary pressure on the value of money.
Communications regarding subscriptions and exchanges should be
addressed to the Subscription Manager, and regarding manuscripts to the
Editor.
* Same rate in US$ for non-Canadian subscriptions.
** Single issues of more than 100 pages: $10.00; exception: volume 21(4) - $17.00.
Published quarterly by:
Department of Entomology
University of Alberta
Edmonton, Alberta, CANADA
T6G 2E3
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Return Undeliverable mail to the address above. Return Postage Guaranteed,
Issued December 1987
QUAES TIONES ENTOMOLOGICAE ISSN 0033-5037
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 23 Number 3 1987
CONTENTS
Clark-Revision of the Anthonomus Subgenus Anthomorphus Weise (Coleoptera:
Curculionidae) 317
Book Review-Mitchell, Andrew W. 1986. The Enchanted Canopy: A Journey to the Last
Unexplored Frontier, The Roofs of the World’s Rainforests 365
Book Notice-Belles, X. 1985. Systematica, filogenia y biogeografia de la subfamilia Gibbiinae
(Coleoptera: Ptinidae) 368
Book Review-D’Abrera, B. 1987. Sphingidae Mundi 369
Book Notice-Cox, J.M. 1987. Pseudococcidae (Insecta: Elemiptera) 371
Craig-A Taxonomic Account of the Black Flies (Diptera: Simuliidae) of the Society Islands -
Tahiti, Moorea and Raiatea 372
REVISION OF THE ANTHONOMUS SUBGENUS ANTHOMORPHUS WEISE
(COLEOPTERA: CURCULIONIDAE)
Wayne E. Clark
Department of Entomology
Alabama Agricultural Experiment Station
Auburn University, Alabama 36849, U.S.A.
Quaestiones Entomologicae
23:317-364 1987
ABSTRACT
The 22 species of the Anthonomus subgenus Anthomorphus Weise are hypothesized to
constitute a monophyletic group on the basis of morphological characters of the adult weevils.
Other adult morphological characters are cited as evidence in proposing a hypothesis of
phylogenetic relationships among the species. The species of Anthomorphus occur in the
Nearctic, Palearctic and Neotropical faunal realms and are known to have hosts in the plant
families Pinaceae, Rosaceae, Malpighiaceae, and possibly Betulaceae. A diagnosis and
description of the subgenus and of each of the species in the subgenus, a key, and illustrations
are presented as aids in identifying the species. Thirteen new species are described: A.
rosadonetoi (Brazil); A. howdenorum (Venezuela); A. rulfoi (Mexico); A., bechyneorum (Brazil,
Honduras, Venezuela); A. stockwelli (Panama); A. diamantinaensis (Brazil); A. camoiranensis
(Venezuela); A. vanini (Brazil); A. bordoni (Mexico, Venezuela); A. haliki (Brazil); A.
chinculticensis (Mexico); A. pimentai Argentina; Bolivia, Brazil) and A. galphimiae (Mexico).
Lectotypes are designated for A. paraguayanus Mustache and A. sulcipygus Champion.
RESUMEN
Caracteres morfoldgicos de los adultos indican que las 22 especies de Anthonomus del subgenero Anthomorphus
constituyen un grupo monofiletico. Se citan otros rasgos morfoldgicos de los adultos como evidencia de una hipdtesis de
las relaciones filogeneticas entre las especies. Las especies de Anthomorphus se encuentran en las Regiones Paleartica,
Nearctica y Neotropica y tienen plantas hospederas en las familias Pinaceae, Rosaceae, Malpighiaceae, y posiblemente
Betulaceae. Se presentan caracteres diagndsticas y una descripcibn del subgenero y de cada especie del subgenero,
ademas de una clave, y ilustracibnes para auxiliar en la identijicacion de las especies. Se describen trece especies nuevas
y se designan lectotipos para dos especies previamente descritas (vease el resumen en ingles).
INTRODUCTION
The Anthonomus subgenus Anthomorphus Weise contains 22 species that occur in the
Nearctic, Palearctic and Neotropical faunal realms. The species have been collected in
association with plants in several families, including Betulaceae, Fagaceae, Malpighiaceae,
Myricaceae, Rosaceae and Pinaceae. Of these, only Malpighiaceae, Rosaceae and Pinaceae
have been established by rearing as actual hosts. Some of the species have been collected on
Malpighiaceae in association with species of Anthonomus in the venustus and unipustulatus
groups (Clark, 1987; Clark and Burke, 1985) and members of the anthonomine genus
Pseudanthonomus. The present study was undertaken to provide descriptions or redescriptions,
illustrations, and a key to facilitate identification of the species. The information on these
species is currently being analyzed to determine their relationships to the Anthonomus
318
Clark
venustus and unipustulatus groups, the genus Pseudanthonomus Dietz and other
Anthonominae known or suspected to have hosts in the Malpighiaceae.
MATERIALS AND METHODS
Specimens of 653 adults, including types of most of the previously described species, were
examined. These were from collections of the following institutions (abbreviations in
parentheses are used to refer to collections in the text): Auburn University Entomological
Collections (AUEM), Auburn, W.E. Clark; Museu Paraense Emilio Goeldi (MPEG), Belem,
W.L. Overal; Museum of Comparative Zoology (MCZC), Cambridge, S. R. Shaw; Illinois
Natural History Survey (INHS), Champaign, J. K. Bouseman; Field Museum of Natural
History (FMNH), Chicago, J.S. Ashe; Texas A&M University (TAMU), College Station,
H.R. Burke; Universidade Federal do Parana (DZUP), Curitiba, G. Rosado-Neto; Deutsches
Entomologisches Institut (IPZE), Eberswalde, L. Dieckmann; Cornell University (CUIC),
Ithaca, J.K. Liebherr; Snow Entomological Museum, University of Kansas (SEMC),
Lawrence, P.A. Ashlock; The British Museum (Natural History) (BMNH), London, R.T.
Thompson; Universidad Central de Venezuela (IZAV), Maracay, L.J. Joly; The American
Museum of Natural History (AMNH), New York, L.H. Herman, Jr.; Canadian National
Collection of Insects and Arachnids (CNCI), Ottawa, D.E. Bright; Museum National
d’Histoire Naturelle (MNHP), Paris, H. Perrin; Carnegie Museum of Natural History
(ICCM), Pittsburg, J.E. Rawlins; California Academy of Sciences (CASC), San Francisco,
D.H. Kavanaugh; Museu de Zoologia (MZSP), Universidade de Sao Paulo, U. Martins;
Centre National de Recherches Agronomiques (LFEV), Versailles, J. d’Aguilar; National
Museum of Natural History (USNM), Washington, D.C., D.R. Whitehead; Purdue University
(PURC), West Lafayette, A.V. Provonsha. The following provided specimens from their
personal collections: C. Bordon (CCBM), Maracay; H.A. Hespenheide (CHAH), Los Angeles;
H. and A. Howden (HAHC), Ottawa; M.A. Ivie (CMIV), Columbus; C.W. O’Brien (CWOB),
Tallahassee; E.L. Sleeper (ELSC), Long Beach.
Measurements were made with the aid of an ocular micrometer in a dissecting microscope
as follows: total length from anterior margin of eyes to elytral apices in lateral view; width
across elytra at widest point; length of pronotum from anterior to posterior margins; length of
rostrum from anteroventral margin of eye to tip, across arc, in lateral view; length of distal
portion of rostrum from antennal insertions to tip in lateral view; width of frons at narrowest
point between eyes; width of base of rostrum just distad of eyes in dorsal view; and width of pro-
and metafemora in anterior view excluding the ventral teeth. The range and, in parentheses, the
mean and sample size of each measurement is given for each species.
Relationships of the species of Anthomorphus to each other were analyzed using PAUP,
Phylogenetic Analysis Using Parsimony, Version 2.4, developed by Swofford (1985), on an
IBM Personal Computer (see Phylogeny section).
Subgenus Anthomorphus Weise
Anthomorphus Weise 1883: 255. Type species (by subsequent designation: Dieckmann 1968: 470): Curculio varians
Paykull.
Paranthonomus Dietz 1891: 199. Type species (by original designation): Anthonomus profundus Le Conte.
Recognition. — The species of Anthonomus in the subgenus Anthomorphus are
distinguished by the following combination of characters:
Anthomorphus Weise
319
1) abdomen of male (Fig. 37) with sternum 5 subquadrately emarginate, leaving
triangular, setose sclerite isolated from posterior margin (except in A. pinivorax );
2) pygidium of male (Figs. 33-36) and of female (Figs. 38-46) sulcate or impressed
(except in A. diamantinaensis and female A. rosadonetoi).
Description. — Male. Length : 1. 8-4.1 mm. Width : 1. 0-2.0 mm. Head : vertex with median fovea, sparsely,
minutely punctate, with narrow, setiform scales; venter with slightly to much broader, more pallid scales, eyes round,
posterior margin slightly elevated so that greatest convexity occurs there. Rostrum : length 1.0-1. 8 x pronotal length; in
dorsal view, sides converging slightly from base to antennal insertions, subparallel or gradually diverging from there to tip;
proximal portion rugose-punctate to distinctly carinate; dorsal margin of lateral rostral groove carinate, directed to middle
of eye; distal portion shallowly punctate to rugose-punctate, length 16-39% of total rostral length; antennal funiculus with
7 articles. Thorax-, sides evenly rounded from subapical constriction; evenly convex from constriction to base; pronotum
densely, coarsely punctate, each puncture giving rise to a narrow scale; broader scales whitish, limited to dorsal midline
and to lateral portions, narrower scales aeneous, forming broad lateromedian vittae; pleuron with punctation and vestiture
as on pronotum, scales on lower portions slightly broader and more pallid. Elytra : humeri not prominent, sides subparallel
in basal 1 /2; striae deep, punctures slightly impinging on interspaces; interspaces more-or-less uniformly convex, sutural
interspaces narrow at extreme base, these and interspaces 2-5 slightly widened and slightly elevated posterior to shallow
obliquely transverse depression; each interspace with median row of narrow to setiform scales and lateral rows of slightly
broader scales directed toward center of interspace. Abdomen : sternum 5 about as long as sternum 4; each sternum with
large, platelike, internal apodeme. Legs: apices of femora truncate, narrowly emarginate; profemur slightly stouter than
metafemur, with large ventral tooth and in some species a smaller, more distal tooth; metafemur with one small acute
ventral tooth; protibia straight to slightly, broadly concave medially on dorsal margin, ventral margin with more-or-less
abruptly delimited median prominence, with or without preapical tooth, apically uncinate; metatibia more nearly straight
than protibia, slightly widened or narrowed at apex.
Female. Length : 2.0-4. 2 mm. Width : 1. 0-2.1 mm. Rostrum : length 1.2-2. 3 x pronotal length; broadly, evenly curved;
length of distal portion 25-49% of total rostral length. Pygidium : with or without dorsomedian impression or sulcus.
Discussion. — The subgenus Anthomorphus was established by Weise (1883: 255) to
include Anthonomus varians (Paykull) and A. perforator (Herbst), along with A. ater
(Marsham), cited as a synonym of the latter. Weise stated that the subgenus was characterized
by the “ unguiculi dentati” (toothed tarsal claws) of the included species. Dieckmann (1968:
471) noted that this characterization was in error and that, in fact, A. varians is the species
with untoothed tarsal claws (the “ unguiculi mutici ’ of Weise). Dieckmann (1968: 474) listed
A. perforator as a “Farbaberration” (color variety) of A. varians and noted that A. ater is a
junior synonym of A. rubi (Herbst). He listed A. varians as type species of the subgenus
Anthomorphus, noting that that species, like the type species of the subgenus Paranthonomus
Dietz, has “ein gefurchtes Pygidium.” On the basis of this pygidial character, Dieckmann
(1968) also included the Palearctic A. pinivorax Silfverberg [as A. pubescens (Paykull)]. He
listed the name Anthomorphus as a synonym of Paranthonomus, but did not explain why he
used the name Paranthonomus for the taxon instead of the older name, Anthomorphus.
The name Paranthonomus was proposed by Dietz (1891) for a subgenus of Anthonomus
including the North American Anthonomus profundus LeConte, A. rubidus LeConte and A.
vulpinus Dietz, all of which were characterized as having a “deep, longitudinal excavation of
the pygidium.” Dietz (1891: 200) also stated that in Europe Paranthonomus is “represented by
A. pubescens, and probably A. pyrenaeus .” (The latter name is also a “Farbaberration” of A.
varians, according to Dieckmann [1968: 474]).
Champion (1903) considered his sulcipygus group, including the Panamanian A. sulcipygus
Champion, to be the same as Dietz’s Paranthonomus. Schenkling and Marshall (1934) listed
A. sulcipygus and 18 additional Neotropical species as members of the subgenus. With the
exception of A. abdominalis Champion, these species do not have the diagnostic characters of
Anthomorphus and they are not retained in the subgenus. The relationships of these other
species have not been satisfactorily determined, although some have been included in recently
completed revisionary studies. For example, A. melanostictus and A. venustus were assigned to
Quaest. Ent., 1987,23 (3)
320
Clark
the venustus group by Clark and Burke (1985), and A. cossonoides was assigned (as a synonym
of A. instabilis Faust) to the gularis group by Clark and Burke (1986a).
Burke (1962: 205-206) placed A. unipustulatus in Paranthonomus, citing the sulcate
pygidium but noting that in that species the sulcus is “shorter and less deeply excavated” than
in A. profundus and A. rubidus. The association of the Peruvian A. sulcatus Kirsch and A.
filicornis Hustache from Guadeloupe with Anthomorphus was not made before the types were
examined in connection with the present study. O’Brien and Wibmer (1982:105) and Wibmer
and O’Brien (1986: 203) listed Paranthonomus as a subgenus of Anthonomus but did not
indicate which species belong to the group.
The following species are included in the subgenus Anthomorphus.
Anthonomus rubidus species group
Anthonomus rubidus LeConte
Anthonomus bordoni new species
Anthonomus paraguayanus Hustache
Anthonomus profundus LeConte
Anthonomus vulpinus Dietz
Anthonomus galphimiae, new species
Anthonomus howdenorum, new species
Anthonomus varians (Paykull)
Anthonomus pinivorax Silfverberg
Anthonomus sulcatus species group
Anthonomus sulcipygus Champion
Anthonomus chinculticensis, new species
Anthonomus vanini, new species
Anthonomus camoiranensis, new species
Anthonomus stockwelli, new species
Anthonomus filicornis Hustache
Anthonomus sulcatus Kirsch
Anthonomus pimentai, new species
Anthonomus bechyneorum , new species
Anthonomus haliki , new species
Anthonomus rulfoi, new species
Anthonomus abdominalis Champion
Anthonomus diamantinaensis, new species
Anthonomus rosadonetoi, new species
Males of the species of Anthomorphus are distinguished by the characters in the following
key. With some exceptions, noted in the treatments of the appropriate species, females are
morphologically uniform throughout the subgenus. This presented problems in determining the
identity of some species previously described from females only. Furthermore, several females
remain among the specimens examined that cannot confidently be associated with identified
males or females. Even though non-genitalic characters are preferentially used in the key, the
male genitalia provide the best diagnostic characters.
Anthomorphus Weise
321
Key to Species of Anthomorphus (males only)
1 Abdominal sternum 5 subquadrately emarginate, with triangular, setose
sclerite isolated from posterior margin (Fig. 37)
1' Abdominal sternum 5 not subquadrately emarginate, with triangular,
setose sclerite isolated from posterior margin
A. pinivorax Silfverberg, p. 330
2 (1) Tasral claws simple, without basal tooth A. varians (Paykull), p. 329
2' Tarsal claws toothed
3 (2') Pygidium sulcate (Figs. 33-36)
3' Pygidium not sulcate A. diamantinaensis, new species, p. 340
4 (3) Metasternum with long, fine, erect, setiform scales
4' Metasternum with scales (somewhat setiform and slightly raised in some,
but not erect)
5 (4) Meso- and metafemora with sparse ventral fringe of long, fine, setiform
scales
5' Meso- and metafemora without ventral fringe of setiform scales
6 (5) Metatibia with ventral fringe of long, setiform scales
A. rulfoi, new species, p. 339
6' Metatibia without ventral fringe of setiform scales
7 (60 Profemur with large tooth distal to ventral tooth; pronotum and elytra with
fine aenescent scales; aedeagus as in Fig. 48
A. bordoni, new species, p. 325
7' Profemur with small or obsolescent distal tooth; pronotum and elytra with
fine whitish scales; median lobe of male genitalia as in Fig. 47
A. rubidus LeConte, p. 323
8 (50 Metatibial uncus with apical prominence; pygidial sulcus broad, not
delimited apically A. galphimiae, new species, p. 328
8' Metatibial uncus without apical prominence; pygidial sulcus narrower,
delimited apically A. howdenorum, new species, p. 328
9 (40 Profemur with distal tooth distinctly separated from large ventral tooth . .
9' Profemur without distal tooth, or with slight indication of distal tooth
arising from basal portion of ventral tooth
10 (9) Metatibia with ventral fringe of long, aenescent, setiform scales in distal
1/2 A. camoiranensis , new species, p. 332
10' Metatibia without ventral fringe of setiform scales
11 (10') Elytra with dark discal macula (indistinct and limited to sutural
interspaces in A. haliki ); strial punctures large, elytral vestiture fasciculate
IT Elytra without dark discal macula; strial punctures smaller, elytral
vestiture not fasciculate
12 (11) Vestiture of prothorax broad scales and strongly differentiated setiform
scales; meso- and metafemora without ventral fringe of long, fine setiform
scales A. bechyneorum, new species, p. 337
12' Vestiture of prothorax elongate, narrow, acuminate scales; meso- and
metafemora with ventral fringe of long, fine, setiform scales
A. haliki , new species, p. 338
2
3
4
5
9
6
8
7
10
17
11
12
13
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322
Clark
13 (1 T) Pygidial channel narrow, shallow (Fig. 35) 14
13' Pygidial channel broader, deeper 15
14 (13) Elytra with common middorsal fascia; protibia with long, sparse, ventral
setiform scales A. ahdominalis Champion, p. 339
14' Elytra without middorsal fascia; protibia without ventral setiform scales .
A. sulcatus Kirsch, p. 334
15 (130 Elytral interspace 3 prominent at extreme base
A. rosadonetoi, new species, p. 341
15' Elytral interspace 3 not prominent at base 16
16 (150 Protibia with preapical tooth; metatibial uncus without apical prominence
A. vanini, new species, p. 332
16' Protibia without preapical tooth; metatibial uncus with apical prominence
A.filicornis Hustache, p. 333
17 (90 Pro- and mesocoxae with long setiform scales; profemur with ventral tooth
displaced distally, without distal tooth; pygidial sulcus with median carina
(Fig. 36) A. pimentai, new species, p. 335
17' Pro- and mesocoxae without setiform scales; profemur with ventral tooth
not displaced distally, with or without distal tooth; pygidial sulcus without
median carina 18
18 (170 Pygidial sulcus narrow, not delimited apically (Fig. 34); head constricted
behind large, prominent eyes, frons delimited posteriorly by transverse
channel A. stockwelli new species, p. 333
18' Pygidial sulcus broad, delimited apically (Fig. 33); head not constricted
behind eyes, frons not delimited by transverse channel 19
19 (180 Elytral integument darkest on large triangular portion of dorsum extended
from humeri to about middle and apically along sutural interspaces 20
19' Elytral integument not darker on triangular portion of dorsum 21
20 (19') Metafemur of male straight, ventral tooth not displaced distally; metatibial
uncus with prominence; protibia without preapical tooth
A. paraguayanus Hustache, p. 325
20' Metafemur of male strongly curved, ventral tooth displaced distally;
metatibial uncus simple; protibia with preapical tooth
A. profundus LeConte, p. 326
21 (19') Aedeagus (Fig. 55) abruptly constricted apically to obliquely truncate
apical prominence A. sulcipygus Champion, p. 331
21' Aedeagus (Fig. 56) constricted distally to symmetrical, rounded apex ....
A. chinculticensis, new species, p. 331
ANTHONOMUS RUBIOUS SPECIES GROUP
Recognition. — The members of the rubidus species group are distinguished by the
following combination of characters:
1) spiculum gastrale of male genitalia with basal portion closely engaging sides of
aedeagus, with ventral keel (Fig. 69) (except in A. pinivorax );
2) aedeagus with midventral, subbasal lobe (Figs. 47-53) (except in A. pinivorax, Fig.
54).
Anthomorphus Weise
323
Discussion. — This group includes the two North American, two Palearctic, and five
Neotropical species. Three monophyletic subgroups are also recognized. One includes A.
rubidus, A. bordoni and A. paraguayanus, all of which have the apical portion of the aedeagus
somewhat asymmetrical (Figs. 47-49). Another includes A. profundus , A. howdenorum and A.
galphimiae. These have the sutural interspaces slightly to markedly expanded subapically. The
last subgroup includes the Palearctic A. varians and A. pinivorax, distinguished from the New
World species by several characters, including possession of sparse, whitish, setiform scales on
the elytra. As indicated above, one of these, A. pinivorax , lacks the diagnostic characters of the
rubidus group. It is assigned to the group on the basis of other characters it shares with A.
varians which indicate that the two are sister species.
Host relationships of the North American species in the rubidus group have not been
determined with certainty, but the most likely hosts are in the plant family Rosaceae and
possibly Betulaceae. Hosts of four of the five Neotropical species are unknown, but at least one
is associated with Malpighiaceae. The Palearctic species are unusual among anthonomines in
having hosts in the family Pinaceae.
Anthonomus rubidus LeConte
Anthonomus rubidus LeConte, 1876: 199. Hamilton 1885: 106. Dietz 1891: 200-201. Fall 1913: 47. Blatchley and Leng
1916: 291. Burke 1962: 206. Holotype: Pennsylvania, male [Penn.] [Type 1959] [J.L. Leconte/Coll.] [A.
rubidus/Lec.] (MCZC).
Recognition (Fig. 1). — This North American species is distinguished by the following
combination of characters:
1) mesosternum, abdominal sterna, and meso- and metafemora of male with long, sparse,
setiform scales;
2) elytra with midbasal macula extended posteriorly on suture (Fig. 2);
3) aedeagus slightly asymmetrical (Fig. 47);
4) endophallus with sparse, minute proximal denticles, with larger, irregular median
denticles (Fig. 47).
It closely resembles A. bordoni from Venezuela from which it is distinguished by the smaller,
less distinct distal profemoral tooth, the longer, more dense, erect, setiform scales on the
metasterum, abdomen, and metafemur and by slight differences in the male genitalia (Figs. 47,
48). It is most likely to be confused with the North American A. profundus from which it is
distinguished by the following characters:
1) vestiture of pronotum and elytra denser (Figs. 1, 2, 5, 6);
2) sutural elytral interspaces without subapical prominences (Fig. 5);
3) pygidial sulcus narrower in male and female (Figs. 33, 38, 39);
4) long setiform scales on male metasternum and meso- and metafemora;
5) aedeagus less strongly asymmetrical (Figs. 47, 50);
6) profemora less strongly inflated, with more well-developed distal tooth;
7) and, protibia without preapical tooth.
As noted by Hamilton (1885: 106), specimens of A. rubidus have been confused in
collections with those of Pseud anthonomus crataegi Walsh. However, as Hamilton noted,
“there should not be much trouble in distinguishing (the two species), as crataegi has only six
joints in the funicle of the antenna, while rubidus has seven - a matter readily determined by
counting them under a microscope.”
Quaest. Ent., 1987, 23 (3)
324
Clark
Description. — Male. Length: 2. 1-2.6 mm (x = 2.4, n=15). Width : 0.9-1. 3 mm (x=1.2, n=15). Head-, eyes
separated by distance ca. 0.6 x width of rostrum at base. Rostrum : slender, length 1.1-1. 5 x (x=1.3, n=15) pronotal
length; broadly, evenly curved; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion finely, sparsely
punctate, length 21-28% (x = 25, n= 15) of total rostral length. Thorax : pronotum with dark, narrow aenescent scales and
with middorsal vitta of long, narrow, pallid whitish scales. Pygidium: median sulcus wider and deeper toward apex. Legs:
profemur slender, ca. 1.3 x stouter than metafemur, with distal tooth that shares common base with larger ventral tooth in
some specimens; protibia without preapical tooth; metatibial uncus long, straight, oblique, truncate.
Female. Length : 2. 2-2. 6 mm (x = 2.4, n=15). Width : 1.1-1. 3 mm (x=1.2, n=15). Rostrum: length 1.2-1. 4 x
(x= 1.2, n= 15) pronotal length; broadly, evenly curved; proximal portion rugose, vestiture limited to proximal 1/3; distal
portion smooth, length 27-35% (x = 30, n=15) of total rostral length. Pygidium (Fig. 38): with broadly rounded
apicodorsal prominence; median sulcus long, straight to slightly wider posteriorly, with long, dense setae. Abdomen:
sternum 5 with posterior margin slightly produced medially.
Plant Associations. — Label data indicate that specimens of A. rubidus have been collected
on the following plants:
Betulaceae
Betula lutea Michx. (under surface of leaves, 1 specimen, McKeever, Herkimer Co.,
New York)
Corylus (20 specimens, Kappa, Illinois; 2 specimens, Haddon Heights, New Jersey)
Fagaceae
Quercus alba (1 specimen, Westerville, Ohio)
Juglandaceae
wild hickory (1 specimen, Tippecanoe Co., Indiana)
Rosaceae
“cherry” (4 specimens, Tippecanoe Co., Indiana)
Prunus demissa D. Dietr. (1 specimen. Bountiful, Davis Co., Utah)
Prunus serotina J.F. Ehrh. (1 specimen, Albemarle Co., Virginia, 2 specimens, Rhea
Co., Tennessee)
“wild cherry” (1 specimen, Tippecanoe Co., Indiana).
Distribution. — This species is widespread in North America from Canada to Florida,
westward to Minnesota and Illinois, with an extralimital record in Utah. In addition to the
holotype of A. rubidus from Pennsylvania, specimens from the following localities were
examined.
CANADA. Quebec. Aylmer (1 female, ICCM). Duparquet (1 female, USNM).
UNITED STATES. Connecticut. New Haven Co.: Wallingford (1 female, CWOB). District of Columbia.
Kennelworth Pond (1 female, TAMU). Florida. Duval Co. (1 female, USNM). Georgia. White Co.: Helen (2 males,
USNM). Illinois. (1 female, FMNH; 1 female, ICCM). Woodford Co.: Kappa (4 males, 3 females, TAMU, 8 males, 5
females, USNM). Indiana. Tippecanoe Co. (1 female, TAMU, 12 males, 3 females, USNM). Iowa. (1 male, USNM).
Kentucky. Fayette Co.: (1 male, USNM). Maryland. Garrett Co.: Deer Park (1 female, USNM). Prince Georges Co.:
Beltsville (1 female, USNM) (1 female, USNM); Bladensburg (1 female, USNM). Washington Co.: Boonsboro (1 male,
3 females, TAMU). Massachusetts. (1 male, SEMC). Adams Co.: Mt. Greylock (1 female, CUIC). Barnstable Co.: East
Sandwich (1 female, USNM). Natick Co.: Sherborn (1 male, MCZC). Michigan. Marquette Co.: Marquette (1 female,
USNM). Midland Co.: (1 female, USNM). Muskegon Co.: Henry (1 female, USNM). Minnesota. Rice Co.: Nerstrand
Woods (1 female, CWOB). New Hampshire. Coos Co.: Carter Dome, White Mountains (4 males, USNM); Carter Notch,
White Mountains (3 females, USNM). Grafton Co.: Franconia (1 male, AMNH). New Jersey. Camden Co.: Haddon
Heights (1 male, 1 female, CNCI). Monmouth Co.: Hornerstown (1 female, AMNH). Morris Co.: Budd Lake (1 female,
AMNH). New York. Herkimer Co.: McKeever (1 female, USNM). Ulster Co.: Slide Mt. (14 males, 1 female, USNM) (2
males, 1 female, USNM). North Carolina. Yancey Co.: Black Mountains (1 male, 1 female, AMNH, 1 female, CASC).
Ohio. (1 male, SEMC). Franklin Co.: Westerville (1 female, ELSC). Hamilton Co.: Cincinnati (1 male, USNM).
Hocking Co.: 8 mi. SW Logan, S.R. 664 (1 female, CMIV). Pennsylvania. Allegheny Co.: Pittsburg (2 males, 2 females,
ICCM). Forest Co.: 1 mi E Neiltown (1 female, USNM). Indiana Co.: Indiana (1 male, AMNH). Northampton Co.:
Wind Gap (1 male, CASC). Westmorland Co.: Jeannette, 15, 20, 28 (1 male, CNCI; 3 males, 1 female, ICCM). York
Co.: 5 mi. N Davidsburg (1 female, USNM). Tennessee. (1 female, USNM). Davidson Co.: Nashville (1 female, USNM).
Great Smoky Mountain National Park (2 males, 1 female, HAHC). Rhea Co.: (2 males, USNM) (1 male, 1 female,
USNM). Sevier Co.: Gatlinburg (1 male, 1 female, USNM). Utah. Davis Co.: Bountiful (1 male, USNM). Virginia.
Albemarle Co.: (2 males, 1 female, USNM), (1 male, USNM). Alexandria Co.: (1 male, USNM). Washington Co.:
Anthomorphus Weise
325
Blacksburg (1 male, USNM). West Virginia. Fort Pendleton (1 female, USNM). The specimens were collected in the
months of May-October.
Anthonomus bordoni, new species
Type Series. — Holotype: Venezuela, male [VENEZUELA: Merida/Jaji/25 June 1983/W.E. Clark and Clark]
[Host 83V-09] (USNM). Paratypes : Mexico, 1 male [MEXICO, N.L. 2200 m. El/Potosi Mt. 8 km. NW, 18/de Marzo
Galeana Aug. 26, 1977 A. Garcia A.] [on pine/& oak]. Venezuela, 4 males, 6 females [VENEZUELA: Merida/Jaji/26
June 1983/W.E. Clark and Clark - ] [Host 83V-09]; 1 male [Venezuela AR/Rancho Grande/ 1 100 m 1 7— V 1 1 1—1965] [F.
Fernando-Y./J. Salcedo/Cols.] [A La Luz]; 1 female [La Grita m./2300. TACHIRA] [VENEZ. Bordon/leg. 25 III
1985]. Total paratypes, 13 (AUEM, CCBM, CWOB, IZAV, TAMU).
Recognition (Fig. 3). — This species is distinguished by the following combination of
characters:
1) metasternum, abdominal sterna, and meso- and metafemora with sparse ventral fringe
of long setiform scales;
2) aedeagus asymmetrical (Fig. 48);
3) elytra with midbasal macula extended posteriorly on suture (Fig. 4);
4) endophallus with minute proximal denticles, with sparse median denticles and large
median tooth (Fig. 48).
It resembles the North American A. rubidus from which it is distinguished by the more distinct
distal tooth on the profemur and, in the male, by the longer, more dense setiform scales on the
metasternum, abdomen, and metafemur. The aedeagus also differs slightly in the two species
(Figs. 47,48).
Description. — Male. Length : 2. 4-2. 8 mm (x=2.6, n = 6). Width : 1. 1-1.4 mm (x=1.3, n = 6). Head : eyes
separated by distance ca. 0.7 x width of rostrum at base. Rostrum : slender, length 1.0-1. 5 x (x=1.4, n = 6) pronotal
length; broadly, evenly curved; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion finely, sparsely
punctate, length 19-30% (x = 26, n = 6) of total rostral length. Thorax : pronotum with elongate, narrow, dark
fulvo-aeneous scales and middorsal vitta of broader, more pallid scales. Pygidium: median sulcus broad anteriorly, deeper
and slightly wider posteriorly, not extended to apex. Legs: profemur narrow, ca. 1.3 x stouter than metafemur; distal tooth
small, acute; protibia with acute preapical tooth; metatibial uncus minute, slightly hooked.
Female. Length: 2.6-2. 9 mm (x = 2.7, n = 8). Width: 1.2-1. 4 mm (x=1.4, n = 8). Rostrum: length 1.4-1. 6 x (x=1.5,
n = 8) pronotal length; broadly, evenly curved; proximal portion rugose, vestiture limited to extreme base; distal portion
smooth, length 29-36% (x=34, n = 8) of total rostral length. Pygidium: broadly rounded apically; median sulcus long,
narrow, shallow, not wide posteriorly, without setae. Abdomen: sterum 5 with posterior margin nearly straight.
Plant Associations. — The type series of A. bordoni was taken on an unidentified tree.
Distribution. — This species is known only from the type series from Venezuela and Mexico.
Specific Epithet. — This species is named in Honor of Carlos Bordon of Maracay,
Venezuela, as a token of appreciation for his friendship and assistance.
Anthonomus paraguayanus Hustache
Anthonomus paraguayanus Hustache 1939: 55-56. Lectotype (here designated): Paraguay, male [Paraguay/Hohenau]
[(2)] [TYPE] [MUSEUM PARIS/ 1949/Col. A. Hustache] [Anthonomus/paraguayanus/m.] (MNHP).
Paralectotypes: Paraguay, 1 male [Paraguay/Hohenau] [Syntypus] [Anthonomus/paraguayanus/co-type m.] [Coll.
DEI/Eberswalde] (IPZE); 1 female [Paraguay/Hohenau] [(2)] [TYPE] [MUSEUM PARIS/ 1949/Col. A.
Hustache] (MNHP). Hustache (1939: 55-56) indicated that this species was described from four females, but the
IPZE “co-type” and one of two MNHP syntypes examined are males.
Recognition. — This species is distinguished by the following combination of characters:
1) metasternum, abdominal sterna, and meso- and metafemora of male without long
setiform scales;
2) aedeagus asymmetrical, constricted to narrowly rounded apex (Fig. 49);
3) metatibial uncus with prominence;
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Clark
4) elytra with midbasal macula extended posteriorly on suture;
5) endophallus with sparse proximal denticles and a large distal tooth (Fig. 49).
It closely resembles the North American A. rubidus and A. bordoni from Venezuela. It is
distinguished from these two species by the absence of long setiform scales on the male femora,
the distinct prominence on the male metatibial uncus, and by differences in the male genitalia
(Figs. 47-49).
Description. — Male. Length : 1.8-2. 5 mm (x=2.2, n = 6). Width : 1. 0-1.1 mm (x=1.0, n = 6). Head : eyes
separated by distance ca. 0.8 x width of rostrum at base. Rostrum : slender, length 1.0-1. 5 x (x = 1.3, n = 6) pronotal
length; broadly, evenly curved; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion finely, sparsely
punctate, length 27-35% (x = 32, n = 6) of total rostral length. Thorax: pronotum with long, narrow aenaescent scales and
middorsal vitta of broader, more pallid fulvous scales. Pygidium : median sulcus narrow posteriorly, wider and deeper
apically. Legs: profemur slender, ca. 1 .4 x stouter than metafemur, with minute distal tooth that shares common base with
larger ventral tooth; protibia without preapical tooth.
Female. Length: 2. 1-2.4 mm (x = 2.2, n = 2). Width: 1.0 mm (x=1.0, n = 2). Rostrum: short, length 1.5 x (x=1.5,
n = 2) pronotal length; slightly, evenly curved; proximal portion smooth, vestiture limited to extreme base; distal portion
smooth, length 34-35% (x = 35, n = 2) of total rostral length. Pygidium: broadly rounded, apicodorsal prominence obsolete;
median sulcus short, shallow, remote from apex, not wider apically. Abdomen: sternum 5 with posterior margin slightly
concave medially.
Plant Associations. — Unknown.
Distribution. — Anthonomus paraguayanus is known only from northern Argentina and
southern Brazil. In addition to the lectotype and paralectotypes from Paraguay, the following
specimens were examined.
ARGENTINA. Misiones. El Dorado (1 female, AMNH).
BRAZIL. Santa Catarina: Nova Teutonia (5 males, 1 female, ELSC, HAHC, MZSP). The specimens were collected
in August, September and November.
Anthonomus profundus LeConte
Anthonomus profundus LeConte 1876: 198. Schwarz 1890: 232. Hamilton 1895: 376. Pierce 1907: 268. Fall 1913: 46-47.
Leng 1920: 322. Schenkling and Marshall 1934: 38. Burke 1962: 206. Hatch 1971: 349-350. Lectotype (designated by
Burke (1984: 265): Illinois, female [(small golden disc)] [Type/1958] [J.L. LeConte/coll.]
[LECTOTYPE/Anthonomus/profundus/LeC. /design. by/H.R. Burke] [A. profundus/Lec.] (MCZC).
Anthonomus vulpinus Dietz 1891: 201. Lectotype (designated by Burke 1984: 262-263): Pennsylvania, male [Pa.]
[Type/1955] [W.G. Dietz/Coll.] [LECTOTYPE/Anthonomus/vulpinus/Dietz/design. by/H.R. Burke] (MCZC).
This synonymy was recognized by Fall (1913: 46-47) who stated that “... the differences given by Dietz... do not look
as important in the specimens themselves as they appear on paper,” as well as by Leng (1920: 322), Schenkling and
Marshall (1934: 38), and O’Brien and Wibmer (1982: 109).
Anthonomus profundus vulpinus Dietz: Blatchley and Leng 1916: 291
Recognition (Fig. 5). — This species is distinguished by the following combination of
characters:
1) metasternum and abdominal sterna, and meso- and metafemora without long setiform
scales;
2) metafemora of male stout, strongly curved, the single small ventral tooth displaced
distally;
3) metatibia of male straight, without midventral prominence;
4) sutural elytral interspaces with subapical prominences that are widely separated in
female (Fig. 39);
5) aedeagus strongly curved in distal 1 /3 (Fig. 50);
6) pygidial sulcus of male broad, slightly narrowed posteriorly, delimited apically (Fig.
33);
7) elytra with midbasal macula extended posteriorly on suture;
8) endophallus with sparse, minute denticles and a large sclerotized plate (Fig. 50).
Anthomorphus Weise
327
It resembles the Mexican A. galphimiae from which it is distinguished by the characters listed
in the section on that species. It is likely to be confused with the North American A. rubidus
but the two are distinguished by the characters listed in the section on the latter.
Description. — Male. Length: 2.8-4. 1 mm (x = 3.3, n=ll). Width-. 1. 2-2.0 mm (x=1.5, n = 1 1 ). Head-, eyes
separated by distance ca. 0.7 x width of rostrum at base. Rostrum: length 1.3-1. 5 x (x= 1.4, n = 1 1) pronotal length; most
strongly curved over antennal insertions; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion
shallowly rugulose-punctate, length 16-25% (x = 21, n = 1 1 ) of total rostral length. Thorax : pronotum with elongate,
acuminate, aeneous setiform scales and middorsal vitta of broader whitish scales. Legs: profemur ca. 1.4 x stouter than
metafemur, distal tooth obsolete or absent; protibia with small, acute preapical tooth; metatibial uncus short, curved,
excavated.
Female. Length: 2. 5-3. 7 mm (x=3.1, n = 9). Width: 1.2-1. 7 mm (x=1.5, n = 9). Rostrum: slender, short, length
1.3-1. 6 x (x=1.5, n = 9) pronotal length, most strongly curved over antennal insertions; proximal portion rugose, vestiture
limited to extreme base; distal portion smooth, length 28-37% (x=31, n = 9) of total rostral length. Pygidium (Fig. 39):
with strongly rounded apicodorsal prominence; median sulcus long, broad, slightly wider apically. Abdomen: sternum 5
with posterior margin straight.
Plant Associations. — According to Pierce (1907: 268) “Mr. Schwarz states that this
species breeds in Crataegus buds.” Schwarz (1890: 232) himself, however, stated that the
species “develops within the fruit of Crataegus crus-galli , the imago appearing in July.” Label
data indicate that specimens of A. profundus have been collected on the following plants (the
Larix record is probably an incidental association):
Rosaceae
Amelanchier canadensis (L.) Medic. (4 specimens, Beltsville, Maryland)
Aronia arbutifolia (L.) Ell. (1 specimen, Beltsville, Maryland)
Aronia " purpuria " (2 specimens, Pemberton, Burlington Co., New Jersey)
Crataegus sp. (2 specimens, Westerville, Franklin Co., Ohio)
Pinaceae
Larix laricina (Du Roi) K. Koch (1 specimen, Volo, Lake Co., Illinois)
Distribution. — This species occurs in eastern Canada and the northeastern United States,
westward to Illinois and southward to Texas, and in California and Oregon. The specimens
from McMinnville, Oregon, referred to by Hatch (1971: 350), were not examined. In addition
to the lectotype of A. profundus , which LeConte (1876: 198) stated was from Illinois, and the
lectotype of A. vulpinus from Pennsylvania, the following specimens were examined.
CANADA. Nova Scotia. Dartmouth (1 male, USNM). Ontario. Mer Bleue (1 male, 1 female, CNCI); Port Credit (1
female, USNM); Toronto (1 female, USNM).
UNITED STATES. California. Trinity Co.: Carrville (1 male, 1 female, CASC). Illinois. Lake Co.: Volo (1 female,
INHS). Indiana. Crawford Co.: (1 female, USNM). Marshall Co.: (1 male, PURC); Starke Co.: (1 male, PURC). Iowa.
Johnson Co.: Iowa City (1 male, 1 female, USNM). Maryland. Garrett Co.: Oakland (4 males, 5 females, USNM).
Prince Georges Co.: Beltsville (1 male, CNCI) (1 female, USNM) (2 males, 2 females, USNM) (1 male, 1 female,
USNM); Priest Bridge (1 male, USNM). Massachusetts. (2 males, 2 females, USNM). Berkshire Co.: North Adams (1
female, USNM). Hampden Co.: Springfield (1 male, USNM) 1 female, USNM). Middlesex Co.: Ashland (1 female,
CASC); Framingham (1 male, 3 females, USNM) (1 male, USNM); Natick (1 male, 1 female, CASC); Sherborn (1
female, CASC); Wilmington (1 female, USNM). Plymouth Co.: Marion (1 male, USNM). Michigan. Eaton Co.: Grand
Ledge (1 female, USNM). Ingham Co.: 2 mi. SW Holt (2 males, TAMU). Ottawa Co.: 1 mi. S. Grand Haven (1 male, 1
female, TAMU). New Jersey. Bergen Co.: Ramsey (2 males, USNM). Burlington Co.: Browns Mills (1 male, 1 female,
CNCI); Pemberton (2 females, USNM). Essex Co.: Caldwell (1 male, 1 female, USNM); Montclair (1 female, USNM);
Newark (1 male, 1 female, USNM). Gloucester Co.: Malaga (2 males, 1 female, USNM) (1 female, USNM); Midwood
(1 male, AMNH). Ocean Co.: Lakehurst (7 males, 5 females, USNM). Union Co.: Berkeley Heights (1 female, USNM);
Elizabeth, 27 Nov (1 female, USNM). New York. New York City (1 male, 6 females, USNM). Nassau Co.: Massapequa
(2 males, 1 female, USNM). St. Lawrence Co.: Cranberry Lake (1 female, USNM). Ohio. Franklin Co.: Westerville (1
male, 1 female, ELSC). Hamilton Co.: Cincinnati (1 female, USNM). Oregon (1 male, 2 females, USNM). Pennsylvania
(4 females, USNM). Clearfield Co.: Clearfield (2 females, USNM). Fayette Co.: Oliver (2 females, USNM). Lycoming
Co.: North Mount (1 female, USNM). Monroe Co.: Canadensis (2 females, USNM). Texas. (1 male, INHS). West
Quaest. Ent., 1987, 23 (3)
328
Clark
Virginia. Greenbriar Co.: White Sulphur Spring (2 males, USNM). Marion Co.: Fairmont (1 female, CASC). Pocahantas
Co.: Cranberry Glades (1 male, CMIV). The Specimens were collected in the months of April-August.
Anthonomus galphimiae, new species
Type Series. — Holotype : Mexico, male [MEXICO: Chiapas/ 27 km SE Teopisca/ 22 Sept. 1981/ Clark and
Coe] [collected on /Galphimia/ glauca Cav., det./W.R. Anderson, 1981] (USNM). Paratypes : Mexico, 4 males, 3 females
[MEXICO: Chiapas/27 Km SE Teopisca/ 22 Sept. 1981/ Clark and Coe] [collected on/ Galphimia/ glauca Cav., det./
W.R. Anderson 1981]; 1 male [MEXICO. Oax. Hwy 175/ 10 Km NE Oaxaca/ 1800 m 16.VI.1979/ H & A Howden],
Total paratypes, 8 (AUEM, HAHC, TAMU).
Recognition (Figs. 7). — This species is distinguished by the following combination of
characters:
1) metasternum and abdominal sterna, but not meso- and metafemora, with long, fine
erect, aeneous, setiform scales;
2) pygidial sulcus of male broad, not narrowed posteriorly, not delimited apically;
3) pygidial sulcus of female broad, deep, wider apically (Fig. 40);
4) aedeagus symmetrical, expanded apically (Fig. 51);
5) elytra with midbasal macula extended posteriorly on suture (Fig. 8);
6) endophallus with sparse, minute proximal denticles, a small distal field of denticles, a
small toothlike sclerite and lightly sclerotized tube (Fig. 51).
It resembles A. profundus in having the pygidial sulcus of the male and female unusually broad
and deep (Figs. 33, 39, 40), but is distinguished from that species by the coarser vestiture, less
strongly inflated, less strongly curved femora that have larger ventral teeth which are not
displaced distally, and by the shape of the aedeagus (Figs. 50, 51).
Description. — Male. Length: 3. 2-3. 5 mm (x = 3.4, n = 6). Width : 1. 5-1.7 mm (x=1.6, n = 6). Head : eyes
separated by distance ca. 0.8 x width of rostrum at base. Rostrum-, length 1.4-1. 5 x (x=1.4, n = 6) pronotal length;
broadly, evenly curved; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion shallowly
rugulose-punctate, length 19-23% (x=22, n = 6) of total rostral length. Thorax-, pronotum with elongate, acuminate,
ferruginous scales and middorsal vitta of broader whitish scales. Legs: profemur ca. 1.1 x stouter than metafemur, with
small, acute distal tooth; protibia with short, acute preapical tooth; metatibial uncus large, with basal prominence.
Female. Female-. 3. 2-3. 3 mm (x = 3.3, n = 3). Width : 1.6-1. 7 mm (x=1.6, n = 3). Rostum: length 1.4-1. 5 x (x=1.5,
n = 3) pronotal length; broadly, evenly curved; proximal portion rugose, vestiture limited to proximal 1/3; distal portion
smooth, length 37-39% (x = 38, n = 3) of total rostral length. Pygidium (Fig. 40): with broad, apicodorsal prominence;
medium sulcus delimited by narrow carina, with long, dense, setae. Abdomen: sternum 5 with posterior margin slightly
produced, with slight apicolateral prominences.
Plant Associations. — The type series was collected on Malpighiaceae ( Galphimia glauca
Cav.).
Distribution. — - This species is known only from the type series from Mexico.
Specific Epithet. — The name of this species is an anagram of the generic name of its host.
Anthonomus howdenorum, new species
Type Series. — Holotype: Venezuela, male [VENEZUELA: Tach.: 3300m. 55 km. NE/ San Cristobal/
V. 17-18. 1974/ H. & W. Howden] (HAHC). Paratypes: Venezuela, 1 male, 1 female [VENEZUELA: H. & A.
Howden]; 1 male [VENEZUELA: Tachira/ Pueblo Hondo/ 28 June 1983/ W.E. Clark and Clark]; 1 female [Paramo la
Negra/ Venezuela, Tachi-/ ra m./ 24-VI-1979] [B. Bechyne/ leg.]. Total paratypes, 4 (AUEM, HAHC, IZAV).
Recognition (Fig. 9). — This species is distinguished by the following combination of
characters:
1) metasternum and abdominal sterna, but not meso- and metafemora, with long, fine,
erect, setiform scales;
2) elytra with subfasciculate white, fulvous and fuscous scales;
Anthomorphus Weise
329
3) sutural elytral interspaces slightly expanded subapically;
4) aedeagus (Fig. 52) strongly asymmetrical;
5) endophallus with minute proximal denticles and a large toothlike sclerite (Fig. 52);
6) elytra with midbasal macula extended posteriorly on suture (Fig. 10).
The aedeagus of this species is similar in form to that of A. profundus (Figs. 50, 52). The
distinctive pattern of strongly differentiated, subfasciculate, white, fulvous and fuscous elytral
scales is unique among known Anthomorphus.
Description. — Male. Length-. 2.8-3. 1 (x = 2.9, n = 3). Width : 1.2-1. 4 mm (x=1.3, n = 3). Head: eyes separated by
distance ca. 0.8 x width of rostrum at base. Rostrum: length 1.4-1. 5 x (x=1.4, n = 3) pronotal length; most strongly
curved over antenna insertions; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion shallowly
rugulose-punctate, length 20-22% (x = 2 1 , n = 3) of total rostral length. Thorax: pronotum with elongate, acuminate
ferruginous scales and middorsal vitta of broader whitish scales. Pygidium: median sulcus narrow, wider and deeper
apically. Legs: profemur ca. 1.3 x stouter than metafemur, with small distal tooth; protibia with large, acute preapical
tooth; metatibial uncus slender, straight, oblique.
Female. Length: 3. 0-3. 2 mm (x=3.1, n = 2). Width: 1.4-1. 5 mm (x= 1.4, n = 2). Rostrum: stout, length 1.5 x (x= 1.5,
n = 2) pronotal length; slightly, evenly curved; proximal portion sulcate, vestiture extensive almost to antennal insertions;
distal portion smooth, length 31-32% (x = 32, n = 2) of total rostral length. Pygidium: with strongly rounded apicodorsal
prominence; median sulcus narrow, deep, not wider apically, with long, dense setae. Abdomen: sternum 5 with posterior
margin slightly produced medially.
Plant Associations. — Unknown.
Distribution. — This species is known only from the type series from Venezuela.
Specific Epithet. — This species is named in honor of Henry and Ann Howden of Ottawa,
Ontario, Canada, in appreciation of their friendship and encouragement.
Anthonomus varians (Paykull)
Curculio varians Paykull 1792: 16. Type Material: Sweden, not examined, lost (Diechmann 1968: 473).
Anthonomus varians (Paykull). Dieckmann 1968: 473-478 (synonyms listed).
Recognition (Figs. 11, 12). — This species is distinguished by the following combination of
characters:
1) elytra with sparse setiform scales;
2) eyes small, round, prominent;
3) femora strongly inflated;
4) rostrum slender, smooth, glabrous from base to tip;
5) tarsal claws simple, without basal tooth;
6) ventral margin of metatibia of male curved;
7) aedeagus symmetrical, narrowed distally (Fig. 53);
8) endophallus unarmed (Fig. 53).
It is distinguished from the other Palearctic species of Anthomorphus, A. pinivorax, by the
curved male metatibia (Dieckmann, 1968: Fig. 38), the unarmed tarsal claws and the
asymmetrical, distally narrowed aedeagus (Fig. 53). It resembles A. pinivorax in several
characters by which the two Palearctic species are distinguished from their New World
relatives. These are the sparse setiform elytral scales, the small round, prominent eyes, the
strongly inflated femora, the slender, smooth, glabrous rostrum and the unarmed endophallus
of the male genitalia.
Description. — (see Dieckmann, 1968: 475).
Plant Associations. — Dieckmann (1968: 475-477) reported that A. varians occurs on
Pinus sylvestris L., P. montana Mill., and occasionally on Picea abies L. Dieckmann also cited
various authors who stated that the larvae develop in the buds of the young shoots or in the
cones, and according to his own observations, in the male inflorescences.
Quaest. Ent., 1987,23 (3)
330
Clark
Distribution .— According to Dieckmann (1968: 477) A. varians is widespread throughout
Europe, the Near East, and central and eastern Siberia.
Anthonomus pinivorax Silfverberg
Anthonomus pinivorax Silfverberg 1977: 14. Replacement name for Curculio pubescens Paykull 1792: 12 (not Fabricius
1775: 131). Dieckmann 1968: 478-479 (lists synonyms). Lectotype: Sweden (designated by Dieckmann 1968: 478),
not examined.
Recognition. — This species is distinguished by the following combination of characters:
1) elytra with sparse setiform scales;
2) eyes small, round, prominent;
3) femur strongly inflated;
4) rostrum slender, smooth, glabrous from base to tip;
5) tarsal claws with basal tooth;
6) metatibia of male straight;
7) aedeagus symmetrical, widened distally (Fig. 54);
8) endophallus unarmed (Fig. 54);
9) abdomen of male with sternum 5 not subquadrately emarginate, without triangular,
setose sclerite isolated from posterior margin (Fig. 37).
It is distinguished from the other Palearctic species of Anthomorphus, A. varians , by the
straight male metatibia (Dieckmann, 1968: Fig. 39), the symmetrical aedeagus that is widened
distally (Fig. 54), and the toothed tarsal claws.
Description. — (see Dieckmann, 1968: 478-479).
Plant Associations. — Dieckmann (1968: 479) reported that A. pubescens occurs on Picea
abies L., Pinus sylvestris L. and P. nigra Arn. He stated that the eggs are laid in the buds of
the young apical shoots and, less frequently, in side shoots. The buds are completely destroyed
by the larval feeding. The presence of the weevils in the young shoots is evidenced by yellowing
of the needles. At the tips of the infested buds the larvae construct cells out of the excrement
and frass that also enclose the needles. Larvae were found in these cells in May and June, and
in July pupae were found there.
Distribution. — This species occurs in central, northern, eastern and southeastern Europe
(Dieckmann, 1968: 479).
ANTHONOMUS SULCATUS SPECIES GROUP
Recognition. — The members of this species group are distinguished by the following
characters:
1) spiculum gastrale of male genitalia with basal portion flat, not closely engaging
aedeagus, without ventral keel (Fig. 70);
2) aedeagus without midventral, subbasal lobe (Figs. 55-68).
Discussion. — This group of 14 Neotropical species is probably paraphyletic. Several
monophyletic subgroups of the group are evident, but relationships among these subgroups
cannot be satisfactorily resolved on the basis of observed characters. Known hosts of the
members of the group are species of Malpighiaceae.
Anthomorphus Weise
331
Anthonomus sulcipygus Champion
Anthonomus sulcipygus Champion 1903: 177. Lectotype (here designated): Panama, male [3] [Sp. Figured.] [V. de
Chiriqui/ below 4,000 ft./ Champion.] [B.C.A.Col.IV.4./ Anthonomus/ sulcipygus/ Champ.] (BMNH).
Paralectotypes : Panama, 2 males, 1 female [Caldera,/ 2400 ft.; Champion.] [(3) (32)] [B.C.A.Col.IV.4./
Anthonomus/ sulcipygus/ Champ.] (BMNH); 1 female [San Lorenzo,/ Panama./ Champion] [A sulcipygus/ Ch.]
(BMNH). Champion 1910: 186. Champion (1903: 177) stated that A. sulcipygus was represented by eight specimens,
all from Panama (“Volcan de Chiriqui” and Caldera). The eight syntypes examined included specimens labelled “V de
Chiriqui, ’’“Caldera,” and “San Lorenzo.” The latter locality (also in the Department of Chiriqui) was not mentioned
in the original description but presumably the specimen bearing that label was among the eight syntypes. Only four of
the original eight syntypes represent the species described here. The other three syntypes are A. sulcatus.
Recognition (Figs. 13, 14). — This species is distinguished by the following combination of
characters:
1) aedeagus with obliquely truncated apical projection (Fig. 55);
2) pygidial sulcus of female long, deep, well-defined (Fig. 41);
3) rostrum of female long, slender, strongly curved (Fig. 13);
4) endophallus with large proximal field of dense, minute denticles, a large tooth-like
sclerite and a more distal, serrate sclerite (Fig. 55).
Characters by which A. sulcipygus is distinguished from the related A. sulcatus and A.
pimentai are discussed under the treatment of the latter species.
Description. — Male. Length: 2. 8-3.4 mm (x=3.0, n = 9). Width : 1. 3-1.6 mm (x=1.4, n = 9). Head : eyes
separated by distance ca. 0.6 x width of rostrum at base. Rostrum: slender, length 1.6-1. 8 x (x=1.7, n = 9) pronotal
length; evenly curved; proximal portion rugose-punctate, sulci obsolete; distal portion Finely, sparsely punctate, length
24-30% (x = 27, n = 9) of total rostral length. Thorax : pronotum with elongate, narrow aeneous scales and middorsal vitta
of broader, whitish scales. Pygidium: median sulcus narrow, deeper and wider apically, not reaching apex. Legs: profemur
ca. 1 .4 x stouter than metafemur; distal tooth obsolete; protibia with acute preapical tooth; metatibial uncus curved, with
slight basal process.
Female. Length: 2.8-3. 2 mm (x=33.0, n=7). Width: 1.3-1. 5 mm (x=1.4, n = 7). Rostrum: length 1. 5-2.2 x (x=1.9,
n=7) pronotal length; strongtly, evenly curved; proximal portion rugose, vestiture limited to extreme base; distal portion
smooth, length 31-47% (x=40, n = 7) of total rostral length. Pygidium: with broadly rounded apicodorsal prominence;
median sulcus long, deep, widened apically, with long, dense setae. Abdomen: sternum 5 with posterior margin nearly
straight.
Plant Association. — Label data indicate that specimens of A. sulcipygus were collected on
Malpighiaceae, Byrsonima coccolobifolia H.B.K. (1 specimen, reared from flower bud,
Fazenda Campininha, Mun. Mogi-Gua^u, Sao Paulo, Brazil).
Distribution. — This species is represented by specimens from widely separated localities in
Mexico, Nicaragua, Panama and southern Brazil. In addition to the type series from Panama,
the following specimens were examined.
BRAZIL. Minas Gerais: Nova Lima, Sao Paulo: Fazenda Campininha, Mun. Mogi-Guaqu (1 female, MZSP).
MEXICO. Veracruz: Coyame, Catemaco (2 males, TAMU). The specimens on which Champion’s (1910: 186)
Managua, Nicaragua, record were based have not been examined. The specimens were collected in the months of
February, June and September.
Anthonomus chinculticensis , new species
Type Series. — Holotype: Mexico, male [MEXICO:/ Chiapas/ Chincultic Ruins/ August 29, 1982/ Clark and
Cave] (USNM).
Recognition. — This species is distinguished by the following characters:
1) aedeagus constricted subapically, rounded at narrow apex (Fig. 56);
2) endophallus with small, obtuse denticles, a large tooth, and a long serrate sclerite (Fig.
56).
It closely resembles A. sulcipygus but differs in characters of the male genitalia. In A.
chinculticensis the aedeagus is narrowed distally, then slightly expanded at the extreme apex
Quaest. Ent., 1987,23 (3)
332
Clark
(Fig. 56), whereas in A. sulcipygus the aedeagus is abruptly constricted to an obliquely
truncate apical prominence (Fig. 55).
Description. — Male. Length: 2.7 mm (n=l). Width : 1.4 mm (n=l). Head : eyes separated by distance ca. 0.6 x
width of rostrum at base. Rostrum : length 1.5 x (n=l) pronotal length; most strongly curved over antennal insertions;
proximal portion rugose-punctate, length 23% (n — 1 ) of total rostral length. Thorax : pronotum with elongate, acuminate,
fuscous scales, without middorsal vitta of broader, more pallid scales. Pygidium: median sulcus narrow, slightly wider and
deeper apically; with shallow, narrow, lateral sulci. Legs : profemur ca. 1.4 x stouter than metafemur, with small distal
tooth that shares common base with large ventral tooth; protibia with small preapical tooth; metatibial uncus large, curved,
oblique, with slight prominence.
Female. Unknowiu
Plant Association. — Unknown.
Distribution. — This species is known only from the type locality from southern Mexico.
Specific Epithet. — The name of this species is derived from the name of the type locality,
the site of a small Mayan pyramid.
Anthonomus vanini , new species
Type Series. — Holotype : Brazil, male [Rio de Janeiro/ Guanabara BRAZIL] [1X63] (MZSP). Paratypes:
Brazil, 2 males [Rio de Janeiro/ Guanabara BRAZIL] [1X63]. Total paratypes, 2 (AUEM, TAMU).
Recognition. — This species is distinguished by the following combination of characters:
1) aedeagus slightly asymmetrical, with narrow apical projection (Fig. 57);
2) endophallus with minute denticles and two large tooth-like sclerites (Fig. 57);
3) metatibia of male without setiform scales.
It resembles A. camoiranensis in possession of an acute apicomedian projection on the aedeagus
(Figs. 57, 58). It is distinguished from that species in lacking a dense apicoventral fringe of
setiform scales on the metatibia of the male.
Description. — Male. Length: 2.9-3.0 mm (x = 3.0, n = 3). Width: 1.2-1. 5 mm (x=1.4, n = 3). Head-, eyes
separated by distance ca. 0.7 x width of rostrum at base. Rostrum : length 1.4-1. 5 x (x= 1.4, n = 3) pronotal length; most
strongly curved over antennal insertions; proximal portion strongly carinate; distal portion finely, sparsely punctate, length
24-26% (x = 25, n = 3) of total rostral length. Thorax-, pronotum with elongate, narrow, aenescent scales and a narrow,
middorsal vitta of broader, more pallid whitish scales. Pygidium: median sulcus narrow, wider and deeper apically. Legs:
profemur ca. 1.3 x stouter than metafemur, with distinct distal tooth; protibia with preapical tooth; metatibial uncus
minute, acute.
Female. Unknown.
Plant Associations. — Unknown.
Distribution. — This species is known only from the type series from southern Brazil.
Specific Epithet. — This species is named for Sergio A. Vanin of Sao Paulo, Brazil, in
appreciation of his friendship and encouragement.
Anthonomus camoiranensis, new species
Type Series. — Holotype: Venezuela, male [VENEZUELA: Bolivar/ Gran Sabana (Camoiran)/ 19 June 1984/
W.E. Clark] [Byrsonima/ crassifolia/ (L.) H.B.K.] (USNM).
Recognition. — This species is distinguished by the following combination of characters:
1) metatibia of male with dense apicoventral brush of long, curved, setiform scales in
distal 1 /2;
2) metatibia of male without apical uncus;
3) aedeagus with narrow apical projection;
4) endophallus with small distal field of moderately large denticles, one slightly larger
than the other (Fig. 58).
Anthomorphus Weise
333
It is distinguished from the closely related A. vanini by characters listed in the treatment of
that species.
Description. — Male. Length-. 2.9 mm (n=l). Width : 1.2 mm (n=l). Head : eyes separated by distance ca. 0.6 x
width of rostrum at base. Rostum: length 1.4 x (n = 1 ) pronotal length; most strongly curved over antennal insertions;
proximal portion rugose-punctate, sulci obsolete; distal portion smooth, sparsely punctate, 35% (n=l) of total rostral
length. Thorax : pronotum with narrow, acuminate aeneous scales and middorsal vitta of broader, whitish scales.
Pygidium: median sulcus narrow, wider and deeper apically, slightly asymmetrical. Legs: profemur ca. 1.2 x stouter than
metafemur, with distinct distal tooth; protibia long, slender, without preapical tooth.
Female. Unknown.
Plant Association. — The holotype of A. camoiranensis was collected on Malpighiaceae
(Byrsonima crassifolia (L.) H.B.K.).
Distribution. — This species is known only from the type locality in southeastern Venezuela.
Specific Epithet. — The name of this species is derived from the name of the type locality.
Anthonomus stockwelli, new species
Type Series. — Holotype: Panama, male [Panama, C.Z./ Coco solo Hosp./ 9°2rN, 79°51'W/ 20 Jan. '72
Stockwell] (USNM). Paratypes: Panama, 1 male [Panama: Canal Zone/ Barro Colorado Is./ 9°10'N 70°50'W]
[16. VII. 1978/ E.M. Fisher]; 1 male [Panama: Colon Prov./ Santa Rita Ridge/ 9°22/N, 79°44'W/ 13 June, '76: E.G.
Riley] . Total paratypes, 2 (CH AH, C WOB) .
Recognition. — This species is distinguished by the following combination of characters:
1) head constricted behind large, prominent eyes (Fig. 15), transversely channelled
behind frons;
2) pygidial sulcus of male narrow, not delimited apically (Fig. 34);
3) aedeagus (Fig. 59) narrowed apically to long, acuminate point;
4) endophallus unarmed (Fig. 59);
5) body form narrow (Fig. 16).
Description. — Male. Length: 2. 2-3.0 mm (x = 2.7, n = 3). Width: 1. 3-1.4 mm (x=1.4, n = 3). Head: eyes
separated by distance ca. 0.6 x width of rostrum at base. Rostrum: length 1.1-1. 4 x (x = 1 .3, n = 3) pronotal length; most
strongly curved over antennal insertions; proximal portion rugose-punctate, sulci obsolete; vestiture obsolete slightly
proximad of antennal insertions; distal portion shallowly, finely punctate, length 17-28% (x= 17-28% n = 3) of total
rostral length. Thorax: pronotum with elongate, acuminate, fulvous to fulvo-ferruginous scales and middorsal vitta of
broader, more pallid whitish scales. Legs: profemur ca. 1.4 x stouter than metafemur, with distinct distal tooth; protibia
without preapical tooth; metatibial uncus minute, oblique.
Female. Unknown.
Plant Association. — Unknown.
Distribution. — This species is known only from the type series from Panama.
Specific Epithet. — The species is named for Henry P. Stockwell, collector of the holotype,
in honor of his contribution to the study of the Curculionidae.
Anthonomus filicornis Hustache
Anthonomus filicornis Hustache 1929: 257. Holotype: Guadeloupe, female [GUADELOUPE/ Gourbeyre/ L Dufau. A.
Hust.] [MUSEUM PARIS/ 1949/ Col. A. HUSTACHE] [Anth./ filicornis/ m.] (MNHP).
Recognition (Figs. 17, 18). — This species is distinguished by the following combination of
characters:
1) aedeagus asymmetrical, narrowed to obtusely pointed apex (Fig. 60);
2) endophallus with minute proximal denticles, with larger denticles and a small
tooth-like distal sclerite (Fig. 60);
3) pygidium of female produced and narrowed apically (Fig. 42);
4) posterior margin of female 5th abdominal sternum subtruncately produced;
Quaest. Ent., 1987,23 (3)
334
Clark
5) pygidial sulcus of male with long lateral setae.
Description. — Male. Length : 2. 4-2. 9 mm (x = 2.7, n=12). Width : 1.2-1. 4 mm (x = 1.3, n=12). Head: vertex
minutely punctate, with whitish, setiform scales; eyes prominent in profile, round, posterior margins not elevated,
separated by distance ca. 0.6 x width of rostrum at base. Rostrum : length 1.4-1. 7 x (x = 1.5, n=12) pronotal length;
slightly curved; proximal portion shallowly rugose, glabrous; distal portion sparsely, shallowly punctate, length 25-32%
(x = 28, n=12) of total rostral length. Thorax: pronotum with narrow, acuminate to setiform, fulvous to aenescent scales
and middorsal vitta of pallid, broader whitish scales. Pygidium: median sulcus narrow, deeper and wider apically.
Abdomen: sternum 5 with broad, subtruncate, posterior marginal prominence. Legs: profemur ca. 1.3 x stouter than
metafemur, distal tooth blunt; protibia with slight preapical tooth; metatibial uncus large, truncate.
Female (Figs. 17,18). Length: 2. 8-3.0 mm (x = 2.7, n=15). Width: 1.2-1. 5 mm (x=1.3, n=15). Rostrum: length
1.5-1. 8 x (x=1.6, n=15) pronotal length; broadly, evenly curved; proximal portion with sulci obsolete, glabrous, except
for setiform scales at extreme base; length of distal portion 30-41% (x = 36, n= 15) of total rostral length.
Plant Associations. — Label data indicate that A. filicornis has been collected on the
following plants:
Malpighiaceae
Byrsonima stipulacea Adr. Juss. (5 specimens, 8 km. S Kilometro 88, Bolivar,
Venezuela);
Byrsonima spicata (Cav.) DC (2 specimens, 3 km. W La Tigrera and 5 km. N Las
Trincheras, Bolivar, Venezuela).
Myricaceae
Myrica splendens (SW) DC (1 specimen, Sainte-Rose, Piton, Guadeloupe).
It seems most likely that the Byrsonima are true hosts but that the Myrica record is an
accidental association.
Distribution. — This species is represented by specimens from the Lesser Antilles, Panama,
Venezuela and Brazil. In addition to the female holotype from Guadeloupe, the following
specimens were examined:
BRAZIL. Minas Gerais: Prata (1 female, CCBM). Pernambuco: Caruaru (1 male, DZUP). Goias: Dianopolis (1
female, MZSP). Sao Paulo: Fazenda Pau d’Alho (1 male, MZSP).
GUADELOUPE. Sainte-Rose, Piton, 300 m (1 male, LFEV).
PANAMA. Panama: Cerro Campana (1 male, USNM). Code: 10 mi SW. Penonome (1 male, SWOB).
VENEZUELA. Bolivar: 8 km. S Kilometro 88 (3 males, 2 females, AUEM); 3 km. W La Tigrera (2 females,
AUEM); 5 km. N Las Trincheras (2 females, AUEM). Delta Amacuro: Isla C. Manamo, 25 km. S Tacupita (1 female,
CCBM). Guarico: Morrocoyes (1 male, IZAV). Lara: Jabon (1 male, AUEM). Portuguesa: Mesa de Cavacas (1 male, 1
female, AUEM). Yaracuy: 7 km. W Nirgua (9 females, AUEM, TAMU). The specimens were collected in the months of
February, April, June-August and November.
Anthonomus sulcatus Kirsch
Anthonomus sulcatus Kirsch 1874: 431. Holotype: Peru, female [Pozuzu / Kirsch] [TYPUS] [Staatl. Museum fur/
Tierkunde Dresden] [Anthonomus/ sulcatus Ksch.] (SMTD).
Recognition (Figs. 19, 20). — This species is distinguished by the following combination of
characters:
1) aedeagus with asymmetrical apical projection (Fig. 61);
2) endophallus with sparse mesal and distal denticles and a large, proximal sclerite (Fig.
61);
3) pygidial sulcus of male narrow, shallow (Fig. 35);
4) pygidium of female with broadly rounded apical prominence, sulcus short, shallow
(Fig. 44).
It is distinguished from the similar A. filicornis by the male genitalia (Figs. 60, 61) and the
female pygidium (Figs. 42, 44). Characters by which A. sulcatus is distinguished from the
Anthomorphus Weise
335
related A. sulcipygus and A. pimentai are listed under the treatment of the latter.
The genitalia of the males of A. sulcatus from Mexico and Panama differ from those from
Brazil in having the apex somewhat constricted before the asymmetrical apical prominence.
There is uncertainty about the identity of this species because the holotype is a female and
the reliability of the diagnostic characters listed cannot be ascertained while the females of
related species (A. vanini, A. camoiranensis, A. stockwelli, and A. chinculticensis ) remain
unknown.
Description. — Male. Length: 2. 0-2. 8 mm (x = 2.5, n=10). Width : 1.0-1. 4 mm (x=1.2, n=10). Head : eyes
prominent, separated by distance ca. 0.6 x width of rostrum at base. Rostrum : slender, length 1.3-1. 6 x (x=1.5, n=10)
pronotal length; broadly, evenly curved; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion finely,
sparsely punctate, length 25-34% (x = 29, n=10) of total rostral length. Thorax-, pronotum with narrow, aeneous scales
and narrow middorsal vitta of broader, more pallid scales. Pygidium (Fig. 35): median sulcus narrow, curved, slightly
wider apically. Legs-, profemur ca. 1.2 x stouter than metafemur; distal tooth obsolete; protibia with long, acute preapical
tooth; mesotibia with ventral, subapical prominence; metatibial uncus short, truncate.
Female. Length : 2.3-2. 8 mm (x=2.6, n= 15). Width: 1. 1-1.4 mm (x= 1.2, n= 15). Rostrum : slender, length 1.3-1. 7 x
(x=1.5, n=15) pronotal length; broadly, evenly curved; distal portion smooth, length 32-44% (x = 37, n=15) of total
rostral length. Abdomen : posterior margin of sternum 5 slightly produce medially.
Plant Association. — Label data indicate that the host plant of A. sulcatus is:
Malpighiaceae, Heteropterys xanthophylla Adr. Juss. (3 females, reared from flower buds,
Belo Horizonte, Minas Gerais, Brazil).
Distribution. — This species is known from widely separated localities in Mexico, Panama,
Peru and Brazil. In addition to the female holotype from Peru, the following specimens were
examined.
BRAZIL. Mato Grosso: Chapada dos Guimaraes (1 female, MPEG). Minas Gerais: Belo Horizonte, UFMG Campus
(3 females, AUEM, MZSP); Cachoeira do Campo (3 females, AUEM, MZSP); ca 5 km. S Concei?ao do Mato Dentro (4
females, AUEM, MZSP); Sao Sebastao das Aguas Claras, Nova Lima (5 males, 20 females, AUEM, MZSP, TAMU);
Sao Sebastao das Aguas Claras, Nova Lima (2 females, AUEM, MZSP); Sao Sebastao das Aguas Claras, Nova Lima (1
male, 2 females, AUEM, MZSP); Serro (4 females, AUEM, MZSP). Rio de Janeiro: Mt. Corcovado (1 male, 2 females,
TAMU). Sao Paulo: Fazenda Pau d'Alho, Itu (1 female, MZSP); Ilha dos Buzios (1 male, MZSP). Santa Caterina: Nova
Teutonia (1 female, ELSC).
MEXICO. Guerrero: 6 mi. NE Taxco (1 male, CWOB).
PANAMA, (from the type series of A. sulcipygus , BMNH) 1 female [(2)] [V. de Chiriqui,/ 25-4000 ft./ Champion.]
[B.C.A.Col.IV.4./ Anthonomus/ sulcipygus/ Champ.] [Type]; 1 male, 2 females [(32)] [V. de Chiriqui,/ 25-4000 ft./
Champion.] [B.C.A.Col.IV.4./ Anthonomus/ sulcipygus/ Champ.]. The specimens were collected in January and
September-December.
Anthonomus pimentai, new species
Type Series. — Holotype: Brazil, male [Brazil: Minas Gerais/ Sao Sebastao das Aguas/ Claras, Nova Lima / 3
Sep 1984 W.E. Clark] (MZSP). Paratypes: Argentina, 1 male [Puerto Iguazu, m 200, MISIONES/ ARGENT. Bordon/
leg. 27 XII 1980], Bolivia, 1 female [BOLIVIA, S.C., 10/ mi. W. Portachuelo/ March 24, 1978 at/ night CW&L
O'Brien]. Brazil, 2 males, 11 females [Brazil: Minias Gerais/ Sao Sebastao das Aguas/ Claras, Nova Lima/ 3 Sep 1984
W.E. Clark]; 1 female [BRAZIL: M.G., Sao/ Sebastao das Aguas/ Claras, Nova Lima/ 16 Dec 1980 R.P. Martins]
[Taken on/ Malpighiaceae/ Banisteriopsis?]; 1 female [Itapetinga/ Bahia Brazil]; 1 female [BRAZIL: Minas Gerais/
Belo Horizonte UFMG/ Campus 4 Oct 1985/ Helco R. Pimenta] [reared from flower/ buds Heteropterys/ umbellata
Adr. Juss. (Malpighiaceae)]; 1 female [BRAZIL: Minas Gerais/ Belo Horizonte UFMG/ Campus 10 Oct 1985/ Helcio
R. Pimenta] [reared from flower/ buds Tetrapterys/ humilis Adr. Juss. (Malpighiaceae)]; 1 male, 8 females [Brazil:
Minas Gerais/ Belo Horizonte UFMG/ campus/ N.S. Domingos] [( Stigmaphyllon / lalandianum) / (C15 19 Apr 82)
(C13 3 Apr 81) (C13 13 Nov 81) (C4 31 Aug 81) (C15 12 Apr 82) (C13 04 Apr 82) (Banisteriopsis/ malifolia/ (15 12
Apr 82) (B17 14 Jan 82)]; 6 females [BRAZIL: Minas Gerais/ Belo Horizonte UFMG/ Campus, Sep 1985/ Helcio R.
Pimenta] [reared from flower/ buds Heteropterys/ xanthophylla Adr. Juss. (Malpighiaceae)]; 4 females [BRAZIL:
Minas Gerais/ Belo Horizonte UFMG/ Campus, 3 Sep 1985/ Helcio R. Pimenta] [reared from flower/ buds
Tetrapterys/ humilis Adr. Juss./ (Malpighiaceae)]; 5 females [BRAZIL: Minas Gerais/ Belo Horizonte UFMG/
Campus, 4 Oct 1985/ Helico R. Pimenta] [reared from flower/ buds Heteropterys/ umbellata Adr. Juss./
(Malpighiaceae)]; 2 males [BRAZIL: Minas Gerais/ Belo Horizonte UFMG/ Campus, 4 Oct 1985/ Helcio R. Pimenta]
[reared from flower/ buds Tetrapterys/ humilis Adr. Juss./ (Malpighiaceae)]; 7 females [BRAZIL: M.G., Belo/
Quaest. Ent., 1987, 23 (3)
336
Clark
Horizonte, UFMG/ Campus 18 May 1985/ R.P. Martins] [reared from flower/ buds Banisteriopsis/ oxyclada Adr.
Juss.)./ Gates (Malpighiaceae)]; 2 males, 1 female [BRAZIL: M.G., Belo/ Horizonte, UFMG/ Campus 27 Apr 1985/
R.P. Martins] [reared from flower/ buds Banisteriopsis/ oxyclada Adr. Juss)./ Gates (Malpighiaceae)]; 1 female
[BRAZIL: M.G., Belo/ Horizonte, UFMG/ Campus 27 Apr 1985/ R.P. Martins] [reared from; Stigmaphyllon/ sp.]; 1
female [Itapetinga/ Bahia Brazil]; 2 males, 5 females [BRAZIL, Mato Grosso/ Sinop (12°31'S,/ 55°37/W) X 1974/ M.
Alvarenga]; 1 male, 1 female [MACAIBA/ R.G. Norte BRASIL/ 22.VII.1951/ M. Alvarenga legit.] [Ex cole<?ao/ M.
Alvarenga]; 6 males, 9 females [Macaiba/ RN Brasil/ (1. XII. 1951) (XII. 1951)/ M. Alvarenga col.]; 3 males, 5 females
[NATAL/ R.G. Norte Brasil/ VII 1951/ M. Alvarenga leg.] [Ex cole^ao/ M. Alvarenga]; 2 males, 1 female [Nova
Teutonia/ SC, Brasil/ XII.1965/ F. Plaumann col.]; 8 males, 8 females [Nova Teutonia,/ Santa Catarina, Brasil
(XII— 1 5—53) (1-5-55) (1-25-55) (XII-25-55) (XII-26-56) (1-15-57) (1-25-57)] [F. Plaumann/ Collr.] [E.L. Sleeper/
Collection]; 1 male [BRASIL: S. Catarina/ Nova Teutonia/ 300-500 m/ 27°irS 52°23'W] [-.X. 1974/ F. Plaumann]; 1
male, 2 females [Dpt0 Zool/ UF-PARANA] [Brasilien/ Nova Teutonia/ 27°11'B. 52°23'L/ Fritz Plaumann/ 300 .
500m/ (X, 1969) (XII 1980)]; 5 males, 6 females [Brasilien/ Nova Teutonia/ 27°11'B. 52°23'L / Fritz Plaumann/ (I
1954) (III 1954) (IV 1954) (IX 1954) (XII 1955)/ 300 . 500 m]; 1 male [Brasilien/ Nova Teutonia/ 27°11/B. 52°23'L/
Fritz Plaumann/ XI 1942/ 300 bis 500 m]; 1 male [S. Roque SP/ 25 XII. 71/ F. Lane Col.]; 1 male [7. VII. 1955/
Barueri/ S. Paulo/ 5465] [K. Lenko leg]; 1 female [BRAZIL: Sao Paulo/ Botucatu, Cerrado/ de Usina 8 Apr 1980/ P.S.
Oliveira] [Taken on/ Byrsonima/ coccolobifolia]; 1 female [Barueri/ S. Paulo - Brasil/ 18. IV. 1955/ K. Lenko col]. Total
paratypes, 128 (AUEM, CCBM, CWOB, DZUP, ELSC, MZSP, TAMU).
Recognition (Figs. 21, 22). — This species is distinguished by the following combination of
characters:
1) profemur long, stout, ventral tooth displaced distally;
2) pro- and mesocoxae and bases of profemora of male with long setiform scales;
3) aedeagus with broad, asymmetrical, emarginate, apical projection (Fig. 62);
4) rostrum of female long, slender, strongly curved;
5) pygidial sulcus of male broad, deep, carinate medially (Fig. 36);
6) endophallus with minute proximal denticles and a long, lightly sclerotized tube (Fig.
62).
The characters of the coxae and male genitalia distinguish males of A. pimentai from those of
the related A. sulcatus and A. sulcipygus. The females of A. pimentai are distinguished from
females of the other two species by the relatively long rostrum. Additionally, the profemur is
considerably less stout in A. sulcipygus and also has a well-developed tooth distal to the large
ventral tooth. Conversely, in A. sulcatus , the profemur is stout, but the stoutness is not
displaced distally and there is an obsolete tooth distal to the large ventral tooth. A. pimentai is
also distinguished from A. sulcatus and A. sulcipygus by characters of the female pygidium,
but these are difficult to assess without dissection. In A. pimentai the impression is shallow but
near the apex (Fig. 43) and visible even in specimens in which the pygidium is but narrowly
exposed. In A. sulcipygus the pygidial channel is long, deep, and relatively well-defined (Fig.
41), whereas in A. sulcatus the channel is short, shallow, and displaced anteriorly so that it is
visible only in specimens with the pygidium broadly exposed (Fig. 44).
The males of A. pimentai from Santa Catarina and from Argentina have shorter setiform
scales on the pro- and mesocoxae than do those from Minas Gerais.
Description. — Male. Length : 2. 4-3.0 mm (x = 2.8, n=15). Width: 1.1-1. 4 mm (x=1.3, n=15). Head: eyes
separated by distance ca. 0.7 x width of rostrum at base. Rostrum: length 1.6-1. 8 x (x= 1.7, n = 15) pronotal length;
broadly, evenly curved; proximal portion sulcate, strongly carinate; distal portion Finely, sparsely punctate, length 22-32%
(x=27, n=15) of total rostral length. Thorax: pronotum with elongate, narrow, aenescent scales and middorsal vitta of
broader, more pallid whitish scales. Legs: profemur ca. 1.4 x stouter than metafemur; protibia with obsolete preapical
tooth; metatibial uncus minute, acute.
Female. Length: 2. 3-3.4 mm (x = 2.8, n=15). Width: 1.1-1. 7 mm (x=1.4, n=15). Rostrum: long, slender, length
2.0-2. 3 x (x=2.2, n=15) pronotal length; strongly, evenly curved; proximal portion shallowly sulcate, vestiture limited to
extreme base; distal portion smooth, length 30-49% (x=43, n = 15) of total rostral length. Pygidium (Fig. 43): with broad,
rounded, apicomedian prominence. Abdomen: sternum 5 with posterior margin slightly produced medially.
Plant Associations. — There are no published host records. Label data indicate that A.
pimentai is associated with the following plants:
Anthomorphus Weise
337
Malpighiaceae
Banisteriopsis malifolia (Nees & Mart) Gates (3 specimens, Belo Horizonte, Minas
Gerais, Brazil)
Banisteriopsis oxyclada Adr. (Juss.). Gates (10 specimens, reared from flower buds,
Belo Horizonte, Minas Gerais, Brazil)
Byrsonima coccolobifolia H.B.K. (1 specimen, Cerrado de Usina, Botucatu, Brazil)
Heteropterys umbellata Adr. Juss. (6 specimens, reared from flower buds, Belo
Horizonte, Minas Gerais, Brazil)
Heteropterys xanthophylla Adr. Juss. (6 specimens, reared from flower buds, Belo
Horizonte, Minas Gerais, Brazil)
Stigmaphyllon lalandianum (6 specimens, Belo Horizonte, Minas Gerais, Brazil)
Stigmaphyllon sp. (1 specimen, reared from flower bud, Belo Horizonte, Minas Gerais,
Brazil)
Tetrapterys humilis Adr. Juss. (7 specimens, reared from flower buds, Belo Horizonte,
Minas Gerais, Brazil)
Distribution. — This species is known only from the type series from southern Brazil,
and northern Argentina.
Specific Epithet. — This species is named for Helcio R. Pimenta of Belo Horizonte,
as a token of appreciation for his help and friendship.
Anthonomus bechyneorum, new species
Type Series. — Holotype: Brazil, [Rio de Janeiro/ Guanabara BRAZIL] [VIII63] (MZSP). Paratypes : Brazil, 1
male, 3 females [Rio de Janeiro/ Guanabara BRAZIL] [ VIII63] ; 1 female [Guanabara, Rio/ de Janeiro Brazil]; 1 male
[Corumba de Goias/ GO, Brasil/ 3 1.1. 3. II. 1962/ J. Bechyne col.]; 1 male, 1 female [Caraguatatuba - SP/ (Res. Flor. - 40
m )/ 2-IV-962 - Martins/ Reichardt & Silva]; 1 male [Caraguatatuba - SP/ (Res. Flor. - 40 m.)/ 22-V. -I. VI. 1962/
Exp. Dep. Zool.]. Colombia, 1 male, 1 female (COLOMBIA) [Villavicencio/ 23-1-76/ J.A. Jimenez] [Semilla/ arbol/
maderable]. Honduras, 1 male [HONDURAS, Atl./ Liberia, 5-IX-1984/ C.W. O'Brien, rainforest]; 1 female
[HONDURAS, Atl./ La Ceiba, CURLA/ 30 Aug. 1984/ C.W. O'Brien]. Trinidad, 2 males [6-1/2 mi. Post, Maracas/
Bay, Trinidad,/ W. I. Aug (8) (13), 1969/ H. & A. Howden]. Venezuela, 3 females [VENEZUELA: Tachira / Pueblo
Nuevo/ 29 June 1983; W.E. Clark and Clark]. Total paratypes, 17 (AUEM, CWOB, HAHC, MZSP, USNM).
Recognition (Fig. 23). — This species is distinguished by the following combination of
characters:
1) elytra with dark middorsal macula (Fig. 24);
2) prothorax with broad scales and narrower, setiform scales;
3) scales on rostrum dense almost to antennal insertions;
4) scales on elytra not fasciculate;
5) aedeagus with asymmetrical, emarginate apical prominence (Fig. 63);
6) endophallus minutely denticulate (Fig. 63);
7) meso- and metafemora without long setiform scales;
8) pygidial sulcus of male shallow, narrow, delimited apically.
The male genitalia of A. bechyneorum are similar to those in A. haliki (Figs. 63, 64), but A.
bechyneorum is smaller in size, has more distinct middorsal elytral macula (Figs. 24, 26), and
lack long setiform scales on the male meso- and metafemora.
The males of A. bechyneorum from Honduras lack the apicomedian emargination of the
apical prominence of the aedeagus.
Bolivia
Brazil,
Quaest. Ent., 1987, 23 (3)
338
Clark
Description. — Male. Length-. 2. 3-3. 3 mm (x = 3.0, n = 7). Width : 1.0-1. 5 mm (x=1.4, n = 7). Head-, eyes
separated by distance ca 0.8 x width of rostrum at base. Rostrum: length 1.3-1. 5 x (x= 1.5, n = 7) pronotal length; most
strongly curved over antennal insertions; proximal portion rugose-punctate, sulci and carinae obsolete; distal portion
shallowly rugulose-punctate, length 19-33% (x = 25, n = 7) of total rostral length. Thorax : pronotum with elongate,
acuminate to setiform aeneous scales and with middorsal vitta of broader, more pallid fulvous scales. Pygidium: median
sulcus narrow, wider and deeper apically. Legs: profemur ca. 1.4 x stouter than metafemur, with small, blunt distal tooth;
protibia with short preapical tooth; metatibial uncus large, curved, with basal prominence.
Female. Length: 3.0-3. 6 mm (x = 3.3, n = 9). Width: 1.4-1. 6 mm (x=1.5, n = 9). Rostrum: stout, length 1.5-1. 7 x
(x=1.6, n = 9) pronotal length; slightly, evenly curved; proximal portion rugose-punctate, vestiture extensive almost to
antennal insertions; distal portion smooth, length 25-32% (x=30, n = 9) of total rostral length. Pygidium: with rounded
apicodorsal prominence; median sulcus ovate, shallow, with spare, elongate scales. Abdomen: sternum 5 with posterior
margin nearly straight.
Plant Associations. — Unknown.
Distribution. — This species is known from the type series from widely separated localities in
Central and South America.
Specific Epithet. — This species is named for J. and B. Bechyne. Their efforts have resulted
in great enrichment of collections of South American Curculionidae and other insects.
Anthonomus haliki, new species
Type Series. — Holotype: Brazil, male [Atibaia/ Sao Paulo./ 18.XI 1971./ F. Halik/ 11655] (MZSP).
Paratypes: Brazil, 2 males, 5 females [Atibaia/ Sao Paulo./ (26.XI 1971.) (18.XI 1971) (17.XI 1971)/ F. Halik/ (11607)
(11608) (11654) (1 1676) (11677) (1 1678) (11679)]. Total paratypes, 7 (AUEM, MZSP).
Recognition (Figs. 25, 26, 64). — This species is distinguished by the following combination
of characters:
1) meso- and metafemora of male with long, fine, setiform scales;
2) pronotum and elytra with pallid olivaceous and slightly narrower aeneous scales;
3) elytral strial punctures oversized, giving pallid scales slightly fasciculate appearance
(Fig. 26);
4) males with scales on rostrum extensive almost to antennal insertions;
5) aedeagus with asymmetrical apical prominence (Fig. 64);
6) endophallus without denticles, with a large, distal, tooth-like sclerite (Fig. 64);
7) pygidial sulcus of male narrow, delimited proximad of apex.
This relatively large Anthomorphus closely resembles A. bechyneorum in the shape of the
aedeagus (Figs. 63, 64). The two species are distinguished by the larger body size, and smaller,
less distinct middorsal elytral macula of A. haliki (Figs. 24, 26). In addition, the meso- and
metafemora of the male of A. haliki have long setiform scales lacking in A. bechyneorum.
Description. — Male. Length: 4.0-4. 1 mm (x = 4.1, n = 3). Width: 2.0 mm (x = 2.0, n = 3). Head: eyes separated by
distance ca. 0.9 x width of rostrum at base. Rostrum: length 1.6-1. 7 x (x=1.7, n = 3) pronotal length; slightly, evenly
curved; proximal portion rugose-punctate, sulci obsolete; distal portion rugulose-punctate, length 18-19% (x= 19, n = 3) of
total rostral length. Legs: profemur ca. 1.4 x stouter than metafemur, with minute distal tooth that arises from base of
large ventral tooth; protibia with short preapical tooth; metatibial uncus large, curved, with basal prominence.
Female. Length: 4. 0-4. 2 mm (x=4.2, n = 5). Width: 1. 9-2.1 mm (x=2.0, n = 5). Rostrum: length 2. 1-2.2 x (x=2.1,
n = 5) pronotal length; slightly, evenly curved; proximal portion rugose-punctate, vestiture obsolete well proximad of
antennal insertions; distal portion rugulose, length 36-40% (x = 38, n = 5) of total rostral length. Pygidium: with rounded
apicodorsal prominence; median sulcus ovate, shallow, with sparse, elongate scales. Abdomen: sternum 5 with posterior
margin nearly straight.
Plant Associations. — Unknown.
Distribution. — This species is known only from the type series from southern Brazil.
Specific Epithet. — This species is named for its collector, F. Halik.
Anthomorphus Weise
339
Anthonomus rulfoi, new species
Type Series. — Holotype: Mexico, male [MEXICO: Tamaulipas/ 4 mi. W.C. Victoria/ (Canon del Navillo)/
November 14, 1985/ P. Kovarik, R. Jones/ and K. Haack] (USNM). Paratypes: Mexico, 1 male, 2 females [MEXICO:
Tamaulipas/ 4 mi. W.C. Victoria/ (Canon del Navillo)/ November 14, 1985/ P. Kovarik, R. Jones/ and K. Haack]; 1
male [Tamazunchale/ 1.28.53 S(an) L(uis) Potosi)./ D.G. Kissinger]; 1 male [Jalapa, Mex./ J.T. Mason]. Total
paratypes, 5 (TAMU, USNM).
Recognition (Figs. 27, 28). — This species is distinguished by the following combination of
characters:
1) metasternum, abdominal sterna and meso- and metafemora with long, fine, setiform
scales;
2) metatibia of male with long, straight, ventral setiform scales in distal 2/3;
3) aedeagus abruptly widened in distal 1 /4 to large lateral lobes that give way distally to
apical projection with apicolateral projections (Fig. 65);
4) endophallus with minute denticles and with a large tooth (Fig. 65);
5) distal profemoral tooth obsolete.
Description. — Male. Length: 2. 5-2.6 mm (x = 2.6, n = 3). Width : 1.2 mm (n = 3). Head : eyes separated by
distance ca. 0.6 x width of rostrum at base. Rostrum : length 1.5-1. 6 x (x=1.6, n = 3) pronotal length; most strongly
curved over antennal insertions; proximal portion shallowly sulcate, feebly carinate; distal portion shallowly
rugulose-punctate, length 20-21% (x = 20, n = 3) of total rostral length. Thorax: pronotum with elongate, acuminate
ferruginous scales and middorsal vitta of broader, white scales. Pygidium: median sulcus broad, slightly wider and deeper
apically. Legs: profemur ca 1.4 x stouter than metafemur, with minute distal tooth that shares common base with larger
tooth; protibia without preapical tooth; metatibia with minute, oblique apical uncus.
Female. Length: 2. 4-2. 5 mm (x — 2.4, n = 2). Width: 1.2 mm (n = 2). Rostrum: slender, length 1.5 x (x=1.5, n = 2)
pronotal length; slightly, evenly curved; proximal portion feebly sulcate, vestiture limited to extreme base; distal portion
smooth, length 37-48% (x = 43, n = 2) of total rostral length. Pygidium: subquadrate apically, broadly emarginate
apicomedially; median sulcus broad, shallow. Abdomen: sternum 5 with posterior margin slightly produced medially.
Plant Associations. — Unknown.
Distribution. — This species is known only from the type series from northeastern Mexico.
Specific Epithet. — This species is named for the late Mexican author Juan Rulfo.
Anthonomus abdominalis Champion
Anthonomus abdominalis Champion 1903: 178. Holotype: Panama, female [San Feliz,/ Panama ./ Champion] [($)]
[B.C.A.Col.IV.4./ Anthonomus/ abdominalis,/ Champ.] [Type] (BMNH).
Recognition (Figs. 29, 30). — This species is distinguished by the following combination of
characters:
1) elytra with common, middorsal fascia of dark fuscous scales (Fig. 30);
2) protibia of male with sparse fringe of long, setiform scales in distal 1 /2;
3) aedeagus with asymmetrical apical prominence (Fig. 66);
4) endophallus without denticles, with a small tooth (Fig. 66);
5) pygidial sulcus of male narrow, shallow.
Description. — Male. Length: 2. 2-2. 3 mm (x = 2.3, n = 2). Width: 1. 0-1.1 mm (x=1.0, n = 2). Head: eyes
separated by distance ca. 0.6 x width of rostrum at base. Rostrum: length 1.4-1. 5 x (x=1.4, n = 2) pronotal length;
broadly, evenly curved; proximal portion shallowly, irregularly sulcate, feebly carinate; distal portion finely, sparsely
punctate, length 23-25% (x = 24, n = 2) of total rostral length. Thorax: pronotum with elongate, acuminate, ferruginous
scales and middorsal vitta of broader, whitish scales. Pygidium: median sulcus narrow, slightly wider and deeper apically.
Legs: profemur ca. 1.4 x stouter than metafemur, with small distal tooth; protibia without preapical tooth; metatibial
uncus large, oblique, excavated, with slight prominence.
Quaest. Ent., 1987,23 (3)
340
Clark
Female. Length : 2. 0-2. 8 mm (x = 2.5, n = 1 1 ). Width : 1.0-1. 4 mm (x=1.2, n=ll). Rostrum : slender, length 1.4-1. 6 x
(x= 1.5, n= 1 1) pronotal length; slightly, evenly curved; proximal portion feebly sulcate, vestiture limited to extreme base;
distal portion smooth, length 35-41% (x = 38, n=ll) of total rostral length. Pygidium (Fig. 45): with narrow, rounded
apicodorsal prominence; median sulcus broad, deep, wider apically. Abdomen : sternum 5 with posterior margin slightly
produced medially.
Distribution. — This species is known from Panama, Trinidad, Venezuela and Brazil. In
addition to the female holotype from Panama, the following specimens were examined.
BRAZIL. Rondonia : Vilhena (1 female, MPEG). Sao Paulo : Barueri (1 female, MZSP).
PANAMA. Canal Zone : Barro Colorado Island (4 females, CWOB); Barro Colorado Island (2 females, CWOB);
Albrook Forest Site, Fort Clayton (1 male, CWOB); 5 mi. NW Gamboa (1 male, 2 females, USNM). Panama : Pacora (1
female, USNM); Panama (1 female, USNM). TRINIDAD. Morne Bleu (1 female, HAHC). VENEZUELA. Aragua :
4.5 km. SE Villa de Cura (1 female, AUEM). The specimens were collected in February, April, May, July, August, and
October-December.
Anthonomus diamantinaensis , new species
Type Series. — Holotype: Brazil, male [Brazil: Minas Gerais/ Diamantina/ 5 September 1984/ W.E. Clark]
(MZSP). Paratypes : Brazil, 3 males, 10 females [Brazil: Minas Gerais/ Diamantina/ 5 September 1984/ W.E. Clark]; 1
male [Brazil: Minas Gerais/ Sao Sebastao das Aguas/ Claras, Nova Lima / 3 Sep 1984, W.E.- Clark]; 1 male, 1 female
[BRAZIL: M.G. Sao/ Sebastao das Aguas/ Claras, Nova Lima/ 16 Dec 80 R.P. Martins] [Taken/ on/ Malpighiaceae/
Banisteriopsis?]; 1 male, 1 female [BRAZIL: M.G., Sao/ Sebastao das Aguas/ Claras, Nova Lima / 27 Feb 81 R.P.
Martins] [Taken/ on/ Malpighiaceae]; 1 male, 3 females [BRAZIL: Minas Gerais/ Belo Horizonte UFMG/ Campus, 4
Oct 1985/ Helcio R. Pimenta] [reared from flower/ buds Heteropterys/ umbellata Adr. Juss./ (Malpighiaceae)]; 2
males, 1 female [BRAZIL: M.G./ Cardeal Mota/ 17 Sept. 1985/ Clark & Martins]; 2 males [BRAZIL: M.G./
Diamantina/ 13 Sept. 1985/ Clark & Martins]; 1 male [BRAZIL: M.G./ Lagoa Santa/ 12 Sep 80, b.f.lll/ R.P.
Martins] [?Taken/ on/ Banisteria/ maritiana]; 3 females [BRAZIL: Minas Gerais/ MG040 Lagoa Santa/ 18 Oct 1985
R.P. Martins/ Helcio R. Pimenta] [reared from flower/ buds Heteropterys/ umbellata Adr. Juss./ (Malpighiaceae)]; 1
male [BRAZIL: M.G./ Mendanha/ 13 Sept. 1985/ Clark & Martins]; 3 males, 2 females [BRAZIL: M.G./ Serro/ 12
Sept. 1985/ Clark & Martins]; 1 female [Pirineus/ GO, Brasil/ 2.11.1962/ J. Bechyne col.]; 2 females [Cristalina,
1200m/ GOIAS, Brasil/ Bordon 16 XI 83]; 1 female [BRASIL: Est. Sao Paulo, Sao Paulo/ 9.XII.1965/ V.N. Alin]; 1
female [BRASIL: Est. Sao/ Paulo, Sao Paulo/ 2. 1968/ V.N. Alin]; 1 male, 2 females [Brazil: Sao Paulo/ Faz.
Campininha/ Mun. Mogi-Guaqu/ 15 November 1979] [R.P. Martins/ Byrsonima/ intermedia/ Cerrado]; 1 female
[BRAZIL: Sao Paulo/ Botucatu, Cerrado/ de Usina 8 Apr 1980/ P.S. Oliveira] [Taken on/ Byrsonima/ coccolobifolia];
3 males, 1 female [BRAZIL: Sao Paulo/ Fazenda Campininha/ Mun. Mogi-Guacu/ 15 Feb 1979 R.P. Martins] [Taken
on/ B./ intermedia]; 1 male [DPTQ ZOOL/ UF-PARANA] [BATATAIS - SP./ BRASIL 4/1/67/ Pe. J. Moure leg]; 1
female [DPT0 ZOOL/ UF-PARANA] [BATATAIS - S P/ BRASIL - 6-II-1966]; 1 female [Faz. Pau d’Alho/ Itu, SP,
Brasil, XI. 1960/ U R. Martins col.]. Total paratypes, 52 (AUEM, CCBM, CHAH, CWOB, DZUP, MZSP, TAMU).
Recognition (Figs. 31, 32). — This species is distinguished by the following combination of
characters:
1) pygidium without median sulcus (Fig. 46);
2) aedeagus abruptly constricted subapically, with slight apical projection (Fig. 67);
3) endophallus with long, thin-walled tubular sclerites (Fig. 67);
4) elytral interspace 3 prominent at extreme base.
The male genitalia of A. diamantinaensis are very similar to those of A. rosadonetoi (Figs. 67,
68), the male of which has a well-developed pygidial sulcus. No characters were found that
distinguish the females of the two species.
Description. — Male. Length: 1. 9-2.4 mm (x=2.2, n=15). Width'. 1.0-1. 2 mm (x=l.l, n=15). Head : eyes
separated by distance ca. 0.8 x width of rostrum at base. Rostrum : length 1.3-1. 8 x (x= 1.4, n= 15) pronotal length; most
strongly curved over antennal insertions; proximal portion deeply sulcate, strongly carinate; distal portion finely, sparsely
punctate, length 24-36% (x = 30, n = 15) of total rostral length. Thorax : pronotum with elongate, narrow, aenescent scales
and middorsal vitta of broader, whitish scales. Pygidium : subtruncate apically. Legs: profemur ca. 1.3 x stouter than
metafemur, without distal tooth; protibia with large, acute preapical tooth; metatibial uncus minute, acute.
Female. Length: 2.0-2. 5 mm (x = 2.3, n=15). Width: 1.0-1. 3 mm (x— 1.1, n=15). Rostrum: short, stout, length
1.3-1. 7 x (x=1.5, n=15) pronotal length; broadly, evenly curved; proximal portion deeply rugose, vestiture limited to
extreme base; distal portion smooth, length 25-42% (x = 32, n = 15) of total rostral length. Pygidium (Fig. 46): with broad,
subquadrate, apicodorsal prominence. Abdomen: sternum 5 with posterior margin slightly produced medially.
Anthomorphus Weise
341
Plant Associations. — Label data indicate that A. diamantinaensis has the following host
plants of the family Malpighiaceae:
Banisteriopsis maritiana (Adr. Juss.) Cuatr. (1 specimen, Lagoa Santa, Minas Gerais,
Brazil).
Byrsonima coccolobifolia H.B.K. (1 specimen, Cerrado de Usina, Botucatu, Sao Paulo,
Brazil)
Byrsonima intermedia (7 specimens, Fazenda Campininha, Mun. Mogi-Guacu, Sao
Paulo, Brazil)
Heteropterys umbellata Adr. Juss. (7 specimens, reared from flower buds, Belo
Horizonte and Lagoa Santa, Minas Gerais, Brazil).
Distribution. — This species is known only from the type series from southern Brazil.
Specific Epithet. — The name of this species is based on the name of its type locality.
Anthonomus rosadonetoi, new species
Type Series. — Holotype : Brazil, male [Rio de Janeiro/ Guanabara BRAZIL] [X-63] (MZSP). Paratype: Brazil,
1 female [Rio de Janeiro/ Guanabara BRAZIL] [X-63] (MZSP).
Recognition. — This species is distinguished by the following combination of characters:
1) male with broad, deep, setose pygidial sulcus;
2) female pygidium not sulcate;
3) aedeagus abruptly constricted subapically, with slight apical projection (Fig. 68);
4) endophallus with long, thin-walled tubular sclerites (Fig. 68);
5) elytral interspace 3 prominent at extreme base.
The only character distinguishing this species from A. diamantinaensis, aside from slight
differences in the aedeagus (Figs. 67, 68), is the sulcate male pygidium. The female of A.
rosadonetoi is indistinguishable from that of A. diamantinaensis.
Description. — Male. Length: 2.2 mm (n=l). Width : 1.1 mm (n=l). Head : eyes separated by distance ca. 0.7 x
width of rostrum at base. Rostrum : length 1.4 x (n=l) pronotal length; most strongly curved over antennal insertions;
proximal portion strongly carinate, with broad, densely punctulate, shallow sulci; distal portion rugulose, length 26%
(n=l) of total rostral length. Thorax : pronotum with elongate, narrow, aenescent scales and middorsal vitta of slightly
broader whitish scales. Pygidium : median sulcus narrow, wider and deeper apically. Legs: profemur ca. 1.3 x stouter than
metafemur, with obsolescent distal tooth; protibia with short, blunt preapical tooth; metatibial uncus minute, acute.
Female. Length: 2.4 mm (n=l). Width: 1.1 mm (n=l). Rostrum: short, stout, length 1.6 x (n=l) pronotal length;
broadly, evenly curved; proximal portion deeply rugose, vestiture limited to extreme base; distal portion smooth, length
33% (n=l) of total rostral length. Pygidium: with broad, subquadrate, apicodorsal prominence. Abdomen: sternum 5 with
posterior margin slightly produced medially.
Plant Association. — Unknown.
Distribution. — This species is known only from the type series from southern Brazil.
Specific Epithet. — This species is named for Germano H. Rosado-Neto of Curitiba, Brazil,
in appreciation of his help and friendship.
Quaest. Ent., 1987, 23 (3)
342
Clark
3 4
Figures 1-4. Habitus, dorsal and lateral views: 1, A. rubidus, male, Haddon Heights, New Jersey; 2, A. rubidus, male.
Kappa, Illinois; 3 and 4, A. bordoni, male, Jaji, Venezuela.
Anthomorphus Weise
343
Figures 5-8. Habitus, dorsal and lateral views: 5, A. profundus, female. Trinity Co., California; 6, A. profundus, male,
Ontario, Canada; 7 and 8, A. galphimiae, male, holotype.
Quaest. Ent., 1987, 23 (3)
344
Clark
Figures 9-12. Habitus, dorsal and lateral views: 9 and 10, A. howdenorum, male, holotype; 11 and 12, A. varians, female,
Ermenonv.
Anthomorphus Weise
345
Figures 13-16. Habitus, dorsal and lateral views: 13 and 14, A. sulcipygus, female, Sao Sebastao das Aguas Claras, Nova
Lima, Minas Gerais, Brazil; 15 and 16, A. stockwelli, male, holotype.
Quaest. Ent., 1987, 23 (3)
Figures 17-20. Habitus, dorsal and lateral views: 17 and 18, A.filicornis, female, holotype; 19 and 20, A. sulcatus, female,
holotype.
Anthomorphus Weise
347
Figures 21-24. Habitus, dorsal and lateral views: 21 and 22, A. pimentai, male, holotype; 23 and 24, A. bechyneorum,
male, holotype.
Quaest. Ent., 1987, 23 (3)
348
Clark
Figures 25-28. Habitus, dorsal and lateral views: 25 and 26, A. haliki , female, paratype; 27 and 28, A. rulfoi, male,
holotype.
Anthomorphus Weise
349
Figures 29-32. Habitus, dorsal and lateral views: 29 and 30, A. abdominalis, female, holotype; 31 and 32, A
diamantinaensis, male, holotype.
Quaest. Ent., 1987,23 (3)
Figures 33-36. Pygidium of male, dorsal view: 33, A. profundus , Pocohantas Co., West Virginia; 34, A. stockwelli,
holotype; 35, A. sulcatus , Sao Sebastao das Aguas Claras, Nova Lima, Minas Gerais, Brazil; 36, A. pimentai, holotype.
Anthomorphus Weise
351
Figure 37. Fifth abdominal sternum of male, ventral view: 37, A.filicornis, 8 km. S Kilometro 88, Bolivar, Venezuela.
Figures 38-46. Pygidium of female, dorsal view: 38, A. rubidus, Tippecanoe Co., Indiana; 39, A. profundus. Trinity Co.,
California; 40, A. galphimiae, paratype; 41, A. sulcipygus, Sao Sebastao das Aguas Claras, Nova Lima, Minas Gerais,
Brazil; 42, A.filicornis, 8 km. S Kilometro 88, Bolivar, Venezuela; 43, A. pimentai, Sao Sebastao das Aguas Claras, Nova
Lima, Minas Gerais, Brazil; 44, A. sulcatus, holotype; 45, A. abdominalis, holotype; 46, A. diamantinaensis,
Diamanthina, Minas Gerais, Brazil.
Quaest. Ent., 1987, 23 (3)
352
Clark
47 48
1
51 52
Figures 47-52. Aedeagus, dorsal view: 47, A. rubidus. Kappa, Illinois; 48, A. bordoni, holotype; 49, A. paraguayanus ,
Nova Teutonia, Santa Catarina, Brazil; 50, A. profundus, Pocohantas Co., West Virginia; 51, A. galphimiae, holotype; 52,
A. howdenorum, holotype.
Anthomorphus Weise
353
57 58
Figures 53-58. Aedeagus, dorsal view: 53, A. varians Ermenonv; 54, A. pinivorax , Europe; 55, A. sulcipygus, Sao
Sebastao das Aguas Claras, Nova Lima, Minas Gerais, Brazil; 56, A. chinculticensis, holotype; 57, A. vanini, holotype; 58,
A. camoiranensis, holotype.
Quaest. Ent., 1987, 23 (3)
354
Clark
Figures 59-64. Aedeagus, dorsal view: 59, A. stockwelli holotype; 60, A. filicornis , 8 km. S Kilometro 88, Bolivar,
Venezuela; 61, A. sulcatus, Sao Sebastao das Aguas Claras, Nova Lima, Minas Gerais, Brazil; 62, A. pimentai, holotype;
63, A. bechyneorum , holotype; 64, A. haliki, holotype.
Anthomorphus Weise
355
Figures 65-68. Aedeagus, dorsal view: 65, A. rulfoi, holotype; 66, A. abdominalis, 5 mi. NW Gamboa, Canal Zone,
Panama; 67, A. diamantinaensis , holotype; 68, A. rosadonetoi , holotype.
Figures 69-70. Spiculum gastrale of male genitalia, ventral view: 69, A. bordoni, holotype; 70, A. vanini, holotype.
Quaest. Ent., 1987,23 (3)
356
Clark
PHYLOGENETIC RELATIONSHIPS
General Considerations
Phylogenetic relationships of the species of Anthomorphus were determined by comparing
the distribution of morphological characters of the adult stage of each of the species.
Twenty-seven of these characters (listed in Table 1) were determined to be apomorphic by
comparison of the species of Anthomorphus with a hypothetical ancestral taxon. This taxon, or
outgroup, was conceptualized after examination of numerous anthonomines, including
members of the grandis, gularis, unipustulatus and venustus groups of the genus Anthonomus
and the Anthonomus subgenus Anthonomorphus (see Clark, 1987 and Clark and Burke, 1985;
1986a, b.). The distribution of apomorphic characters (presence indicated by a score of “1,”
absence by a score of “0”) among the 22 species of Anthomorphus is depicted in Table 2.
The characters were analyzed using the PAUP computer programs for the species consistent
with the distribution of characters determined to be apomorphic, thus minimizing the number
of required hypotheses of homoplasy. The result, compromised by weighting some characters to
give a pattern only partly consistent with that produced by PAUP, is the phylogenetic tree
depicted in Fig. 71 .
The analyses are in three separate sets: 1) analyses of all species of Anthomorphus together;
2) separate analyses of the species assigned to the rubidus group; and 3) separate analyses of
the species assigned to the sulcatus group. The same outgroup (lacking apomorphic characters,
Table 2) was specified in each analysis. The three sets of analyses are discussed separately in
the following paragraphs.
Analyses, set 1 . These analyses provided the rationale for recognizing two species groups, a
monophyletic rubidus group and a paraphyletic sulcatus group. An initial PAUP analysis
considered all of the species of Anthomorphus with a unique set of apomorphic characters (i.e.,
one each of each of pairs of taxa 6 and 15, 9 and 11, 10 and 19, and 13 and 18 in Table 2 was
deleted from the analysis), using the MULPARS option. This analysis produced more than 100
equally parsimonious trees. A strict consensus tree derived from these trees indicated that the
species assigned to the rubidus group, except for A. varians and A. pinivorax, belong to a single
monophyletic group. A subsequent PAUP analysis (also using the MULPARS option) treated
the same species as did the first, except that A. pinivorax (taxon 14) was excluded. The
consensus tree derived from the trees produced by this analysis placed A. varians in the rubidus
group at the position indicated in Fig. 71. The problematical A. pinivorax is depicted in Fig. 71
as the sister group of A. varians on the basis of synapomorphy in characters 1, 2, 7, 24, 26, even
though subsequent PAUP analyses weighting these characters failed to move that species to the
rubidus group.
Set 2, the rubidus group. As constructed by the analyses described as set 1, this group is
justified on the basis of synapomorphous characters 12 and 13 (see Table 1). The species in this
group (taxa 4, 12-14, 16, 18, 21 and 23) were subjected to a PAUP analysis using the
ALLTREES option without weighting. This analysis produced a single tree with the topology
depicted in the upper portion of Fig. 71.
The problems encountered in formulation of this group stem from the fact that A. varians
and A. pinivorax seem to have diverged considerably from the New World stock from which
they must have been derived. The greatest problem is that A. pinivorax does not exhibit the
apomorphic characters (12 and 13) indicating monophyly of the group to which it is assigned.
The problem is compounded by the absence in A. pinivorax of the emargination of sternum 5 of
Anthomorphus Weise
357
3 10 20
12 13
14
4 15
27
•(10)
(10)
(15)
1 2 7 24 26
21
(11 12 13)
22
16
17
(22)
8 11
19
18 23
(10
5 25
(8)
rubidus
bordoni
paraguayanus
profundus
galphimiae
howdenorum
varians
pinivorax
sulcipygus
chinculticensis
vanini
camoiranensis
stockwelli
f ilicornis
sulcatus
pimentai
bechyneoruin
haliki
rulf oi
abdominalis
diamantinaensis
rosadonetoi
XX characters
(XX) homoplasious characters
71
Figure 71. Phylogenetic tree depicting hypothesized relationships of the species of Anthomorphus.
Quaest. Ent., 1987, 23 (3)
358
Clark
the male abdomen with the associated isolated triangular sclerite (apomorphic character 11),
one of the characters considered to be synapomorphic in Anthomorphus. Furthermore, A.
varians is autapomorphic for Anthomorphus in possession of simple tarsal claws. It is possible
that A. varians and A. pinivorax are not sister species and that the apomorphic characters they
share (1, 2, 7, 24 and 26) are due to convergence incidental to adaptation to similar host plants
(both species are unusual among known anthonomines in having hosts in the family Pinaceae).
It seems unlikely, however, that two independent host shifts involving relatively distantly
related species would result in this unusual host association.
Set 3, the sulcatus group. A PAUP analysis was performed on the species assigned to the
sulcatus group (taxa 2, 3, 5-10, 11, 15, 17, 19, 20, deleting one each of each of the redundant
pairs of taxa 6 and 15, 9 and 11 and 10 and 19). This analysis also produced more than 100
equally parsimonious trees. The strict consensus tree derived from these trees was accepted in
part. This tree indicated that A. abdominalis , A. rulfoi, A. bechyneorum , A. haliki and A.
pimentai form a monophyletic group. This grouping was accepted even though there is no
synapomorphy to support it. An analysis of these taxa to the exclusion of all of the others except
the outgroup, using the ALLTREES option, produced three trees, the topology of one of which
is produced as part of Fig. 71. The consensus tree also depicted A. diamantinaensis on a branch
of its own as the sister group of all of the remainder of the sulcatus group. That species,
however, is depicted in Fig. 71 as forming a monophyletic group with A. rosadonetoi on the
strength of shared possession of characters 5 and 25, of which the latter is considered the
strongest evidence of the suggested relationship. This arrangement requires the assumption of
the loss of character 8 in A. diamantinaensis. The consensus tree based on the trees produced in
the initial analysis of all of the species of Anthomorphus , described above as “Set 1,” placed
two of the species assigned to the sulcatus group, A. sulcipygus and A. chinculticensis , in the
position of sister group to the rubidus group. There is no character evidence to support this and
it is not incorporated into Fig. 71. Thus, the sulcatus group stands as an incompletely resolved
paraphyletic group.
Further resolution of the phylogenetic relationships of the species in the sulcatus group
cannot be made without additional characters. At this point it does not seem advisable to
extend the character matrix to include additional adult morphological characters that probably
would only add homoplasies.
ACKNOWLEDGMENTS
Thanks are extended to the individuals and institutions listed in the Materials and Methods
Section for the loan of specimens used in this study. H.R. Burke initially suggested the group as
suitable for revision and assisted in gathering literature and specimens. Host plants of some of
the species (Malpighiaceae) were identified by William R. Anderson, University of Michigan.
Special thanks are extended to R. Parentoni Martins and Helcio R. Pimenta, Universidade
Federal de Minas Gerais, for assistance in collecting and rearing Anthomorphus and other
anthonomines. Some specimens were collected by the author travelling with financial support
from the “Biosystematics and Ecology of the Boll Weevil” research project, U.S. Department of
Agriculture Broad Form Cooperative Agreement #12-14-7001-73 with Auburn University.
H.R. Burke, G.W. Folkerts and M.C. Wooten provided critical review of the manuscript. This
paper is published as Alabama Agriculture Experiment Station Journal Series No. 15-861105.
Anthomorphus Weise
359
REFERENCES CITED
Blatchley, W.S., and C.W. Leng. 1916. Rhynchophora or weevils of North Eastern America,
pp. 1-682. Indianapolis, Indiana.
Burke, H.R. 1962. Studies on the genus Anthonomus in North and Central America
(Coleoptera: Curculionidae). I. Some new and little known species from Mexico.
Southwestern Naturalist 7(3-4): 202-210.
Burke, H.R. 1984. Lectotype designations for species of North American Anthonomus
described by W.G. Dietz, H.C. Fall and J.L. LeConte (Coleoptera: Curculionidae). The
Coleopterists Bulletin 38(3): 257-266.
Champion, G.C. 1903. Biologia Centrali-Americana. Insecta. Coleoptera. Rhynchophora.
Curculionidae. Curculioninae (part), vol. 4, pt. 4, pp. 145-312.
Champion, G.C. 1910. IBID, Curculioninae (concluded) and Calandrinae, vol. 4, pt. 7, pp.
79-221.
Clark, W.E. 1987. Revision of the unipustulatus species group of the genus Anthonomus
Germar (Coleoptera: Curculionidae). Coleopterists Bulletin 41(1): 73-88.
Clark, W.E., and H.R. Burke. 1985. Revision of the venustus species group of the genus
Anthonomus Germar (Coleoptera: Curculionidae). Transactions of the American
Entomological Society 111: 103-170.
Clark, W.E., and H.R. Burke. 1986a. Revision of the gularis species group of the genus
Anthonomus Germar (Coleoptera: Curculionidae). The Coleopterists Bulletin 40(1): 1-26.
Clark, W.E., and H.R. Burke. 1986b. Phylogeny of the species of the Anthonomus subgenus
Anthonomorphus Dietz, with discussion of relationships with Anthonomus grandis
Boheman (Coleoptera: Curculionidae). Journal of the Kansas Entomological Society 59(3):
508-516.
Dieckmann, L. 1968. Revision der westpalaearktischen Anthonomini (Coleoptera:
Curculionidae). Beitrage zur Entomologie 17(3/4): 377-564.
Dietz, W.G. 1891. Revision of the genera and species of Anthonomini inhabiting North
America. Transactions of the American Entomological Society 18: 177-276.
Fabricius, J.C. 1775. Systema entomologiae,... [30] + 832 pp. Flensburgi; Lipsiae.
Fall, H.C. 1913. A brief review of our species of Magdalis, with notes and descriptions of other
North American Rhynchophora. Transactions of the American Entomological Society 39:
23-72.
Hamilton, J. 1885. Remarks on some species of Coleoptera, with supplementary descriptions.
The Canadian Entomologist 17: 103-106.
Hamilton, J. 1895. Catalogue of the Coleoptera of Southwestern Pennsylvania, with notes and
descriptions. Transactions of the American Entomological Society 22: 317-381.
Hatch, M.H. 1971. The beetles of the Pacific Northwest. University of Washington
Publications in Biology, vol. 16, xiv + 662., (Part V: Rhipiceroidea, Sternoxi, Phytophaga,
Rhynchophora, and Lamellicornia.)
Hustache, A. 1939. Curculionides nouveaux de ^Argentine et autres regiones
Sud-Americaines. Anales de la Sociedad Cientifica de Argentina 128: 39-124.
Kirsch, T.F.W. 1874. Beitrage zur Kenntniss der Peruanischen Kaferfauna auf Dr.
Abendroth's Sammlungen basirt. Berliner Entomologische Zeitschrift 18: 385-432. (Drittes
Stuck).
LeConte, J.L. 1876. In LeConte and Horn.
Quaest. Ent., 1987,23 (3)
360
Clark
LeConte, J.L., and G.W. Horn. 1876. The Rhynchophora of America, north of Mexico.
Proceedings of the American Philosophical Society 15(96): i-xvi + 455 pp.
Leng, C.W. 1920. Catalogue of the Coleoptera of America, north of Mexico, x + 470 pp.
Mount Vernon, N.Y.
O'Brien, C.W., and G.J. Wibmer. 1982. Annoted checklist of the weevils (Curculionidae sensu
lato) of North America, Central America, and the West Indies (Coleoptera:
Curculionidae). Memoirs of the American Entomological Institute (34): i-ix, 1-382.
Paykull, G. von. 1792. Monographia curculionum sueciae. Edman, Upsala. [viii] + 151 p.
Pierce, W.D. 1907. On the biologies of the Rhynchophora of North America. Studies from the
Zoological Laboratory, The University of Nebraska, Lincoln. 78: 247-319.
Schwarz, E.A. 1890. Food-plants and food-habits of some North American Coleoptera.
Proceedings of the Entomological Society of Washington 1: 231-234.
Schenkling, S., and G.A.K. Marshall. 1934. Coleopterorum Catalogus, Pars 139,
Curculionidae: Anthonominae, pp. 3-83; Laemosaccinae, pp. 1-8. (vol. 29).
Swofford, D.L. 1985. PAUP: Phylogenetic Analysis Using Parsimony. Users Manual, Illinois
Natural History Survey, Champaign, Illinois.
Silfverberg, H. 1977. Nomenclatoric notes on Curculionidae (Coleoptera). Notulae
Entomologicae 57: 13-14.
Weise, J. 1883. Notizen iiber Riisselkafer. Deutsche Entomologische Zeitschrift 27: 254-256.
Wibmer, G.J., and C.W. O'Brien. 1986. Annoted checklist of the weevils (Curculionidae sensu
lato) of South America (Coleoptera: Curculionidae). Memoirs of the American
Entomological Institute (39): i-xvi, 1-563.
Anthomorphus Weise
361
Table 1. Apomorphic characters analyzed to produce phylogenetic tree in Fig. 71 (see Table 2)
1 Eyes small, round, prominent
2 Rostrum slender, smooth, glabrous from base to tip
3 Elytra with triangular basal macula
4 Sutural elytral interspaces expanded subapically
5 Elytral interspace 3 prominent at extreme base
6 Elytral striae large, deep
7 Scales on elytra sparse, setiform, whitish
8 Pygidium of male sulcate
9 Pygidium of female with sulcus broad, apical, posteriorly undelimited
10 Metasternum and abdominal sterna of male with long setiform scales
1 1 Sternum 5 of male abdomen emarginate, leaving setose, triangular sclerite
isolated from posterior margin
12 Basal portion of spiculum gastrale embracing median lobe, keeled ventrally
13 Aedeagus with midventral, subbasal lobe
14 Aedeagus with apical portion asymmetrical
15 Aedeagus strongly asymmetrical
16 Aedeagus expanded at extreme base
17 Aedeagus with acute apicomedian projection
18 Aedeagus narrowed to base, lateral plates thick
19 Aedeagus with asymmetrical, medially emarginate, apical projection
20 Endophallus with spinose proximal plate with s-shaped component, and
with a more distal spine
21 Endophallus with large distal spine and large serrate proximal spine
22 Endophallus with small distal spines, a larger distal spine, and a ribbed
plate
23 Endophallus without denticles, with one small spine
24 Endophallus unarmed
25 Endophallus with long, thin-walled tubular sclerites
26 Femora inflated
27 Meso- and metafemora of male with long, ventral, setiform scales
Quaest. Ent., 1987, 23 (3)
362
Clark
Table 2. Data matrix for the phylogenetic tree in Fig. 71
Anthomorphus Weise
363
INDEX TO NAMES OF TAXA
(Synonyms in italics)
FAMILY GROUP TAXA
Anthonominae, 318
Betulaceae, 317, 324
Fagaceae, 317, 324
Juglandaceae, 324
Malpighiaceae, 317-318, 328, 331,
334-335,337,341
Myricaceae, 317, 334
Pinaceae, 317, 327
Rosaceae, 317, 324, 327
GENERA AND SUBGENERA
Anthomorphus Weise, 317-320, 329, 356,
358
Banisteriopsis Rob. in Small, 335-336,
340
Byrsonima Rich, ex H.B.K., 334, 336, 340
Corylus L., 324
Cretaegus L., 327
Heteropterys H.B.K., 335, 340
Larix Mill., 327
Paranthonomus Dietz, 317-320
Pseudanthonomus Dietz, 317-318
Tetrapterys Cav., 335
SPECIES AND SUBSPECIES
abdominalis Champion, Anthonomus,
319- 320, 322, 339, 349, 351, 355, 358
abies L., Picea, 329-330
arbutifolia (L.) Ell., Aronia, 327
ater (Marsham), Anthonomus, 319
bechyneorum, new species, Anthonomus,
320- 321, 337-338, 347, 354, 358
bordoni, new species, Anthonomus,
320-321, 323, 325, 342, 352, 355
camoiranensis, new species, Anthonomus,
320-321,332,335,353
canadensis (L.) Medic., Amelanchier, 327
chinculticensis, new species, Anthonomus,
320,331,335,353,358
coccolobifolia H.B.K., Byrsonima, 331,
337, 341
crataegi Walsh, Pseudanthonomus, 323
demissa D. Dietr., Prunus, 324
diamantinaensis, new species,
Anthonomus, 319-321, 340-341, 349,
351,355, 358
filicornis Hustache, Anthonomus, 320,
322,333-334, 346,351,354
galphimiae, new species, Anthonomus,
320-321,323, 328,352
glauca Cav., Galphimia, 328
haliki, new species, Anthonomus,
320-321, 337-338, 348, 354, 358
howdenorum, new species, Anthonomus,
320-321,323,328, 344, 352
humilis Adr. Juss., Tetrapterys, 337
instabilis Faust, Anthonomus, 320
laricina (Du Roi) K. Koch, Larix, 327
lutea Michx., Betula, 324
malifolia (Nees & Mart), Banisteriopsis,
337
maritiana (Adr. Juss.) Cuatr.,
Banisteriopsis, 341
montana Mill., Pinus, 329
nigra Arn., Pinus, 330
oxyclada Adr. (Juss., Banisteriopsis, 337
paraguayanus Hustache, Anthonomus,
320, 322-323, 325-326
perforator (Herbst), Anthonomus, 319
pimentai, new species, Anthonomus, 320,
322, 331, 335-336, 347, 350-351, 354,
358
pinivorax Silfverberg, Anthonomus,
319-323, 329-330, 353, 356, 358
profundus LeConte, Anthonomus,
318- 320, 322-323, 327-328, 343,
350-351
profundus vulpinus Dietz, Anthonomus,
326
pubescens (Paykull), Anthonomus, 319,
330
rosadonetoi, new species, Anthonomus,
319- 320, 322, 341,355,358
rubi (Herbst), Anthonomus, 319
rubidus LeConte, Anthonomus, 319-321,
Quaest. Ent., 1987,23 (3)
364
Clark
323-325,327, 342,351
rulfoi, new species, Anthonomus, 320-321,
339, 348,358
serotina J.F. Ehrh., Prunus, 324
spicata (Cav.) DC, Byrsonima, 334
splendens (SW) DC, Myrica, 334
stipulacea Adr. Juss., Byrsonima, 334
stockwelli, new species, Anthonomus, 320,
322.333.335.345.354
sulcatus Kirsch,- Anthonomus, 320, 322,
331,334,336, 346,354
sulcipygus Champion, Anthonomus,
319-320, 322, 331-332, 335-336, 345,
351,353,358
sylvestris L., Pinus, 329-330
umbellata Adr. Juss., Heteropterys, 337,
341
vanini, new species, Anthonomus, 320,
322.332.335.353.355
varians (Paykull), Anthonomus, 319-321,
323, 329-330, 344, 353, 356, 358
varians Paykull, Curculio, 318, 329
vulpinus Dietz, Anthonomus , 319-320,
327
xanthophylla Adr. Juss., Heteropterys,
335,337
Book Reviews
365
BOOK REVIEW
Mitchell, Andrew W. with forward by Gerald Durrell, 1986. The Enchanted Canopy: A
Journey of Discovery to the Last Unexplored Frontier, the Roofs of the World’s Rainforests.
Macmillan Publishing Company. New York. 225 pages, 99 colour photographs, one black
and white photograph, one text figure, subject index. Price $29.95. U.S.A.
A few months before my first trip to the tropical rainforests, I accepted with great pleasure
a chance to review a book on the subject of rainforest canopies, the inside front cover of which
began: “you are about to enter the earth’s last uncharted frontier”, hopefully referring to
rainforests and not the book itself. Opening the cover, I found an incredible collection of superb
colour photographs, many of them entomological, but portraying among them all conceivable
facets of the rainforest environment. I flipped through the pictures numerous times (and
wondered if the picture at the top of page 99 was in sideways) before settling down to the text,
prefaced by a brief, if uninspired, forward by Gerald Durrell.
Andrew Mitchell is an adventurer, naturalist, and television and film producer, and his book
is a semi-popular treatment of life, both human and otherwise, in the world’s rainforests. It is
not scientific, but it does claim to represent the cutting edge of rainforest knowledge, and is
therefore worthy of close scrutiny. The preface includes a vivid description of daybreak in the
Bornean forest canopy, and the first chapter then presents a summary of the location and status
of the world’s rainforests. The tone is set, with emphasis on conservation, primates, and caving
ropes. Apparently, the best way to reach the canopy is via ropes of the same sort used by
spelunkers when lowering themselves into caves. In a harness which can be moved up and down
on a main rope, biologists can suspend themselves at any level of the canopy, dangling with both
arms free to take pictures, notes, or specimens. This is indeed interesting, but by the end of the
book the wonders of caving ropes have been repeated ad nauseam , along with accounts of
construction of walkways in the canopy. The second chapter is concerned with animal
locomotion in the canopy. It begins with an odd argument that nightmares involving a fall from
a great height, a common theme for most of us, might represent a sort of racial memory from
our arboreal ancestry: one of Mitchell’s erroneous views of human nature and evolution, which
in my opinion form the intellectual core of this book. A discussion follows of the use of regular
routes through the canopy by branch-running mammals, which reminded me of the routes
taken in aspen forests around Edmonton by red squirrels. Throughout the text, naturalist
readers will be reminded of temperate examples of many of the supposedly unique features of
the rainforest canopy. A few inaccuracies also surface in this chapter, notably near the middle
of page 44 where one gets the impression that lorises are not primates.
The third chapter, entitled “The Hanging Gardens”, deals with epiphytic plants and their
associated fauna, and is highlighted by a fascinating discussion of arboreal roots. It is more
interesting than the preceding two chapters, perhaps on account of its dramatic, law-of-the
jungle theme. The next chapter, “A Place to Live”, continues the theme of dramatic
competition among the forest creatures. The most entomologically interesting part of this
chapter is a discussion of passion flower vines and their defences against egg-laying by
heliconiid butterflies. Chapter five, “Tree-top Voices”, points out the difficulty of
communicating in the dense growth of the canopy, and surveys the diversity of animal calls
which have evolved to use the “sound window” between 125-600 Hz, in which calls can
penetrate the forest to maximum advantage. A lengthy discussion of primate calls leads to a
brief discussion of orangutan sexuality, in which Mitchell uncritically cites reports of male
Quaest. Ent., 1987,23 (3)
366
Book Reviews
orangutans raping human women, and being brought to “tribal long-house parties to indulge in
some bizarre activities, so much so that there was for a time a fear that venereal disease might
enter the wild population.” Primatologists I questioned on this topic found it as difficult to
believe as I did, although orangutans are apparently the only non-human primates which
engage in forced copulation. Similar sexual prowess was once mistakenly attributed to male
gorillas (for an historical summary, see “Man and Apes” by Ramona and Desmond Morris,
1966, Hutchinson and Co., Ltd., London, pp. 54-83) and this belief may tell us more about our
perceptions of apes than it does about great ape behaviour. This point aside, the fifth chapter is
well-focussed and interesting, a welcome change for the better. “The Flowering of the Forest”,
the sixth chapter, begins by outlining some of the puzzles surrounding the study of canopy
flowers. How do conspecifics achieve cross-pollination, and some, synchronized blooming, when
they are spread few and far between among other trees? Just as this story is picking up
Mitchell launches into another advertisement for caving ropes, followed by scattered natural
history notes having something or other to do with pollination. The next chapter, “Feasting in
the Tree Crowns”, emphasizes plant dispersal and seed success, not feeding by animals as the
title suggests, and includes numerous fascinating accounts of adaptations of canopy plants and
animals. It wanders a bit (for example into a discussion of nesting in hornbills), misidentifies
the beetle figured on page 171 as a scarab (it is a chrysomelid), and anthropomorphically refers
to “terrified” ants having their brains eaten away by parasites, “sending them mad”.
By the time I reached Chapter Eight (“Killers in the Canopy”) it was clear that Mitchell
views nature as a dramatic stage upon which violent battles are fought; a typical approach for
television nature programmes, and programmers. Nonetheless, I enjoyed the interesting survey
of research on carrion feeding vultures and their keen sense of smell, allowing them to find
carrion hidden beneath the canopy. The last two chapters deal with people and their
relationship with the rainforest. The “Tree People” presents a survey of people who “have not
altogether lost the art of climbing”, and thus begins with an anthropological fallacy, bordering
on racism. Uncritical references to the Tasaday tribe, which is now widely considered an
anthropological hoax, further tarnish Mitchell’s credibility. The entire chapter seems to reflect
his perception of native people as noble savages, in harmony with their surroundings and in
sharp contrast with the ignorance of Western culture. Anthropologists now recognize this belief
as a prime example of Western ignorance. The book ends with a plea for conservation, in a
chapter with two titles; one in the table of contents (“A Future Above the Tree Tops”) and one
in the text (“Above the Future Forest”), both of which sound a bit religious to me. Mitchell
makes a provocative point when he suggests that promotion of nature reserves in third world
countries is simply “a new form of colonialism”, which is true in a strictly political and
sociological sense, notwithstanding good biological reasons for such endeavours. The text then
produces the only two typographic errors I could find (“scaffoldng” on p. 236, and “Mavins” on
p. 247), and a string of endorsements for “The Living Planet”, David Attenborough (“the
world’s most famous naturalist”), Operation Raleigh, and a few other adventurous canopy
projects, all of which involve “arboreal naturalists”, fulfilling their “deeper felt mission”. A
complex metaphor summarizes our plight. Man is cutting off the branch he is sitting on in the
great clock of the rainforest which tells us that time is up, and that we must fix the clock
without knowing how it works.
Reading this book is like taking a guided hike through the rainforest canopy with a good
naturalist. Along the way he points out curious plants and animals, and tells fascinating stories
about them, making the hike much m<!fre informative than it would have been without him.
Book Reviews
367
When we return to the starting point, we have picked up his contagious enthusiasm for the
plight of this environment, and feel a strong need to do something about it. As a former
interpretive naturalist myself, I admire his technique, but I also know how tempting it is to
present an embellished, oversimplified version of biology in order to motivate the people on your
hike. Popularization is not inherently bad, but it carries an intellectual responsibility, which
Mitchell has not entirely heeded.
Why review such a book in the first place, you might ask? Certainly it is unfair for a
practising biologist to pounce on a popular account as if it were a scientific treatise. On the
contrary, I argue that popular science is more important than that. Those of us interested in
evolutionary theory, and in the study of ecology know all too well that many of our colleagues
have received most if not all of their training in these fields from their television sets, and the
magazine rack. Well-educated, intelligent people will read this book, and will find little wrong
with its outlook. It is extremely rare to see evolutionary theory portrayed accurately in science,
let alone in the popular press. By understanding popular conceptions of ecology and evolution,
we can identify sources of some of the misconceptions which arise in classrooms, biological
journals, and politics.
Competition plays a strong role in Mitchell’s book. Survival of the fittest, and the struggle
for existence both imply the overriding importance of competition, and so evolution is seen by
many people as nothing but competition. This makes for good stories. It also reinforces the idea
that all life is struggling, presumably to become more advanced, which means more
mammalian, or more human. In Chapter 4 we find that “amphibians suffer from the fact that
their skins are permeable to water.” Should we feel sorry for them, happily wrapped in our own
water-tight epidermis? In the next chapter, we see that gibbons are on the right track, since
they “have now chosen a family way of life.” The Great Chain of Being still figures
prominently in popular ideas.
Then, there is natural selection, which apparently requires sex and rainforests. One poor
plant, mentioned on page 87, “is denied the genetic diversity that sex would bring, and seems to
be locked into an evolutionary dead end.” On page 218, we find the incredible statement that
“without freedom of choice, natural selection is made sterile and evolution is brought to an
end.” By freedom of choice he means genetic diversity, but these concepts are not identical.
Freedom of choice implies free will, or self determination. Natural selection is never prevented
from choosing, it can only be restricted in the number of choices available at a given time. But
“the heartland of natural selection”, the rainforest, is being destroyed. What is natural
selection, in Mitchell’s view? On page 232, Mitchell reflects: “it is hard to believe that mere
accident, moulded by a length of time beyond the imagination of our short-lived human minds,
could realize such diversity.” A creationist could have written that sentence, and creationists
could quote it in support of their cause, if Mitchell were a more prominent evolutionist.
Mis-quoting might not be necessary. Indeed, “it is hard to believe ...”
Evolutionary theory is not based on accident, it is based on selection and heritable variation.
It is not based on sex, or there would be no parthenogenetic or asexual organisms. It is not
based on rainforests, or there would be a wasteland outside my window in Edmonton right now.
But how are non-biologists to know this, if it is the privileged knowledge of a scientific elite?
The political battle between creationism and evolutionary thought no longer needs biology,
since it is a popular concept of evolution which is under attack, and a lay public which defends
it. I have criticised Mitchell’s presentation of biology, but why should I assume that he is
accountable to biologists? Perhaps this is no longer our concern, and perhaps we cannot change
Quaest. Ent., 1987,23 (3)
368
Book Reviews
either side of public opinion.
I will not end on a critical note. We cannot change the popular view of evolution, and we
cannot save the rainforests from today’s population pressures. Mitchell’s book reflects our grief,
and our collective confusion about the processes which operate in our world. On page 187,
Mitchell states; “entomologists, I find, appear uninterested in most things unless insects are
involved.” Appearances, I hope, have proven themselves deceiving.
John H. Acorn
Department of Entomology
University of Alberta
BOOK NOTICE
BELLES, X. 1985. Sistematica, filogenia y biogeografia de la subfamilia Gibbiinae
(Coleoptera: Ptinidae). Treballs del Museu de Zoologia, Num. 3, 94 pp. Ajuntament de
Barcelona. (Parc de la Ciutadella Apt. de Correus 593, 08003- Barcelona, Spain).
This number, the third to be published in a series that began in 1979, is an excellent
example of modern systematic entomology, treating a taxon that is of interest to applied
entomologists as well as to taxonomists and evolutionary biologists, generally. The taxonomic
section contains excellent line drawings of male genitalia and habitus illustrations of many
species. Dot maps complement statements in the text about geographical range of each species.
The treatments of phylogenetic and biogeographic aspects of the gibbiine ptinids seem to be
very well done. A valuable feature of this publication for anglophones is an English two-page
summary of the Spanish text.
Treballs del Museu de Zoologia is a new series continuing the “Trabajos del Museo de
Zoologia de Barcelona”. An English translation of an editorial preceding the text of Belles
explains that the series is intended for publication of extensive monographic research works on
any zoological group. Authors are invited to write in such a way that the text will be “easily
understood by specialists of any branch of Biology”. Because the editorial was translated into
English, I assume that the journal is likely to accept manuscripts in languages other than
Spanish. The potential importance of this series is thus apparent to authors seeking a place of
publication for first class manuscripts in systematic biology.
G. E. Ball
Book Reviews
369
BOOK REVIEW
D’Abrera, B., 1987. Sphingidae Mundi. E.W. Classey Ltd., Faringdon, Oxon., U.K. ix + 226
pages, inch Appendix, generic index, species index. 97.50 pounds sterling (inclusive)
(approximately CAN $220.00).
This latest contribution from Mr. D’Abrera is a significant piece of work for those of us with
a love of the Sphingidae. This is not a revision of the family; it is designed to enable the reader
to identify hawk moths by comparison with the figures. There are no keys or illustrations of
genitalia.
The family is covered very well in this book. The author estimates the family contains 1050
species in approximately 200 genera; of these, only three genera and 124 species are not
illustrated here (diagnoses are provided for six of these species). From this standpoint, this is
the most complete work available on the Sphingidae.
The quality of the figures for which the author is responsible is excellent; leafing through the
plates is a delight in itself! Species are illustrated life size, the figures are clear, and colours are
accurately represented. Described subspecies are listed in the text, and although most are not
illustrated, a brief diagnosis is usually provided. An attempt has been made to show the known
variation of some species, and the dorsal and ventral surfaces of many species are illustrated. A
general distribution and some biological notes are provided. The author has included figures of
some species he has not seen in an Appendix; the quality of these figures varies, but most are
adequate.
Several taxonomic changes have been proposed: four names are proposed as new synonyms;
four names are resurrected from synonymy; one new subspecies is described; one new name is
proposed; and one lectotype is designated.
My only major complaint with this book is that, for the price, it is not particularly well
organized. The arrangement of figures on the plates may look nice in a collection, but doing this
in a book leaves a great deal of unused space. Considering the bulk of the cost of production
must be from producing the plates, leaving unused space only increases the number of plates,
and thus increases the cost of the book. The quality of the figures and the ability of the reader
to identify the species is not affected by the number or arrangement of specimens on the plate.
Since the author bore the entire cost of production himself, I am surprised a more economical
format was not adopted. Also, for several reasons, I can not see that anything has been gained
by including some figures in an Appendix; the author has not consistently placed all figures for
which he is not responsible into the Appendix (see photographs of Philodia astyanor Boisduval,
page 128, and Temnora kala Darge, page 148, and the painting of Barbourion lemaii (Le
Moult), page 89); the author appears to have forgotten which species are in the Appendix (see
below); and the species illustrated in the Appendix are well separated from section of text
relevant to them. With better organization of the plates, the species figured in the Appendix
could easily have been incorporated into the existing plates, thus reducing the number of plates
required (and the cost of the book).
In addition to the organization of the plates, I found it difficult to cross reference the plates
with the text. Finding the section of text relevant to a given species required scanning for a
specific name on one to three pages. Numbering the plates and adding a number to the species
name of each plate (with a corresponding number in the text) would make it much easier to
associate text and plates.
Quaest. Ent., 1987, 23 (3)
370
Book Reviews
Finally, I noted that the format used for presenting data about the distribution and number
of species in each genus was changed on page 86 for no apparent reason. This is a minor point,
but I prefer the text to be consistent throughout.
The book has several other small defects. First, the checklist of genera has not been
consistently followed in the text. Neococytius Hodges, Ampeloeca R. & J. and Sphingidites
Kernbach are listed in the checklist, but do not appear in the text. ( Ampeloeca is treated as a
synonym of Darapsa Walker; Sphingidites is a fossil.) Chlaenogramma Smith and Isogramma
R. & J. (= Autogramma Jordan) from the checklist are listed in the text as synonyms of
Manduca Hiibner and Ceratomia Harris respectively. Cressonia Grote & Robinson is used in
the checklist and on the plate, but Amorpha Hiibner (the correct name) is used in the text.
Compsogene R. & J. from the checklist is equivalent to Amblypterus Hiibner in the text; the
latter does not appear in the checklist and should be spelled Amplypterus (it is the correct
name for this genus). On other matters in the checklist, Neoclanis Carcasson is listed twice,
and the author of Adhemarius is cited as “Hiibner” in the checklist and “Oiticica” in the text.
Second, there are many typographical errors in the text, and although most are minor, few
pages of text have no such errors. The plates are much better in this regard, but several spelling
errors were noted.
Third, several oversights were noted. The text indicates the following species are illustrated
in the Appendix, but figures were not found in the copy before me: Protaleuron rhodogaster R.
& J., Cephonodes banksi Clark, Cephonodes novebudensis Clark, Macroglossum buruensus
Holland, Macroglossum albolineata Clark, Macroglossum bruini Clark, Macroglossum
haslami Clark, Macroglossum moecki Rlitimeyer, and Macroglossum stevensi Clark. The
following species are figured in the Appendix, but this is not mentioned in the text:
Oxyambulyx immaculata Clark, Platysphinx bituberculatus Darge, Polyptychus ferroseus
Gehlen, Baniwa yavitensis Lichy, Xylophones haxairei Cadiou and Theretra tessmanni
Gehlen. Three species are figured on the plates which are not mentioned in the text: “P.
boisduvali” (p. 67), “7/. sounder si" (p. 115) and “ Platysphinx dorsti” (Appendix).
Polyptychoides grayi assimilis R. & J. is figured on page 69; this is not mentioned in the text
(p. 74), and the character in the wing maculation cited as diagnostic for this subspecies does
not agree with the figure. Leucostrophus hirundo (Gerstaecker) is renamed L. alterhirundo by
the author (p. 168), but L. hirundo is used on the plate. Panacra cingulata Clark from the text
(p. 136) has apparently been called P. angulata on the plate. The figure captions for Daphnis
dohertyi Rothschild (p. 127) and Maasenia heydeni (Saalmiiller) (p. 139) indicate that more
than one subspecies has been described for these species, but none are listed in the text.
Adhemarius gannascus dentoni Clark from the text (p. 49) is treated as a full species on the
plate (the author believes this subspecies is a separate species). In the text, Hemeroplanes
gracilis Jordan (p. 100) also appears in the combination Callionima gracilis Jordan (p. 102).
Finally, “Butler, 1877” is cited as the author of Sphinx perelegans Hy. Edwards (p. 40).
I also object very strenuously to the author’s treatment of “ Cocytius cluentius Cramer, 1775
stat. rev.” (p. 22). Since this is the type species of Neococytius Hodges, what the author has
done, in effect, is synonymize Neococytius and Cocytius. However, he has done this without
even citing the name Neococytius in the text, and the discussion of characters upon which this
decision was based is entirely inadequate to allow the reader to make an informed decision
about the validity of Neococytius. His arbitrary action is scientifically unacceptable and is
likely to cause confusion in the mind of the average reader. Personally, I do not think this
synonymy should be accepted until D’Abrera has provided a much more detailed explanation
Book Reviews
371
for this proposed change.
In spite of these criticisms, this book is an important synopsis of the Sphingidae of the world,
and will be indispensible to those of us who collect hawk moths. Unfortunately, the hefty price
tag may discourage many people from buying this book.
E.R. Fuller
Department of Entomology
University of Alberta
BOOK NOTICE
COX, J. M. 1987. Pseudococcidae (Insecta: Hemiptera). Fauna of New Zealand [number]
11, 230 pages. Science Information Publishing Centre, DSIR, P. O. Box 9741, Wellington,
New Zealand (US$ 29.82).
The 116 species of Pseudococcidae (67 newly recorded and 61 new to science) known to
occur in New Zealand are treated in about 100 pages of text, complemented with 119 pages of
excellent illustrations, showing both habitus and detailed structural features of many species.
Only females are keyed and described, since males are known for very few species. Known host
plant records are listed for each mealy bug species in the text, and the mealy bugs recorded for
each known host plant are listed in an appendix.
This volume is another excellent contribution to the entomological literature, meeting in all
respects the high standards required by the Editorial Advisory Group and the Series Editor of
the “Fauna of New Zealand”.
G. E. Ball
Frontispiece. Tuauru River, Mahina, Tahiti. A typical habitat of larvae of Simulium tahitiense Edwards and S. oviceps
Edwards.
Quaestiones Entomologicae 1987, 23 (3)
A TAXONOMIC ACCOUNT OF THE BLACK FLIES (DIPTERA: SIMULIIDAE) OF THE
SOCIETY ISLANDS - TAHITI, MOOREA AND RAIATEA
Douglas A. Craig
Department of Entomology
University of Alberta
Edmonton, Alberta, T6G 2E3 Quaestiones Entomologicae
Canada 23:372-4291987
ABSTRACT
The taxonomy of black flies of the Society Islands - Tahiti, Moorea and Raiatea is revised.
With the exception of one new species, Simulium anatolicum n. sp., subgenus Hebridosimulium
Grenier and Rageau, all species are assigned to the subgenus Inseliellum Rubtsov, for which a
diagnosis is given. An amended diagnosis is provided for Hebridosimulium. Thirteen species
are recognised in the Society Islands, of which eight (S. admixtum n. sp., S. arlecchinum n. sp.,
S. cataractarum n. sp., S. exasperans n. sp., S. lotii n. sp., S. malardei n. sp., S. mesodontium n.
sp., S. neoviceps n. sp.) are described as new from Tahiti, one (S. opunohuense n. sp.) from
Moorea, and one (S. castaneum n. sp.) from Raiatea. The three previously described species
(S. tahitiense Edw., S. oviceps Edw. and S. cheesmanae Edw.) are fully redescribed. Four
species groups are recognised. The malardei - group contains S. malardei, and S. teruamanga
Craig and Craig from Rarotonga; the opunohuense - group contains S. lotii and S.
opunohuense; the oviceps - group contains S. admixtum, S. arlecchinum, S. castaneum, S.
cataractarum, S. mesodontium, S. neoviceps, S. oviceps and Simulium sp. The tahitiense -
group contains S. exasperans, S. tahitiense and Simulium “IIS”. The other species are left
ungrouped. Keys are provided for known larvae, pupae and adults of the Tahitian Simuliidae.
Brief information is given on habitats.
RESUME
Une revision de la taxonomie des mouches noires des lies Societe - Tahiti, Moorea et Raiatea est presentee. A
Texception d’une nouvelle esp'ece, Simulium anatolicum n. sp., sous-genre Hebridosimulium Grenier et Rageau, toutes les
especes sont regroupees dans le sous-genre Inseliellum Rubtsov. Une diagnose est produite pour le sous-genre Inseliellum,
et une diagnose revisee est presentee pour Hebridosimulium. Treize especes sont reconnues pour les lies Societe: dont huit
nouvelles especes (S. admixtum n. sp., S. arlecchinum n. sp., S. cataractarum n. sp., S. exasperans n. sp., S. lotii n. sp., S.
malardei n. sp., S. mesodontium n. sp., S. neoviceps n. sp.) decrites pour Tahiti, une (S. opunohuense n. sp.) pour Moorea,
et une (S. castaneum n. sp.) pour Raiatea. Les trois especes restantes (S. tahitiense Edw., S. oviceps Edw. et S.
cheesmanae Edw.) sont entierement redecrites. Quatre groupe-especes sont reconnus. Le growpc-malardei contenant S.
malardei et S. teruamanga Craig et Craig, de Rarotonga; le growpe-opunohuense (S. lotii et S. opunohuense); le
groupe- oviceps (S. admixtum, S. arlecchinum, S. castaneum, S. cataractarum, S. mesodontium, S. neoviceps, S. oviceps et
Simulium sp), et la groupe- tahitiense contenant S. exasperans, S. tahitiense et Simulium “IS”. Les especes restantes
demeurent non-groupees. Des clefs d’identification sont presentees pour les larves, nymphes et adultes connus pour
Tahiti. Les habitats sont brievement decrits.
INTRODUCTION
Simuliidae occur on many of the world’s volcanic and continental islands (Crosskey, 1981).
With few exceptions, such as large islands like New Zealand and New Guinea (Smart and
Clifford, 1965; Dumbleton, 1973b; McLea and Lambert, 1983), the number of simuliid species
374
Craig
living on individual islands is low. Even for an ecologically diverse and geologically old island
such as New Caledonia (Holloway, 1979) only two taxa of simuliids are known, despite
moderately intensive collection (Bedo, 1977; 1984).
When simuliids were first reported from Polynesian islands, few taxa were recognized
(Edwards, 1927; 1933; 1935) and the taxonomic situation was not definite as Edwards (1927)
indicated by comments about S. cheesmanae.
While larvae of the Tahitian S. tahitiense and S. oviceps were common and relatively well
described, larvae of S. cheesmanae were unknown. Grenier and Rageau (1960) provided more
extensive descriptions for S. tahitiense and S. oviceps , as well as descriptions of new larval and
pupal material, suggesting that it might be that of S', cheesmanae ; but still a relatively simple
taxonomic situation.
While studying the number of larval instars of S. tahitiense and S. oviceps (Craig, 1975a), I
too, described new larval material, but realized it was probably not that of S. cheesmanae
because of its small size, since S. cheesmanae is the largest Tahitian simuliid. At that time I
also collected larval material for karyotyping. Rothfels (in Craig, 1975a) commented that S.
tahitiense and S. oviceps were remarkably close to each other chromosomally. A reconstructed
phylogeny of the Tahitian Simuliidae then seemed very simple (Craig, 1975b). Rothfels (pers.
comm., 1974) subsequently collected more material himself which included at least one taxon
not related to those then known.
In 1980, 1981 and 1983, 1 collected other species of simuliid in Tahiti which were confirmed
karyologically by Rothfels (in Craig, 1983) and it became apparent that the Rarotongan,
Marquesan and Tahitian species were unique in the Simuliidae by possessing heterogametic
females. The taxonomic situation then became far from simple. As can be seen by this present
work, many of the new species are morphologically distinct in the immature stages, and an
intriguing question is why were they not discovered sooner? In retrospect, two aspects of work
on Tahitian Simuliidae provide the answer. Firstly, both S. tahitiense and S. oviceps larvae
prefer the abundant larger, open rivers of Tahiti, while the newly discovered species are in more
specialized habitats, such as cascades, small heavily-shaded streams, or from the only lake-fed
river on the island. Earlier collections, as judged from locality labels, were simply made from
the large rivers which had easier access and probably provided more satisfying results in terms
of numbers of specimens. The new species sometimes occur in very low densities, and several
are known, unfortunately, only from unique specimens. Secondly, S. tahitiense is
morphologically variable and the limits of the variation were not then well known.
Consequently, S. exasperans , S. lotii and S. malardei had been collected previously by earlier
workers and me, but were not recognized.
One of the major difficulties with Tahitian Simuliidae has been association of life stages.
Larvae of most species (both karyotyped and not) are morphologically quite distinct, with
pupae less so and the adults difficult to distinguish without detailed preparation of their
genitalia. Therefore, great care was taken to ensure that adults reared from pupae were
correctly associated to their larvae. With as many as four species occurring sympatrically, and
with little to distinguish some of the later stages, it is remarkable that the original association of
stages of S. tahitiense and S. oviceps was correct. In large part that was because material taken
over the years was from large populations from preferred habitats of those species. However,
the probability of S. tahitiense being confused with S. exasperans n. sp. existed because that
species is so similar to S. tahitiense and it occurs in the same habitats, at quite high frequency
in many places. Likewise, pupae of S. lotii n. sp. are easily confused with small pupae of S.
Society Islands Black Flies
375
exasperans. This raises a serious problem with collections of pinned adults of Tahitian
simuliids. Many such specimens are severely collapsed because the adults are small and cannot
withstand drying. Therefore many exterior characters cannot be easily seen. Without
associated pupal exuviae, and without detailed preparation of the genitalia, specific
identification of such pinned adults is almost impossible. This particularly applies to S.
tahitiense and S. exasperans. Indeed, in some collections of pinned reared adults, associated
pupal exuviae show that S. exasperans had previously been identified as S. tahitiense.
Consequently the work of correctly identifying present collections of pinned material will be
onerous, if not impossible. This makes reared material valuable, and such is essential for future
collections.
Of considerable importance for correct association were specimens undergoing larval/pupal
moults, which allowed characters of both stages to be definitively associated. A few pupal
exuviae from reared adults retained the larval exuviae and these specimens assumed even more
importance.
Some larval material which is karyologically distinct cannot yet be morphologically
characterized. Rothfels (in Craig, 1983) showed two distinct cytotypes, “IIS” and
“IS-complex”, however, the larval remains from the karyological examination are not adequate
to provide sufficient morphological details to erect species for these distinct cytotypes. Larvae
of “IIS” are presently morphologically indistinguishable from S. tahitiense and those of
“IS-complex” cannot be distinguished from S. cataractarum.
Interpretation of the type status of previously described species has been difficult because
Edwards failed to label his type specimens. It was necessary to infer the types and the status
from data labels on the material in the British Museum (Natural History); the lectotypes
designated here are from series of syntype specimens so determined by R.W. Crosskey
(BMNH). For S. oviceps, no Tahitian data were explicitly given for any specimen and syntype
status was inferred by Crosskey from Mumford and Adamson’s locality data and by back
reference to material originally mistakenly identified as S. bussoni (Edwards, 1927).
In comparison to Northern Hemisphere simuliids, little is known about the biology of the
simuliids from the Society Islands, but Schroder (1985, 1988) has made an important
contribution by studying larval distribution, habitats and feeding behaviour.
MATERIAL
More than 200 adults were reared from pupae for this study and a similar number of pinned
adults from various institutions were also examined. Approximately 250 larvae were identified
to species, but many hundreds of larvae of the commoner species were available.
Codens for institutions where material is deposited:
BMNH Department of Entomology, British Museum (Natural History), Cromwell
Road, London, SW7 5BD England.
BPBM Bernice P. Bishop Museum, P. B. 19000- A, Honolulu, Hawaii, 96819.
U.S. A.
CNCI Canadian National Collection of Insects, Arachnids and Nematodes,
Biosystematics Research Centre, Ottawa, Canada. K1 A 0C6.
DAC Personal collection. D. A. Craig, Department of Entomology, University of
Alberta, Edmonton, Canada. T6G 2E3.
Quaest. Ent., 1987, 23 (3)
376
Craig
DSIR New Zealand Arthropod Collection, Entomology Division, DSIR, Private
Bag, Auckland, New Zealand.
IP Institut Pasteur, 25 et 28, Rue du Docteur-Roux, 75724 Paris, France.
MNHP Department d’Entomologie, Museum National d’Histoire Naturelle, 45
Rue de Buffon, 75005, Paris, France.
UASM Strickland Museum, Department of Entomology, University of Alberta,
Edmonton, Canada. T6G 2E3.
USNM United States National Museum, NHB 168, Washington, D. C. 20560,
U. S. A.
METHODS
Collection and Treatment of Specimens
When possible, two collections of larvae were made, one into Carnoy’s fixative for eventual
karyological examination, the other into 90% ethanol to extrude the anal papillae. For scarce
material, fixation in Carnoy’s took precedence.
Adults were reared from pupae placed in tubes with damp filter paper. After emergence,
adults were allowed to mature before preservation in 90% ethanol along with their pupal
exuviae. Most of this material was critical point dried. Although colours are slightly muted, this
is a preferred technique for it results in minimal collapsing. Most museum collections of pinned
specimens of Polynesian simuliid adults, which were allowed to dry naturally, collapsed and are
of little taxonomic use.
Larvae and pharate pupae for examination by light microscopy were dealt with following
Currie (1986). Cleared material was placed in glycerine-jelly as a temporary mountant during
examination. Material for scanning electron microscopy was prepared according to Craig and
Borkent (1980).
Notes about Structural Features
Determination of homologous hypostomal teeth is not without difficulty when all the teeth
are similar in shape and length as in S. oviceps, or when only the median tooth is differentiated
as in S. lotii. Following Currie (1986), I distinguish between apical teeth formed from the
dorsal wall of the hypostoma and the lateral serrations formed from the ventral one. For most
hypostoma a distinct line marks the two structures (Figs. 36, 37, 39, 40). A prominent apical
tooth in a lateral position is termed “lateral tooth - 1 tth”, e.g., as in S. malardei. Such a
decision results in three lateral teeth and two paralateral teeth per side (Fig. 40).
Of particular importance taxonomically is presence or absence of thoracic tubercles on the
pupae. When only late last instar larval material is available, with care, some of the pharate
pupal cuticle can be dissected out with the pupal gills. Tubercles can then clearly be seen with
phase microscopy (Fig. 73). In many older preserved specimens, larval cuticle has separated
from the underlying pharate pupa and is relatively easy to remove. The pharate pupal carcass
can then be examined with a scanning electron microscope (Fig. 72).
Adult Tahitian Simuliidae are relatively difficult to identify and normally require detailed
preparation of their genitalia.
Water Characteristics
To determine amount of suspended material larger than 0.45 n m in the water (Lacey and
Lacey, 1983), 500-1000 ml of water were filtered through previously weighed Millipore®, Type
Society Islands Black Flies
377
tjl* jo' 151*25' I5I*20' 149* 55' U9* 50' 149* 4j'
Society Islands
Q
tee
|slo°
Mooroa
Tahiti
Mo
Windward Islands
149* JS* 149" JO" 149* 25* 149*20' 149* IS' 149* »'
Map 1. Localities and distribution of Black Flies in the Society Islands. 1 = Simulium admixtum n. sp.; 2 = 5.
anatolicum n. sp.; 3 = 5. arlecchinum n. sp.; 4 = 5. castaneum n. sp.; 5 = 5. cataractarum n. sp.; 6 = 5. cheesmanae
Edwards; 7= 5. exasperans n. sp.; 8 = 5. lotii n. sp.; 9 = 5. malardei n. sp.; 10 = 5. mesodontium n. sp.; 11=5'.
neoviceps n. sp.; 12 = 5. opunohuense n. sp.; 13=5. tahitiense Edwards; 14 = 5. oviceps Edwards; 15 = Simulium
“IIS”; 16 = Simulium sp.
Quaest. Ent., 1987, 23 (3)
378
Craig
HA filters. These were then dried and reweighed. The pH of water was determined with a
portable, Fisher, Digital pH meter, Model 609.
Localities and Locality Data
Polynesian place names are used for most localities. Where alternative spellings are
available, both are given initially. Localities are listed in order from Papeete clockwise (i.e.,
north, east; Map 1) with the district given first and italicized. Tahiti is formed from two extinct
volcanic centres, with the larger western land mass termed “Tahiti-nui”, the smaller,
“Tahiti-iti”. Localities on Tahiti-iti are so indicated. Names, longitudes and latitudes were
taken from the maps - “Edition Provisoire 1958, 1:40,000, Ministere des Travaux publics et des
Transports, Institut Geographique National, France” and “Carte Touristique, 1:100,000,
Edition 3, 1977, Institut Geographique National, France”. Where distances from the coast, or
other definitive statement is made on older collections, longitude and latitude was determined
for those localities. Data for a given locality are listed chronologically. Where detailed localities
are not given on labels, these are listed first under “Tahiti”. The station numbers ( e.g Stat.
#3) given on some labels refer to data in the author’s field notes.
Species Criteria
Possession of a unique character state, or unique combination of character states, was taken
as an indication of reproductive isolation and specific status. In most instances, morphological
specific status agrees with cytospecies determined by Rothfels (in Craig, 1983). Choice of types
for three species was dictated by existence of syntypes. Otherwise, the life stage most likely to
be encountered and most easily identified was chosen. Larvae were the preferred stage for types
because of relative ease of identification. Whenever possible, karyotyped larval material
(Rothfels in Craig, 1983) has been deposited in the institution holding the type.
Data from labels of type specimens are cited in the manner following O’Hara (1983). Labels
are listed from the top down, with each label enclosed in quotation marks; lines of text within a
label are indicated by a slash mark. Repositories for all types and material examined are given
in parentheses.
Species Groups
Species which appear to be morphologically closely related have been assigned to species
groups. At present these are based mainly on pupal and larval characteristics. Some species,
such as S. cheesmanae for which only adults are known, are not placed in any group.
Figures
In Figures 26-35 of larval head capsules, rays of labral fans have been omitted for clarity, as
have the hypostomal sensilla in Figures 36-44.
TAXONOMIC TREATMENT
Key to Female Adults of Tahitian Simuliidae
Use of this key requires preparation of the adults’ genitalia. Females of S. admixtum, S.
arlecchinum, S. malar dei, S. mesodontium and S. neoviceps are unknown.
1
Large-sized species: body length 2. 5-3. 8 mm; antennae and legs distinctly
yellow
5.
Society Islands Black Flies
379
V Moderate to smaller-sized species: body length 1. 2-2.1 mm; antennae
yellow basally; legs dark, or with yellow/pale pattern 2
2 (T) Body: blackish-brown; length 1.2-1. 6 mm 3
2' Body: brownish; length 1.5-1. 9 mm 4
3 (2) Body: black. Genitalia: cerci and anal lobe small; lateral sclerite of genital
fork simple; sternite VIII with median pigmented region ovoid (Fig. 6) .
S. lotii n. sp., p. 388
y Body: brownish-black. Genitalia: cerci and anal lobe large; lateral sclerite
of genital fork with accessory arm; sternite VIII with median pigmented
region broad and diffuse (Fig. 8) S. oviceps Edwards, p. 405
4 (20 Genitalia: spermatheca without reticulate pattern; lateral sclerite of genital
fork with anterior projection sharply rounded (Figs. 5,9) 6
4' Genitalia: spermatheca with slight reticulate pattern; lateral sclerite of
genital fork with anterior projection blunt (Fig. 3)
S. cataractarum n. sp., p. 400
5 (1) Calcipala: expanded enormously. Anepisternal membrane: haired
(Hebridosimulium). Genitalia: cerci narrow basally and apically;
hypogynial valves locked basally to anal lobes; sternite VIII with median
pigmented region deeply concave (Fig. 1) S. anatolicum n. sp., p. 382
5' Calcipala: not so expanded. Anepisternal membrane: bare ( Inseliellum ).
Genitalia: cerci broadly rounded apically; hypogynial valves free of anal
lobe bases; sternite VIII with pigmented region diffuse, subrectangular
medially (Fig. 4) S. cheesmanae Edwards, p. 385
6 (4) Sternite VIII: with median pigmented region small; genital fork stem
broader; anal lobes bare (Fig. 5) S. exasperans n. sp., p. 392
6' Sternite VIII: with median pigmented region distinct, extended laterally;
genital fork stem slim; anal lobes hairy (Fig. 9)
S. tahitiense Edwards, p. 394
Key to Male Adults of Tahitian Simuliidae
Use of this key requires preparation of the adults’ genitalia. Males of S. admixtum, S.
arlecchinum, S. mesodontium and S. neoviceps are unknown.
1 Large-sized species; body length 3.0 mm (Fig. 11)
S. cheesmanae Edwards p. 385
Y Moderate to small-sized species; body length less than 2.5 mm 2
2 (1') Gonocoxa: median surface straight to slightly concave, no medial
projection (Figs. 10-16) 3
2' Gonocoxa: median surface convex, with medial projection (Fig. 17)
Simulium sp. p. 408
3 (2) Small-sized species; body length less than 1.5 mm 4
y Moderate-sized species; body length between 1.6-2. 5 mm 5
4 (3) Ventral plate: heart-shaped; bare; anterior arms subparallel to curving
medially; anteriomedian notch indistinct (Fig. 14)
S. malardei n. sp., p. 387
4' Ventral plate: broad; posteromedian region hairy; anterior arms diverging;
anterior median notch rounded, distinct (Fig. 15)
Quaest. Ent., 1987, 23 (3)
380
Craig
S. oviceps Edwards, p. 405
5 (3') Ventral plate: broadly rounded anteromedially; median notch broad (Figs.
10, 12, 16)
5' Ventral plate: protruded anteriorly; median notch deeply incised (Fig. 13)
S. lotii n. sp., p. 388
6 (5) Ventral plate: anterolateral arms flanged distally (Figs. 10, 12)
6 ' Ventral plate: anterolateral arms smooth distally (Fig. 16)
S. tahitiense Edwards, p. 394
7 (6) Ventral plate: posterolateral margins concave distally, cone-shaped, hairy
medially S. exasperans n. sp., p. 392
1' Ventral plate: posterolateral margins slightly concave, posterior apex
broad, hairy posteriorly S. cataractarum n. sp., p. 400
Key to Pupae of Tahitian Simuliidae
Pupae of S. admixtum, S. cheesmanae and S. neoviceps are unknown.
1 Thoracic tubercles: present (Fig. 18); pupal cocoon: either boot- or
slipper-shaped
V Thoracic tubercles: absent (Fig. 19); pupal cocoon: slipper-shaped
2 (1) Gill filaments: shorter than thorax; postscutellum not cone-shaped in
lateral view (Fig. 18); cocoon boot-shaped S. anatolicum n. sp., p. 382
2' Gill filaments: as long or longer than thorax; postscutellum cone-shaped in
lateral view (Figs. 20, 21); cocoon slipper-shaped
3 (F) Dorsal gill filament: shorter than other filaments . . S. lotii n. sp., (p. 388)
and S. cataractarum n. sp., (p. 400)
3' Dorsal gill filament: as long as other filaments v
S. exasperans n. sp., p. 392 and S. malardei n. sp., p. 387
4 (2') Thoracic tubercles: rounded; dorsal gill filament longer than thorax,
sometimes reflexed posteriorly (Fig. 21) . ... S. tahitiense Edwards, p. 394
4' Thoracic tubercles: pointed; dorsal gill filament shorter than thorax and
other filaments, rarely long (Fig. 20) S. oviceps Edwards, p. 405
5 (3) Pharate pupal material indicates that S. arlecchinum n. sp.. (p. 398) and S.
mesodontium n. sp., (p. 403) may also have short dorsal gill filaments and
lack thoracic tubercles.
Key to Mature Late Instar Larvae of Tahitian Simuliidae
The larvae of S. cheesmanae and S. anatolicum are unknown.
1 Labral fan: fully developed; head spot pattern positive (Fig. 35). Posterior
abdomen: expanded gradually (Fig. 22), or expanded ventrally (Fig. 23).
Anal sclerite extended laterally
F Labral fan: rays reduced in number or size, or virtually absent (Figs. 32,
34, 66, 67); head spot pattern partially (Figs. 28, 32), or completely
negative (Figs. 29, 34). Posterior abdomen: hemispherical, or abruptly
expanded laterally at segment VI (Fig. 25). Anal sclerite extended
laterally, or almost, or compfetely extended around anal proleg (Fig. 24)
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381
ov/cc/75-group (p. 397) 2
2 (1') Labral fan stem: shorter than antenna. Line of hypostomal teeth convex
(Fig. 43); median tooth rarely prominent (Fig. 41) 3
2' Labral fan stem: longer than antenna. Line of hypostomal teeth concave;
lateral teeth and median tooth prominent (Fig. 38) 4
3 (2) Labral fan: with ca. 12 small rays. Hypostoma: not protruding markedly
anteriorly. Head sensilla: numerous (Fig. 66). Anal sclerite: extended
laterally, or further ventrally 5
3' Labral fan: virtually absent (Fig. 32). Hypostoma: protruding markedly
anteriorly. Head sensilla: not numerous (Fig. 67). Anal sclerite: extended
completely around anal proleg, sometimes fused
S. neoviceps n. sp., p. 403
4 (2') Anal sclerite: extended almost complete around anal proleg (Fig. 24).
Head spot pattern: distinctly positive (Fig. 29)
S. cataractarum n. sp., p. 400
4' Anal sclerite: extended only laterally. Head spot pattern: negative 6
5 (3) Abdominal cuticle: lacking tubercles. Hypostoma: teeth subequal in size
(Fig. 43) S. oviceps Edwards, p. 405
5' Abdominal cuticle: covered with ovoid, greyish tubercles (Fig. 63).
Hypostoma: sublateral teeth, laterad to median tooth, small (Fig. 41) ....
S. mesodontium n. sp., p. 403
6 (4') Head cuticle: corrugated and rugose (Fig. 26). Head spot colour pattern:
8-shaped S. admixtum n. sp., p. 397
6' Head cuticle: not corrugated and rugose. Head spot colour pattern:
H-shaped (Fig. 27) S. arlecchinum n. sp., p. 398
7 (1) Body pale: length less than 4.5 mm. Posterior abdomen: expanded
gradually (Fig. 22). Anal papillae: simple 8
7' Body dark: length longer than 4.5 mm. Posterior abdomen: expanded
abruptly ventrally. Anal papillae: simple or complex (Fig. 23) 9
8 (7) Sublateral hypostomal teeth: three per side (Fig. 40)
S. malardei n. sp., p. 387
8' Sublateral hypostomal teeth: more than three per side (Fig. 39)
S. lotii n. sp., p. 388
9 (7X) Anal papillae: simple S. exasperans n. sp., p. 392
9 ' Anal papillae: complex S. tahitiense Edwards, p. 394
Subgenus Hebridosimulium Grenier and Rageau
Hebridosimulium Grenier and Rageau, 1961a: 96 (as genus). Type species: Simulium jolyi Roubaud, 1906 (by original
designation).
Simulium ( Hebridosimulium ) Grenier and Rageau. Crosskey, 1967: 27.
Diagnosis. — As for Crosskey (1967), but with the following modifications necessitated by
additional material from Fiji and Vanuatu and by present description of a new species from
Polynesia. Adult: with or without distinct “cell” basal to anterior branch of media (MA);
cibarial pump without armature; hypogynial valves and anal lobes coadapted basally;
spermatheca with slight or heavy pattern, with or without clear area around junction of sperm
duct.
Quaest. Ent., 1987,23 (3)
382
Craig
Pupa: dorsal thoracic tubercles rounded, lateral and posterior tubercles pointed; gill with
eight or ten simple elongated filaments; scutellum not pointed dorsally.
Simulium ( Hebridosimulium ) anatolicum n. sp.
(Figs. 1, 18,69)
Types. —
Holotype. Female: pinned; reared, with pupal cocoon and exuviae as subsidiary material;
label data:- “Holotype”, “Simulium (H.)/ anatolicum Craig/ det./ D.A. Craig 1986”, “Tahiti,
Mahaena, Mahape R./ 17° 34’ 00” S. 149° 20’ 33" W./ alt. 700m. 14-vi-1980,/ D.A. &
R.E.G. Craig. Stat. #6.” (MNHP). Head, genitalia and pupal exuviae in vial. Left gill of pupa
as slide mount, same data as adult (MNHP). Small portion of pupal exuviae used for scanning
electron microscopical examination (DAC).
Specific epithet.— The specific epithet is based on the Greek word “anatolios”, meaning
“eastern”, in reference to this new record well to the east of other Hebridosimulium.
Diagnosis. —
Adult: large; yellowish; mandibles small; wing with no basal “cell”, veins Rl, Sc, lacking
hair; spermatheca, pattern tuberculate, unpigmented region at junction of sperm duct.
Pupa: sternite V hooks close-set.
Description. —
Adult female. (Single reared specimen). Body: generally yellowish-brown. Length: 2.5 mm. Head: grayish-brown,
width 0.98 mm, length 0.69 mm. Eyes: yellowish-grey; interocular distance 0.33 width of clypeus; frontal angle, 115
degrees; ommatidia 0.014 mm in diameter, ca. 34 and 45 respectively across and up eye in middle row. Frons: bulging
ventrally over sunken antennal bases, densely covered with silver scale-like hairs. Clypeus: concolorous with ventral region
of frons, slightly longer than wide, densely covered with silver hairs medially; lateral hairs dark, extended beyond bases of
maxillary palpi. Postocciput: densely covered with silver hairs extended to eye margin, scattered long black hairs extend
over posterior margin of eye. Antenna: yellowish; length 0.63 mm. Mouthparts: length, 0.50 head depth; mandibles with
ca. 30 very fine teeth; lacinia with ca. 25 pronounced retrorse teeth; maxillary palpus (distal articles absent from
specimen), basal articles flattened and densely covered with black hairs; sensory vesicle of third article elongated
occupying 0.50 width of article, opening of vesicle 0.66 width of vesicle, ca. 25 bases of sensilla visible. Cibarial pump:
space between proximal arms flat, smooth, 1.3 times as wide as deep. Thorax: rich brown; length 1.4 mm; postpronotum,
contiguous area with scutum creamy, remainder of scutum uniformly brown, moderate vestiture of silver hair, in some
views three darker vittae; scutellum yellowish, with sparse black and golden hairs; postnotum brown medially and
posteriorly, yellow laterally; anepisternal membrane with distinct pale yellow flattened hairs; katepisternum bare;
mesepimeral hair tuft black, sparse. Wing: length 2.9 mm, maximum width 1.3 mm; veins pale yellow, hairs black; stem
vein hair tuft black, covering vein; basal section of vein R with hairs dorsally and ventrally; Rl clear dorsally; Sc clear
dorsally, with hairs ventrally; area basal to vein MA without distinct clear “cell”. Halter: creamy-white. Legs: coxae,
femura and tibiae yellow, distal portion of tibiae brown; fore basitarsus six times longer than wide, brown anteriorly; hind
basitarsus parallel sided, seven times longer than wide; calcipala enormously extended to 0.66 length of 2nd tarsomere; 3rd
tarsomere with two long hairs extended to pretarsal claws. Pretarsal claw: smoothly curved, simple, no basal tooth.
Abdomen: generally black with dense dark scales anteriorly, pale scales posteriorly on each segment; basal fringe of shiny
dark golden hairs extended to abdominal segment II; tergites III— VII lighter in colour than others; sternite VII distinct,
narrower anteriorly; pleural regions with dense silver scales. Genitalia (Fig. 1): complex; cercus, sharply ovoid distally in
ventral view; anal lobes straight sided medially, almost in contact under hypogynial valves, divergent posteriorly almost at
right angles; valves divergent only slightly posteriorly, closest approach before apex, bluntly rounded posteriorly; sternite
VIII with posterolateral edges of valves very thin; bases of valves and anal lobes interlock, with anterolateral edge of anal
lobe overlapping ventrally base of valve; genital fork stem smooth, thin, with slight anterior knob, lateral arms divergent at
right angles, lateral sclerite acutely triangular. Spermatheca (destroyed during preparation for illustration): dark brown;
distinct turberculate pattern; region at duct junction unpigmented.
Adult male. Unknown.
Pupa. Length; 3.6 mm. Gill (Fig. 18): 1.4 mm in length, eight filaments, branching directly out of base; two ventral
filaments branching from short stem, single median Filament directed anterolaterally from base; dorsal branch dividing
into three, two of these divide again, medial filament single. Head and thoracic cuticle (Fig. 69): with rounded black
tubercles, those more lateral and posterior, pointe^-and hooked. Thorax: trichomes trifid. Abdomen: tergite II with spine
combs anteriorly; tergites III and IV with eight recurved hooks posteriorly; last tergite small, with spine field anteriorly;
Society Islands Black Flies
383
caudal sclerite with distinct terminal spines; sternites V and VI each with six hooks. Cocoon: boot-shaped, 4.6 mm long, 1.6
mm wide, 1.9 mm high; length of ventral lip 1.9 mm; opened at 45 degrees to base, lateral and posterodorsal edges well
formed, anteroventral edge rough, with small fenestrae.
Larvae. Unknown.
Bionomics. — The pupa from which the adult was reared came from grass trailing in a
small, rapidly flowing, open stream. Velocity was ca. 80 cm/s and temperature 29° C.
Simulium tahitiense and S. lotii were taken from the same locality.
Distribution. — The subgenus Hebridosimulium was previously known only from Fiji and
Vanuatu ( = New Hebrides), where it is widely distributed. This new record from Tahiti,
represents a range extension of some 3500 km.
Phylogenetic Relationships. — Dumbleton considered the simuliid from Fiji to represent a
species separate from S. jolyi of Vanuatu ( = New Hebrides). Crosskey (1974), on the other
hand, maintained that S. laciniatum and S. jolyi were conspecific subspecies.
Since the morphological character state differences between adults and pupae of the taxa in
Hebridosimulium are of the same nature as those which separate species within Inseliellum
and which are corroborated karyologically (Rothfels in Craig, 1983), it seems logical to
consider the three taxa in Hebridosimulium as species. Further, the great distances between
the respective islands and likely lack of vicariant genetic interchange among their simuliid
faunas for considerable time ( e.g ., maximum six to 13 million years between Fiji and Vanuatu.
Doutch, 1981), adds weight to the probable specific status of these taxa. For the present, I
hypothesize specific status for the Tahitian Hebridosimulium.
The discovery of a single specimen from a subgenus unexpected in Tahiti initially suggested
that it was a contaminant from collections made either in Fiji or Vanuatu. However, S.
anatolicum is morphologically distinct from S. jolyi and S. laciniatum. The pupa of S.
anatolicum has only eight filaments, not ten, indicating that perhaps it is more closely related
to the Fijian S. laciniatum. Further material from Tahiti, and detailed karyological and
morphological examination of extensive collections in my possession from both Fiji and
Vanuatu may help resolve this situation.
Rothfels (pers comm., 1986), commented that S. anatolicum might be the cytospecies he
had found on Tahiti and termed “forbidden” (in Craig, 1983). Apparently “forbidden” and
Hebridosimulium had similar karyological characteristics.
Subgenus Inseliellum Rubtsov
Inseliellum Rubtsov, 1974: 275 (as genus). Type species: Simulium oviceps Edwards 1933, by original designation and
monotypy (see Crosskey, 1987 for discussion).
Simulium (Inseliellum) Rubtsov. Craig and Craig, 1986: 357. Crosskey, 1987: 388.
Rubtsov (1974) did not provide a diagnosis for Inseliellum beyond stating that the
characteristics were as given for S. oviceps by Grenier and Rageau (1960: 734-735). Since
Craig (1975a, b; 1977; 1983), Crosskey (1987), and this work, show fairly conclusively that
Inseliellum is not just restricted to S. oviceps , a more detailed diagnosis is provided.
Diagnosis. —
Adult Female. Cibarial pump: unarmed. Wing: basal section of radius haired. Thorax:
scutum without conspicuous pattern; pleural membrane bare; katepisternum bare. Legs: fore
tarsus slender, fore basitarsus five to seven times as long as its greatest width; calcipala,
extended to 0.50 length of adjacent tarsomere; pedisulcus not prominent; claws with large basal
tooth. Abdomen: covered with pale scales, semishining or dull on last few segments; sternite VII
clearly developed. Genitalia: hypogynial valves rounded or slightly angulate apically in lateral
Quaest. Ent., 1987, 23 (3)
384
Craig
view; anal lobes normal; cerci rounded to angulate; spermatheca with faint pattern, clear area
at junction of sperm duct. Heterogametic.
Description. —
Adult Male. Upper eye: ommatidia not exceptionally enlarged, 12-22 rows. Thorax: scutum without bold pattern.
Legs: hind basitarsus slightly dilated. Genitalia: gonostyles simple, tapered, with single substantial terminal spine; ventral
plate not toothed, broadly subtriangular, with moderately well developed shoulders, basal arms short, directed anteriorly,
rounded or angular; median sclerite simple, rod-shape; parameres long and slender.
Pupa. Gill: shorter than, or as long as, pupa; four to eight fine filaments, dorsal filaments variable, shorter than, or as
long as others. Thorax: tubercles present or absent, rounded or pointed; trichomes simple; scutellum pointed or rounded
dorsally. Abdomen: onchotaxy normal; tergites I and V bare, II— IV with hooks, VI-VIII with spine combs; terminal spines
small to absent. Cocoon: slipper-shaped.
Larva. Head: margins from essentially parallel to strongly convex; antenna either shorter than labral fan stem or just
longer; labral fans normal to strikingly reduced, rays variable, from normal numbers and shape, to reduced in number,
and/or reduced in size to essentially absent; spot pattern ranging from positive to negative; frontoclypeal apotome variable
in shape, from wide posteriorly, to narrow; apotome cuticle variable, ranging from normally smooth to strongly corrugated,
and/or with raised sensillar sockets; sensilla number normal or very numerous; postgenal cleft variable, from broadly
rounded to essentially absent; hypostoma variable, ranging from 12-15 apical teeth, with median and lateral teeth
prominent and three sublateral teeth per side, or four or five sublateral teeth per side and only median tooth prominent, or
no teeth prominent, or median tooth absent; hypostoma produced anteriorly normal amount, to strongly produced, four to
eight hypostomal sensilla, normally lying in rows parallel to hypostomal edge, or tightly grouped; mandibles normal, to
short and robust, serration various, from subequal sides, to one side half length of other, sides concave or flat, apex sharp;
basal sensillum either on base of serration or on mandible. Abdomen: posterodorsal cuticle with small trichoid, bifid, trifid,
or stellate sensilla; cuticle either smooth, tuberculate or with laterally elongate ovoid, low tubercles; general shape either
normal, gradually increased and decreased in size, broadcast at segment VI or VII. or expanded abruptly ventrally at
segment VI and abruptly posteriorly at segment VIII, or expanded abruptly laterally at segment VI and decreased
gradually posteriorly; posteroventral tubercles present or absent. Anal papillae complex with secondary papillae, or one or
all papillae simple. Rectal scales absent. Posterior circlet of hooks with 60-108 rows of 10-17 hooks.
Bionomics. — Eggs laid on rocks (Craig, 1983), or vegetation (Schroder, pers. comm.,
1987). Larvae and pupae found on rocks or vegetation in a great variety of habitats, ranging
from large, fast open rivers to small, densely-shaded streams; in trickles of water and in leaf
packs on upstream surfaces of rocks (Craig and Craig, 1986).
Geographic Distribution. — Polynesia: Huahine, Marquesas, Moorea, Rarotonga, Raiatea,
Tahiti (Craig, 1983; Craig and Craig, 1986; Sechan in Klein et al., 1983). Possibly Micronesia:
Guam, Palau, Truk (Crosskey, 1987).
Phylogenetic Relationships. — Inseliellum morphologically is close to Eusimulium and
Pomeroyellum and is difficult to separate from those taxa. As Grenier and Rageau (1960)
indicated, adult characteristics of Tahitian species of Inseliellum place it very close to
Eusimulium Roubards as defined by Edwards. Edwards (1935) commented that the Tahitian
and Marquesan simuliids formed a distinct subgroup within Eusimulium , but he did not state
his basis for that comment. Under a scheme suggested by Freeman and de Mellion (1953),
Grenier and Rageau ( loc . cit.) suggested that the three then known Tahitian species belonged
to Division A. Group I, of Simulium. They further comment that the Tahitian species cannot
be placed in Eusimulium as defined (Rubtsov, 1956), because of lack of conical ventral
tubercles in the larvae. However, these structures are present, although not as well developed as
in Eusimulium. Edwards (1932) commented that the difference between typical Eusimulium
species and Tahitian species is the latter’s complete absence of hair on the upper surface of the
Rs vein and eight pupal gill filaments. Edwards further noted that in these features they
resembled Enderlein’s Gomphostibia. He was not aware that some of the Marquesan species
have the typical Eusimulium arrangement of four pupal gill filaments (Sechan in Klein et al.,
1983). Rubtsov (1974) states that S. oviceps and S. tahitiense have affinities with
Eusimulium. Craig (1983) briefly considers the similarity of Polynesian Simuliidae to
Pomeroyellum and the possibility that ancestral Polynesian simuliids may have entered the
Society Islands Black Flies
385
Pacific from the West.
Because of the problem of indistinct morphological limits to the subgenera, the main feature
that will probably establish monophyly of Inseliellum is karyological i.e., heterogametic
females. Establishing the sister taxon to Inseliellum for outgroup analysis for cladistic purposes
will probably be difficult.
Simulium ( Inseliellum ) cheesmanae Edwards
(Figs. 4, 11)
Simulium cheesmanae Edwards, 1927: 242. Lectotype female (by present designation), TAHITI. Edwards, 1935: 38.
Smart, 1945: 502.
Simulium (Inseliellum) cheesmanae Edwards. Crosskey, 1987: 388.
Types. —
Lectotype. Female: pinned. By present designation; label data:- “Lectotype”, “Syntype”,
“Society Is. Tahiti./ 8.3.25. E. Cheesman./ BM 1925-464”, “Simulium cheesmanae Edw.
SYNTYPE 9/ det. R.W. Crosskey 1979”, “Simulium (I.)/ cheesmanae /det /D.A. Craig
1986” (BMNH). Missing right antenna. Both wings with damaged anterior veins.
Paralectotypes - designated from syntypes. Female: pinned; head and genitalia cleared; in
vial; same data as for lectotype, but date “5.5.25” (BMNH). Female: cleared; in vial; same
data as for lectotype, but date “10.3.25” (BMNH). Male: cleared; in vial; label data:- “Mataeia
Aug 1928 A Tonnoir” (BMNH).
Diagnosis. —
Adult female: large; antennae and legs yellow.
Description. —
Adult Female. Body: generally dark brown; length 2. 9-3. 8 mm. Head: dark brown, with distinct areas pollinose; width
1.1 mm, depth 0.8 mm. Eyes: interocular distance slightly less than 0.33 width of clypeus; frontal angle, 95 degrees;
ommatidia 0.023 mm in diameter, ca. 27 and 52 respectively, across and up eye in middle row. Vertex: dark brown
ventrally to frons, regions adjacent to eyes silvery-pollinose extended down between antennae; vestiture of sparse golden
hairs. Clypeus: as long as wide; silvery-pollinose. Post-ocular hairs: barely extended anteriorly to eye margin. Antenna:
length 0.71 mm; dark yellow, lighter basally. Mouthparts: length less than 0.33 head depth; mandibles insubstantial, with
only fine apical serrations; lacinia with ca. 26 pronounced, pigmented retrorse teeth; palpus, 0.8 mm long, distal article 0.4
mm long; third article with distal medial angle distinct, sensory vesicle occupying 0.33 width of third article, 0.50 width of
vesicle, ca. 10-14 sensilla sockets barely visible. Cibarial pump: space between proximal arms slightly less than twice as
wide as deep, rounded with slight medial projection. Thorax: dark brown to black; length 1.6-1. 8 mm; postpronotal lobes
distinct with broadly rounded anterolateral margins, slightly pollinose in dorsal view; scutum uniformly very dark brown,
faint vittae in some views; scutellum and postnotum concolourous with scutum; scutellum subshiny, with hairs on posterior
edge directed anteriorly; postnotum prominent, pollinose in some views; pleuron light brown. Wing: length 3. 7-4. 2 mm,
maximum width 1.4-1. 9 mm; basal cross veins dark brown; stem vein yellow, hair tuft black. Halter: yellow. Legs:
generally yellow, fore- and meso-tarsi dark brown; metabasitarsus and distal metatarsomere dark brown to black, distal
region of metatibia darker; pretarsal claw, smoothly curved with moderately sharp apex, prominent basal tooth slightly
longer than 0.50 length of claw. Abdomen: generally dark brown; basal fringe of golden hairs extended to abdominal
segment III; tergites I-V with sparse golden hairs, remainder of tergites slightly pollinose under denser golden hairs;
pleural and sternal regions greyish. Genitalia (Fig. 4): cercus, blunt apically in ventral view, narrow basally; anal lobe
twice as broad as long; hypogynial valves very broad, median edges virtually straight, extended well over anal lobe; genital
fork stem smooth and narrow, slightly wider anteriorly, lateral arms broad and smooth, lateral sclerite triangular, anterior
apex forming 90°; sternite VIII slightly sclerotized for full width with median rectangular region more heavily sclerotized.
Spermatheca: slightly ovoid; heavily pigmented; no reticulate pattern; unpigmented region at junction of sperm duct
normal.
Adult Male (Single Specimen). Body: generally brownish-black; length 3.1 mm. Head: width 1.3 mm, depth 0.9 mm.
Eyes: upper ommatidia orangey-red; 0.068 mm in diameter, 13 and 15 respectively up and across the eye; lower ommatidia
dark brownish-red, 0.025 mm in diameter. Frons: slightly more than 0.33 head width; dark black-brown, slightly pollinose.
Clypeus: dark brown; 0.20 as wide as head. Antenna: length 0.7 mm; scape, pedicel and basal portion of first flagellomere
yellowish, remainder light brown; first flagellomere as long as scape and pedicel; whole antenna slender in appearance.
Mouthparts: very short, 0.16-0.20 as long as head depth; mandibles possibly absent; lacinia without retrorse teeth, apex
hairy; palpus 0.69 mm long, apical article 0.35 mm in length, sensory vesicle in third article 0.33 as wide as article, opening
Quaest. Ent.. 1987,23 (3)
386
Craig
0.33 as wide as vesicle, ca. 10 sensilla sockets visible. Cibarial pump: space between proximal arms deeply U-shaped, with
smooth edge. Thorax: length 1 . 1 mm; postpronotal lobe with vestiture of golden scales; scutum uniformly black, vestiture
as for postpronotal lobe; scutellum and postscutellum concolourous with scutum; scutellum with sparse black hairs,
otherwise bare. Wing: length 2.7 mm, maximum width 1.3 mm. Halter: yellowish. Legs: yellowish-brown; vestiture of
dense yellowish-brown hairs; hind leg with femur 3.0 times longer than greatest width, tibia 3.3 times longer than width;
pretarsal claw with ca. 22 grappling hooks. Abdomen: dark brown with pilose areas; basal fringe of long golden hairs
extended to abdominal segment III. Genitalia (Fig. 11): gonocoxa approximately twice as long as maximum width;
gonostylus ca. 0.33 as wide as long, slightly curved with one blunt terminal spine, slightly longer than wide; ventral plate
extensively haired medially, cone-shaped posteriorly, posterolaterally concave, anterior arms concave anteromedially,
anteromedian notch broad.
Pupa. Unknown.
Larva. Unknown.
Bionomics. — The females of this species are the largest of the Tahitian simuliids. Although
J. L. Gressitt (label data) has taken specimens of S. cheesmanae at higher altitudes in Malaise
traps, adults are also collected at lower altitudes flying around humans, where the simuliids are
very conspicuous because of their colour. The largest specimen ever taken was probing the skin
of a man’s neck. The substantial teeth on the laciniae indicate that the females may be capable
of blood feeding. It is possible that this is the species which gave rise to the intriguing, older
reports of Tahitian simuliids feeding on humans (Edwards, 1927; Cheesman, 1932).
Phylogenetic relationships Although Grenier and Rageau (1960) suggested that the
immature stages of the species named later herein as S. lotii, might be those of S. cheesmanae ,
this is clearly not so since all stages of S. lotii have been associated. Furthermore, individuals of
S. lotii are amongst the smallest of Tahitian simuliids. It is interesting that no S. cheesmanae
adults were obtained from the many pupae reared during this study. Large larvae, distinct from
those of S. tahitiense , the next largest Tahitian simuliid, have not been found. This suggests
that S. cheesmanae either occurs at very low frequencies and the adults are concentrated by
their host-seeking behaviour around man, or they have an unusual habitat yet to be discovered,
or both. The male is assumed to be that of S. cheesmanae because of its association with
females of that species and because of its large size.
Material Examined. — In addition to types, I have seen the following:
Tahiti. 03-viii-25, L.E. Cheesman (1 2, BMNH); 05-viii-25, L.E. Cheesman (1 2, BMNH); 00-i-68, J.C. Hitchcock (10 2,
USNM).
Papeete , Fautaa (sic.) V. alt. 1000', 23-viii-28, A.M. Adamson (1 2, BPBM); Fautaua V. alt. 1500', ll-xi-28, A.M.
Adamson (1 2, BPBM); Mt. Aorai. alt. 4000-5000', 16-ix-34, E.C. Zimmerman (1 2, BPBM); Fare Rau Apa. 00-i-60,
N.L.H. Krauss (1 2, BPBM); Fautaua V. alt. 25 m, 5/1 l-vii-61, J.L. Gressitt (1 2, BPBM); NW. ridge, Mt. Aorai. alt.
1400-1450 m, 10-vii-61, J.L. Gressitt (1 2, 1 3, BPBM); Fautaua R., Bain Loti. 17° 33' 15" S. 149° 33' 00" W.
08-vii-80, D.A. & R.E.G. Craig (1 2, CNCI); Fautaua R„ Papeete. 17° 35' 30" S. 149° 31' 42" W. l-vii-81, D.A. Craig
(1 2, MNHP).
Faaa, Papearii. alt. 600' , 09-xi-28, A.M. Adamson (1 2, BPBM); Tipaerui V ( = V. de la Reine), 17° 33' 20" S. 149°
25’ 45" W. 12-ix-28, A.M. Adamson (1 2, BPBM).
Papenoo , Papenoo V. 17° 36' 22" S. 149° 25' 08" W. 25-X-28, A.M. Adamson (1 2, BPBM); Papenoo V., 13-ii-61,
N.L.H. Krauss (1 2, USNM).
Papara, Papara V. (?=Taharuu V), alt. 750', 21-xii-28, A.M. Adamson (1 2, BPBM); Mumford & Adamson (1 2,
BMNH).
malardei - group
Diagnosis. —
Pupa: thoracic tubercles, present or absent, if present rounded.
Larva : hypostomal sublateral teeth, four or less; paralateral teeth, two or more; posterior
abdominal segments with dorsal tubercles.
This group may eventually include some of the Marquesan simuliids.
Society Islands Black Flies
387
Included taxa. — Simulium teruamanga Craig and Craig (Craig and Craig, 1986);
RARATONGA: Simulium malardei n. sp.; TAHITI.
Simulium (Inseliellum) malardei n. sp.
(Figs. 14, 22, 40,51,59)
Simulium “new”. Craig, 1983: 534.
Types. —
Holotype. Larva: last instar; in alcohol; label data:- “Holotype”, “S. malardei ”, “Tiarei, west of Trou du Souffleur,/
17° 31’ 30" S. 149° 23 ' 32" W./ alt. 10m, 17-ii-87, D A. Craig” (MNHP).
Paratypes. Male: reared; pinned; with pupal exuviae and cocoon as subsidiary material; cleared, in vial; label data:-
“Paratype”, “Simulium (I.)/ malardei Craig/det./ D.A. Craig 1986”, “TAHITI, Tahiti-iti, Tautira, /E. trib. Vaitepiha
R.,/ 17° 46’ 58" S. 149° 10' 39" W./ alt. 50m, 17-ii-83,/ D.A. Craig. Stat. #61” (MNHP). Female: reared; pinned;
with pupal exuviae and cocoon as subsidiary material; label data:- as for male (MNHP). Pupa: label data:- “Paratype”,
“Simulium (I.)/ malardei Craig/ det./ D.A. Craig 1986”, “Tahiti-iti, Tautira, E. trib. Vaitepiha R.,/ 17° 46' 30" S. 149°
10' 20" W. alt. 50m,/ 19-vii-80, D.A. Craig.” (1, MNHP). Larva: as slide mount; label data:- “Paratype”, “Simulium
(I.)/ malardei Craig/ det./ D.A. Craig 1986”, “Tahiti-iti, Plateau de Taravao, Trib. Vaitepahua R., 17° 46' 25" S. 149°
15' 26" W./ alt. 640m, 10-vii-73,/ D.A. Craig” (CNCI). Larva: as slide mount; label data:- “Paratype”, “Simulium (I.)/
malardei Craig/ det./ D.A. Craig 1986”, “Tahiti-iti, Tautira, E. trib. Vaitepiha R.,/ 17° 46' 30" S. 149° 10' 20" W. alt.
50m,/ 19-vii-80, D.A. & R.E.G. Craig. Stat. #12” (BMNH). Larva: last instar; as slide mount; label data:- “Paratype”,
“Simulium (I.)/ malardei Craig/ det./ D.A. Craig 1986”, “Tahiti-iti, Tapuaemaui, Fanaa R.,/ 17° 47' 20" S. 149° 17'
25" W. 1 5-ii- 1 984, P. Schroder.” (BMNH). Larva: last instar; as slide mount; label data:- “Paratype”, “S. malardei”,
“Tahiti-iti, Tautira, E. trib. Vaitepiha R.,/ 17° 46' 58" S. 149° 10' 39" W./ alt. 50m, 02-vii-8 1 ,/ D.A. & R.E.G. Craig”
(MNHP).
Larvae: in alcohol; label data:- “Paratype”, “Simulium (I.)/ malardei Craig/ det./ D.A. Craig 1986”, “Tahiti-iti,
Plateau de Taravao, Trib. Vaitepahua R.,/ 17° 46 ' 25" S. 149° 15 ' 26" W. 10-vii-73,/ alt. 640m, D.A. & R.E.G. Craig”
(9, BMNH; 8, MNHP). Larvae: penultimate instar; label data:- “Paratype”, “Simulium (I.)/ malardei Craig/ det./ D.A.
Craig 1986”, “Tahiti-iti, Tautira, E. trib. Vaitepiha R.,/ 17° 46' 30" S. 149° 10' 20" W./ alt. 50m, 19-vii-80, D.A.
Craig” (3, CNCI; 1 MNHP). Larvae: penultimate; label data:- “Tiarei, immediately west of Trou du Souffleur, 179 31 '
30" S. 149° 23 ’ 32" W. alt. 10m, 17-ii-87,/ D.A.C.” (3, BMNH; 3, BPBM; 3, CNCI; 4, MNHP; 3 UASM; 3, USNM).
Specific epithet. — Because of the probable phylogenetic importance of this species, it is
named in honour of the Institut Territorial de Recherches Medicales “Louis Malarde”,
Papeete, Tahiti. The personnel of this institute and that of ORSTOM, Papeete, provided
assistance without which this study would not have been completed.
Diagnosis. —
Pupa: dorsal gill filaments long; thoracic tubercles absent. Larva: hypostoma with three
sublateral teeth per side only.
Description. —
Adult Female. Unknown.
Adult Male (single reared specimen). Body: generally blackish-brown; length 0.8 mm. Head: width 0.8 mm, depth 0.6
mm. Eyes: upper ommatidia 0.03 mm in diameter, 18 and 22 respectively across and up eye; lower ommatidia, 0.01 mm in
diameter. Clypeus: 0.20 as wide as head. Antenna: length 0.7 mm. Mouthparts: 0.25 length of head depth; mandibles
slender, with apical hairs only; lacinia short with small retrorse teeth; palpus 0.7 mm long, distal article 0.3 mm, sensory
vesicle spherical, occupying only 0.50 of width of third palpal article, opening less than 0.33 width of vesicle, ca. 10
sensillar sockets visible. Thorax: length 0.7 mm; scutum evenly jet black; postpronotal lobe lighter, vestiture of even
silver-golden scales; scutellum broadly cone-shaped with obtuse angle, vestiture of long pale hairs crossing at midline;
postscutellum light brown; pleuron brown; anepisternal membrane pale. Wing: length 1.7 mm, maximum width 0.6 mm;
stem vein hair tuft small, but distinct; basicostal vein with distinct black hairs. Halter: white. Legs: generally brown;
pretarsal claw smooth, basal tooth clear, 0.33 as long as claw, ca. 24 grappling hooks on each claw. Abdomen: generally
dark brown; basal fringe of brown hairs extended to abdominal segment III; tergite II pale medially, light brown laterally,
other tergites evenly brown. Genitalia (Fig. 14): gonocoxa 2.0 times as long as basal width; gonostylus ca. 2.5 times as long
as basal width, terminal spine prominent; ventral plate heart-shaped, bare, anterior arms robust and heavily pigmented,
medial notch shallow.
Pupa. (From above male, and one fully developed pharate pupa.). Length: 2.0 mm. Gill: 1.4 mm in length; branching
pattern typical; eight slender filaments, subequal in length. Thorax: tubercles absent; cuticle straw coloured and shiny.
Abdomen: tergite I bare, with lateral pigment patches; tergite II with small hooks; tergites III and IV with substantial
hooks; tergite V bare; tergites VI-VIII with small hooks in single row; sternite III bare; sternites IV-VII with four equally
Quaest. Ent., 1987,23 (3)
388
Craig
spaced hooks; caudal spines very short.
Larva. Last instar with dark pupal gills; length 5.7 mm. Body: light brownish-grey with creamy intersegmental
regions. Head capsule (Fig. 31): broadest at stemmata, narrowing posteriorly, but not conspicuously so; generally pale in
colour; frontoclypeal apotome broadest posteriorly; head-spots colour pattern positive, but faint; anteromedian,
anterolateral and posteromedian spots fused, posterolateral spot faint; sensilla number normal; cervical sclerites distinct,
but partially fused to postocciput. Stemmata: surrounded by clear area, extended posterodorsally to ecdysial suture.
Antenna: extended well beyond labral fan stem, total length 0.39 mm, distal article 0.12 mm, annulation between basal
and medial articles insubstantial. Labral fans: with 29 rays, ca. 0.73 mm in length; six or seven medial rays shorter and less
robust; microtrichia of median rays 1.1 times as long as ray width, no microtrichial pattern (Fig. 59). Hypostoma (Fig.
40): heavily pigmented; median tooth slightly shorter than lateral teeth; three sublateral teeth per side; lateral teeth
prominent; one to two paralateral teeth; six to seven lateral serrations per side; hypostomal sensilla, five per side in row.
Postgenal cleft: squarish, 1.3 times as wide as deep, rough anteriorly. Postgenal bridge: 1.3 times as long as cleft (Fig. 31).
Mandible: apex (Fig. 51) with 10-11 spinous teeth between apical teeth and mandibular serration; distance between
serration and spinous teeth as long as width of serration base; serration higher than width as base, anterior edge twice as
long as posterior, concave; basal sensillum 0.50-0.33 length of serration, on distinct base. Maxillary palpus: 2.8 times as
long as width at base. Mandibular phragma: extended ventrad to 0.50 depth maxillae base. Abdomen: segments increased
gradual in size to segment V, then to maximum size, but not suddenly (Fig. 22); posterodorsal cuticle with evenly spaced
spine-like tubercles, trichoid and trifid sensilla, 4.5 /xm and 3.1 nm long respectively; terga VI with two anterodorsal
tubercles (Fig. 22); smaller tubercles on anterior segments (absent from earlier instars); posteroventral tubercles very
small. Anal sclerite: with posteroventral arms 2.0 times longer than dorsolateral arms; lateral accessory sclerite consisting
of two groups of semifused sclerites, dorsal group larger, or posteroventral arms extended almost around anal proleg.
Circlet of hooks: 60 rows of hooks, 14 hooks per row. Anal papillae simple (Fig. 22).
Bionomics. — All material known is from vegetation in small, densely shaded streams and
trickles of water. At the Vaitepiha R. site, particulate material in the water was 0.6 mg/L.
Noticeable was that the water was clear even during heavy rain.
Phylogenetic Relationships. — Simulium malardei is the only Tahitian simuliid possessing
six sublateral hypostomal teeth in the larva. This plesiomorphic character state resembles that
of the Marquesas Island simuliids (Sechan in Klein et al., 1983). Tubercles on the
posterodorsal abdomen of the larvae is a character state shared with larvae of the Rarotongan
S. teruamanga (Craig and Craig, 1986).
Material Examined. — In addition to types I have seen the following;
Tiarei , immediately west Trou du Souffleur, 17° 31 ' 30" S. 149° 23' 32" W. alt. 10m., 17-ii-87, D.A. Craig (6 early
instar larvae, DAC).
opunohuense - group
Diagnosis. —
Pupa : thoracic tubercles absent; gill with dorsal filaments shorter than others.
Larva\ generally pale; hypostomal median tooth prominent, others not; postgenal cleft
squarish; anal papillae simple.
Included taxa. — Simulium lotii n. sp. TAHITI; Simulium opunohuense n. sp. MOOREA.
Simulium (Inseliellum) lotii n. sp.
(Figs. 6, 13,30, 39, 50)
Simulium sp. ? cheesmanae Edwards. Grenier and Rageau, 1960: 733; Grenier and Rageau, 1961b: 174.
Simulium sp. Craig, 1975a: 470.
Simulium sp. “light”. Craig, 1983: 534. Schroder, 1985: 5.
Types. — A larva has been chosen as the holotype because definitive association of stages
was difficult. The holotype comes from Bain Loti, where morphologically identical specimens
are karyologically distinct from sympatric species (Rothfels in Craig, 1983).
Holotype. Larva: last instar; slide mount; label data:- “Holotype”, “Simulium (I.)/ lotii
Craig/ Tahiti, Papeete, Fautaua R., Bain Loti,/ 17° 33’ 15” S. 149° 33’ 00" W./ ll-vii-80,
Society Islands Black Flies
389
D A. & R.E.G. Craig.” (MNHP).
Paratypes. The reared adults chosen as paratypes have associated pupal exuviae closely
matching pharate pupal material taken from last instar larvae. However, there is a possibility
that some of these specimens could be of S. exasperans.
Female: pinned; reared; with pupal exuviae and cocoon as subsidiary material; head and
genitalia in vial; label data:- “Paratype”, “Simulium (I.)/ lotii Craig/det ./ D.A. Craig 1986”,
“Tahiti, Papeete, Fautaua R.,/ Bain Loti/ 17° 33' 15" S. 149° 33’ 00" W. / 08-vii-80,/ D.A.
& R.E.G. Craig. Stat. #2” (MNHP). Female: pinned; reared; with pupal exuviae and cocoon
as subsidiary material; genitalia in vial; label data:- “Paratype”, “Simulium (I.)/ lotii
Craig/det./ D.A. Craig 1986”, “Tahiti, Tiarei, Fareteuira R./ Cascades de Faarumai,/ 17°
32 T 10" S. 149° 23’ 48" W./ 21-vii-80,/ D.A. & R.E.G. Craig. Stat. #15” (BMNH). Male:
pinned; reared; with pupal exuviae and cocoon as subsidiary material; label data:- “Paratype”,
“Simulium (I.)/ lotii Craig/ det./ D.A. Craig 1986”, “Tahiti, Mataiea, Vaihiria R./ 17° 41 '
31" S. 149° 25’ 02" W./ alt. 250m. 17-vii-80,/ D.A. & R.E.G. Craig. Stat. #11” (MNHP).
Alcohol material: karyotyped larval remains; label data:- “Paratype”, “Simulium (I.)/ lotii
Craig/ det./ D.A. Craig 1986”, “Tahiti, Papeete, Fautaua R./ Bain Loti,/ 17° 33 ' 15" S. 149°
33’ 00" W./ 8-vii-80,/ D.A. & R.E.G. Craig. Stat. #2” (4, MNHP). Karyotyped larval
remains; label data:- as above, but date “ll-vii-80/ Stat. #5” (3, BMNH; 4, CNCI). Larvae:
label data:- “Paratype”, “Simulium (I.)/ lotii Craig/ det./ D.A. Craig 1986”, “Tahiti, Paea,
Aoua St., 20-vi-73, D.A. Craig.” (5, CNCI). Pupa & larvae: label data:- “Paratype”,
“Simulium (I.)/ lotii Craig/ det./ D.A. Craig 1986”, “Tahiti, Papeete, Fautaua R., Bain Loti,
17° 33 ’ 15" S. 149° 33 ' 00" W. alt. 92m. 15-vii-74, D.A. & R.E.G. Craig” (1 pupa, 3 larvae,
DSIR). Larvae: label data:- “Paratype”, “Simulium (I.)/ lotii Craig/ det./ D.A. Craig 1986”,
“Tahiti, Papeete, Fautaua R., Bain Loti, 1 4-iv-6 1 , J.N. Belkin” (6, USNM). Larva: label
data:- “Paratype”, “Simulium (I.)/ lotii Craig / det./ D.A. Craig 1986”, “Tahiti, Haapape,
Tuauru R.,/ 17° 32’ 10" S. 149° 29' 15" W. alt. 60m. 09-vii-80/ D.A. & R.E.G. Craig. Stat.
#3.” (1,UASM).
Specific epithet. — In reference to “Bain Loti”, a pool in the Fautaua River, where this
species was first collected. This pool is well known in Tahiti because of the famous French
writer and traveller, Pierre Loti. In “Marriage of Loti” (Loti, 1887), “Loti” spent many hours
dallying in a pool (now Bain Loti) of the Fautaua River with his beautiful young bride Rarahu.
Diagnosis.- —
Larvae : small; pale; head spot pattern positive, spots lightly fused.
Description. —
Adult Female. Body: generally black; length 1.6 mm. Head: black; width 0.6 mm, depth 0.4 mm. Eyes: dark grey;
interocular distance 0.50 width of clypeus; frontal angle, 88 degrees; ommatidia 0.013 mm in diameter, ca. 27 and 37
respectively across and up eye in middle row. Vertex and frons: subshiny, black, slightly pilose. Clypeus: 0.25 head width;
concolourous to slightly lighter than frons, slightly pilose with vestiture of sparse hairs. Antenna: length 0.4 mm; scape,
pedicel and basal portion of first flagellomere lighter in colour than remainder. Mouthparts: length 0.50 head depth;
mandibles shorter than labrum, with ca. 20 very fine teeth; lacinia with ca. 16 retrorse teeth; palpus 0.4 mm long, distal
article 0.08 mm long, proximal article concolourous with clypeus, sensory vesicle occupying 0.50 width of third article,
opening 0.50 width of vesicle, ca. 15 sensillar sockets visible. Cibarial pump: space between proximal arms U-shaped, as
wide as deep, smooth. Thorax: length 0.3 mm; postpronotum brown, junction with scutum distinct; scutum uniformly
black-brown, subshiny with vestiture of very sparse silver scales; scutellum pale yellow, vestiture of very sparse hairs,
posterolateral edges slightly concave; postscutellum, concolourous with scutellum. Wing: length 1.7 mm, maximum width
0.7 mm; stem vein hair tuft not prominent. Halter: pale yellow. Legs: forelegs paler than others; pretarsal claw with basal
tooth 0.50 length of claw. Abdomen: generally dark brownish-grey; basal fringe of insubstantial golden hairs extended
back to abdominal segment II; tergites increasing in size posteriorly, subshiny; tergites VII and VIII with long dark hairs.
Genitalia (Fig. 6): cercus, bluntly cone-shaped in ventral view; hypogynial valves divergent smoothly posteriorly, almost in
contact distally, smoothly rounded apically; stem of genital fork long and thin, lateral sclerites triangular with acute
anterior apex; sternite VIII medial pigmented area subrectangular. Spermatheca: slightly ovoid; no pattern; not strongly
Quaest. Ent., 1987, 23 (3)
390
Craig
pigmented; unpigmented area at junction of sperm duct normal.
Adult Male. Body: generally black; length 2.3 mm. Head: width 0.7 mm, depth 0.6 mm. Eyes: upper ommatidia 0.05
mm in diameter, 18 and 20 per row respectively, up and across eye; lower ommatidia 0.03 mm in diameter, 34 and 28 per
row respectively, up and across eye. Clypeus: black; 0.25 as wide as head; vestiture of long black hairs. Antenna: uniformly
dark grey; length 0.45 mm. Mouthparts: 0.66 length of head depth; labrium pale laterally; mandibles with ca. 20 fine
teeth; lacinia with ca. 15 retrorse teeth; palpus, uniformly blackish-brown, 0.38 mm long, sensory vesicle occupying 0.50
width of third palpal article, opening less than 0.50 width of vesicle, ca. 15 sensillar sockets visible. Thorax: length 0.3 mm;
scutum and postpronotum uniformly matt black, anterior vestiture of long golden scales; scutellum lighter than scutum,
vestiture of dense black hairs; postscutellum black; pleuron dark brown. Wing: length 1.6 mm, maximum width 0.8 mm.
Halter: dark grey. Legs: dark; pretarsal claw smoothly curved and slender, ca. 19 grappling hooks on each claw. Abdomen:
black; basal fringe of long black hairs, extended to abdominal segment IV; tergites V-VI with pale areas laterally; sternites
with yellowish-brown mottled area. Genitalia (Fig. 13): gonocoxa 2.3 times as long as basal width; gonostylus ca. 0.50 as
wide as long, curved with one blunt terminal spine; ventral plate extensively haired, broadly rounded posteriorly,
posterolateral edges concave, distinct lateral angle, anterior arms heavily pigmented, anteromedial area domed with deep
medial incision.
Pupa. Length: 2.4 mm. Gill: length 1.6 mm; branching pattern normal, dorsal pair of filaments 0.50 length of
contiguous pair; markedly varied, both same length, or one, or other shorter, length from 0.25-0.66 length of other
contiguous pair, a few with one filament on that branch (i.e., seven filaments in total). Thorax: tubercles absent; cuticle
light brown, shiny. Abdomen: tergite I bare; tergite II with small hooks and trichoid sensilla; tergites III— IV with
substantial hooks; tergite V bare; tergites VI-VIII with small hooks; tergite IX with field of small spines; sternites IV and
V with two pairs of hooks; sternites VI and VII with hooks evenly spaced; caudal spines, small to absent.
Larva. Length: last instar with dark pharate pupal gills, 4. 3-4.9 mm. Body: greyish-brown with pale intersegmental
regions as broad as colour bands. Head capsule (Fig. 30): essentially parallel-sided; generally pale to light brown, variable;
frontoclypeal apotome broadest at posterior, pale anteriorly, light brown posteriorly; spot pattern positive with
anteromedian, anterolateral and posterolateral spots lightly fused; cervical sclerites distinct; sensilla, normal number.
Antenna: longer than labral fan stem, distal article 0.1 mm, proximal articles 0.47 mm; basal article light brown. Labral
fans: with 31-33 rays, ca. 1.1 mm in length, the six medial rays less substantial; microtrichia of medial rays 2.2 times
longer than ray width; microtrichial pattern only on medial rays, with 6-7 small microtrichia interspersed with larger
microtrichia. Hypostoma (Fig. 39): median tooth prominent; other teeth subequal in length, tips in straight line, four
sublateral teeth per side, two paralateral teeth, three or four small lateral serrations and four to six hypostomal sensilla per
side; lateral and paralateral teeth with tines. Postgenal cleft: squarish. Postgenal bridge: three times longer than cleft.
Mandible: apex (Fig. 50) with nine to 10 spinous teeth; area between teeth and serrations as wide as serration base;
serration as high as basal width, anterior edge 1 .5 times longer than posterior edge, edges straight; sensillum small, not on
serration. Maxillary palpus: 2.0 times as long as width at base. Mandibular phragma: extended ventrad to 0.50 depth of
maxilla base. Abdomen: segments I-V increased slightly in size to VI, segment VII markedly larger, segment VIII largest;
posteroventral tubercles almost absent; posterodorsal cuticle with low ovoid tubercles, and trichoid sensilla, 22.5 jam long.
Anal sclerite: posteroventral arms 2.0 times longer than dorsolateral arms; lateral accessory sclerite fused to posteroventral
arm via narrow junction. Circlet of hooks: 92 rows of hooks, 15-16 hooks per row. Anal papillae: simple.
Bionomics. — Simulium lotii is widespread on Tahiti and occurs with S. tahiliense, S.
exasperans and S. oviceps in shaded, larger rivers and streams. There is a tendency for the
larvae to be on vegetation (Schroder, 1985).
Phylogenetic Relationships. — This species was first recognised by Grenier and Rageau
(1960) as Simulium sp. ( = ? cheesmanae). Examination of their material (Craig, 1983),
showed that it was conspecific with material described as Simulium sp. (Craig, 1975a) and
material termed “light” by Craig (1983). The pupae, and female genitalia of material described
here agree with the material described by Grenier and Rageau (1960) and which I have
examined. However, the ventral plate of the male described here is more cone-shaped apically
than that of the male described by Grenier and Rageau ( loc . cit.). This leaves open the
possibility of a misassociation either by Grenier and Rageau, or me, or, the presence of another
species similar to S. lotii.
Material Examined. — In addition to the types, I have seen the following:
Papeete , Faataua (sic) R., Bain Loti, 17° 33 ' 15" S. 149° 33 ’ 00" W. 00-ix-1954, 00-vi-1959, Grenier, P. and J. Rageau
(slide material of pharate male, IP). 1 l-vii-74, D.A. & R.E.G. Craig (larvae in alcohol, BMNH; BPBM; CNCI; DSIR).
Pirae, Pirae R., undated material, Grenier, P. and J. Rageau. (slide material of pharate female, IP).
Tiarei, Fareteuira R., Cascades de Faarumai, 17° 32' 10" S. 149° 23' 48" W. 2 1 -vii-80, D.A. & R.E.G. Craig (1 2, 1 6,
DSIR).
Mataiea, Vaihiria R„ 17° 41’ 31" S. 149° 25' 02" W. alt. 250 m. 17-vii-80, D.A. & R.E.G. Craig (1 9, 1 <3, BMNH; 1 <5,
Society Islands Black Flies
391
CNCI).
Simulium ( Inseliellum j opunohuense n. sp.
(Figs. 7, 33, 42, 54)
Types. —
Holotype. Larva: last instar; as slide mount; label data:- “Holotype”, “Simulium (I.)
opunohuense Craig/ det./ Craig 1986”, “Polynesia, Moorea, /Opunohu V. Marae de Titiroa,
17° 32 ’ 00” S. 149° 49 ' 40” W./26-ii-84, P. Schroder” (MNHP).
Paratypes. Larvae: Penultimate instars; as slide mounts; other earlier instars in alcohol;
label data:- “Paratype”, “Simulium (I.) opunohuense Craig/ det./ Craig 1986”, “Polynesia,
Moorea, Opunohu V.,/ nr. road leading to Pao Pao, 17° 31 ' 42” S. 149° 50' 00” W./26-ii-84,
P. Schroder” (1 slide, 5 larvae, BMNH; 1 slide, 5 larvae, MNHP).
Specific epithet.— Named after the Opunohu Valley, Moorea, where the larvae and
presumably the adult were found.
Diagnosis-
Female adult: jet black; sternite VIII without median pigmented region. Larva: head spot
pattern positive, spots separate; accessory sclerite of anal sclerite absent.
Description-
Adult Female (single specimen, see Phylogenetic Relationships). Body: evenly jet black; length 2.5 mm. Head: width
0.6 mm, depth 0.5 mm. Eyes: interocular distance 0.33 width of clypeus; frontal angle 45 degrees; ommatidia 0.02 mm in
diameter, ca. 26 and 36 respectively across and up eye in middle row. Clypeus: as wide as long, 0.20 as wide as head,
densely pilose, vestiture of sparse golden hairs. Antenna: length 0.4 mm; scape and pedicel dark orangey-brown;
flagellomeres dark. Mouthparts: length 0.33 head depth; mandibles as long as labrum, with ca. 14-16 small sharp teeth;
lacinia as long as labrum, with ca. 24 very sharp retrorse teeth; palpus, 0.6 mm long, distal article 0.2 mm long, sensory
vesicle elongate, occupying 0.50 length and 0.50 width of third article, opening 0.50 width of vesicle, ca. 50+ sensilla
sockets visible. Cibarial pump: space between proximal arms U-shaped, 1.5 times wider than deep, smooth. Thorax: length
0.9 mm; postpronotal lobe slightly pilose; scutum evenly jet black, vestiture of sparse golden hairs, denser posteriorly;
scutellum and post scutellum concolourous with scutum; scutellum with slightly concave sides, vestiture of long black
hairs; postscutellum slightly pilose. Wing: length 2.0 mm, maximum width 0.9 mm; stem vein hair tuft insubstantial.
Halter: dark yellow. Legs: dark with dense vestiture of fine yellow hairs; pretarsal claw very slender, basal tooth longer
than 0.50 length of claw. Abdomen: evenly black; basal fringe of golden hairs extended back to abdominal segment II.
Genitalia (Fig.7): cercus in ventral view bluntly rounded; hypogynial valves diverging slightly then converging posteriorly,
extended 0.50 way along anal lobe; stem of genital fork smooth thin, with angulate anterior apex, dark brown with pigment
continuing posteriorly into fork; lateral sclerites sharply triangular, anterior apex forming acute angle; sternite VIII
pigmented evenly over full width, slightly darker medially. Spermatheca: ovoid; slight reticulate pattern; not heavily
pigmented; clear area at junction of sperm duct substantial .
Adult Male.- Unknown.
Pupa, (based on pharate pupal material from holotype larva). Gill: length 2.1 mm; eight filaments; branching pattern
typical, two dorsal-most filaments equal in length, 0.50 length of other filaments. (Possibly variable; one filament shorter
and one almost absent). Thorax: tubercles probably absent.
Larva. Last instar larva with dark pharate pupal gills; length 2. 8-3. 4 mm. Body: pale grey with narrow intersegmental
regions. Head capsule (Fig. 33): generally pale to light brown; frontoclypeal apotome pale anteriorly, light brown
posterolaterally; spot pattern positive, anteromedian and posteromedian spots fused, occasionally central spots surrounded
by very light brown; cervical sclerites small, but distinct. Stemmata: with posterior light brown area. Head cuticle:
essentially smooth, corrugated laterally; sensilla number normal. Antenna: longer than labral fan stem; distal, proximal
articles and antennal base light brown. Labral fans: with 29 substantial and three insubstantial rays, ca. 0.73 mm. in
length, darker than head; microtrichia of medial rays up to 2.0 times longer than ray width, pattern of seven or eight small
microtrichia between larger ones. Hypostoma, postgenal cleft and bridge (Fig. 33, 42): essentially as for S. tahitiense\ five
hypostomal sensilla per side. Mandible: apex (Fig. 54) with eight spinous teeth; distance between spinous teeth and
mandibular serration smaller than serration base width; serration as high as basal width, sharply pointed and with concave
sides; lengths of anterior and posterior sides equal; sensillum peg-like. Maxillary palpus: 3.0 times as long as width at base.
Mandibular phragma: extended ventrad to 0.33 depth of maxilla base. Abdomen: segments I-IV increased smoothly
laterally to maximum size at segments V and VI; posteroventral tubercles very small; posterodorsal cuticle with ovoid
scales with longitudinal striae, and with trichoid sensillae 6.9 jam long. Anal sclerite: posteroventral arms 1.5 times longer
than dorsolateral arms; accessory sclerite absent. Circlet of hooks: 81 rows of hooks, 14-16 hooks per row. Anal papillae:
Quaest. Ent., 1987,23 (3)
392
Craig
simple.
Bionomics. — - The holotype larva was collected from a small, heavily shaded rocky stream,
the bottom of which was covered with red epilithic algae. The paratype larvae came from
further downstream and were on trailing roots of trees. The water was 5 cm deep with velocity
ca. 50 cm/s (P. Schroder, pers. comm., 1987).
Phylogenetic Relationships. — Larvae of S. opunohuense are morphologically very similar
to those of S. lotii , differing mainly in the more distinct head spot pattern. However, the female
adults of the two species differ in a number of characteristics, such as colour of body and legs,
and shape and sclerotization of parts of the genitalia. Cytological examination will no doubt
show these two taxa to be very closely related. The single adult collected by Puleston, while
probably of S. opunohuense , was not included in the types series because the association to the
immature stages was not definitive. The possibility remains that if this unique female has not
been correctly associated, S. opunohuense may be conspecific with S. lotii and the adult is then
that of an undescribed species.
Material Examined. — In addition to types I have seen the following:
FR. POLYNESIA , Opunohu Bay (word “Bay” crossed out), vii.6.1959, D.E. Puleston (9 pinned; head and genitalia
cleared and in vial, USNM).
tahitiense - group
Diagnosis. —
Pupa: scutellum bluntly pointed dorsally. Larva: body expanded rapidly ventrally at
segment VI, decreased abruptly at segment VIII (Fig. 23); head spot pattern positive; margins
of head capsule not markedly convex (Fig. 35).
Included taxa. — S. tahitiense Edwards, S. exasperans n. sp. and Simulium “IIS”;
TAHITI.
Simulium (Inseliellum) exasperans n. sp.
(Figs. 5,12,19,49)
Types. —
Holotype. Female: pinned; reared; with pupal exuviae and cocoon as subsidiary material;
label data:- “Holotype”, “Simulium (I.)/ exasperans Craig”, “Tahiti, Punaauia, Punaruu R., /
17° 38 ’ 20" S. 149° 35 ' 05" W. /alt. 120m. 07-vii-80, / D.A. Craig. Stat. #1” (MNHP).
Paratypes. Adults: pinned; reared; with exuviae and cocoons as subsidiary material; label
data as for holotype (2 <3, BMNH; 1 <3, BPBM; 1 9, 2 <3, MNHP). Male: pinned; reared; with
exuviae and cocoon as subsidiary material; label data:- “Paratype”, “Simulium (I.)/ exasperans
Craig”, “Tahiti, Papeete, Fautaua R., Bain Loti, /08-vii-80, 17° 33 ’ 15" S. 149° 33 ’ 00" W./
D.A. Craig. Stat. #2” (1 <3, MNHP). Females: pinned; reared; with exuviae and cocoons as
subsidiary material; label data as for Bain Loti, but date as “1 l-vii-80.Stat. #5” (1 9 each,
BMNH; BPBM; CNCI; MNHP). Male: slide mount; label data as for Bain Loti, but date:-
“08-vii- 1 974” (DAC). Female: pinned; reared; with exuviae and cocoon as subsidiary material;
label data:- “Tahiti, Tiarei, Fareteuira R., Cascades de Faarumai,/17° 32’ 10" S. 149° 23'
48" W./ 2 1 -vii-80, / D.A. & R.E.G. Craig. Stat. #15” (1 9, BMNH). Male and female:
pinned; reared; with exuviae and cocoons as subsidiary material; label data:- “Tahiti-iti,
Tautira, Vaitepiha R.,/17° 46’ 30" S. 149° 10’ 21" W./ alt. 50m. 19-vii-80,/ D.A. & R.E.G.
Craig. Stat. #13” (1 9, 1 <3, MNHP). Males and females: pinned; reared; with exuviae and
Society Islands Black Flies
393
cocoons as subsidiary material; label data:- “Tahiti, Mataiea, Vaihiria R.,/ 17° 41 ’ 31" S.
149° 25' 02" W./ alt. 250 m. 17-vii-80, /D.A. & R.E.G. Craig. Stat. #11” (2 9, 1 each
BMNH; BPBM; CNCI; MNHP; 1 9, DSIR). Female: pinned; reared; with exuviae and cocoon
as subsidiary material; label data:- “Tahiti, Mataiea, E. Lac Vaihiria, /17° 40’ 58” S. 149°
24' 51” W./ alt. 430m. 22-vii-80, /D.A. & R.E.G. Craig. Stat. #16” (1 9, MNHP). The
following material was previously identified as S. tahitiense. Female: pinned; reared; with
exuviae and cocoon as subsidiary material; label data:- “Tahiti, Baini (sic) /Loti /April 14,
1961. /J.N. Belkin #35” (1 9, USNM). Female: pinned; reared; with exuviae and cocoon as
subsidiary material; label data:- “Tahiti, Paea, / March 30 1961, /D.G. Basio #26” (1 9,
USNM). Female: pinned; reared; with exuviae and cocoon as subsidiary material; label data:-
“Tahiti, Paea,/ April 2 1961, / J.N. Belkin #28” (1 9, USNM). Pupa: with larval exuviae; label
data:- “Tahiti, Punaruu R., / 1969. B. Hocking” (MNHP). Larva: last instar; label data:-
“Tahiti, Mataiea, Vaihiria R., / 17° 41 ’ 31" S. 149° 25’ 02" W./ alt. 250m. 17-vii-80,/D.A.
& R.E.G. Craig” (BMNH). Pupal and larval material: as slide mounts; two last instar larvae;
label data:- “Tahiti. Paea, Aoua St., /20-vi-73, D.A. Craig” (BMNH; MNHP). Alcohol
material: Pharate adult; label data:- “Papeete, Fautaua R., Bain Loti,/ 17° 33 T 15" S. 149°
33’ 00" W./ 08-vii-80, D.A. & R.E.G. Craig. Stat. #2” (CNCI). Larvae: label data:- “Tahiti,
Paea, Aoua Str., /20-vi-73, D.A. Craig” (larvae, 2 pupae each BMNH; BPBM; MNHP;
larvae, CNCI; DSIR; USNM). Larvae: label data:- “Tahiti, Paea,/ ‘Robinson’s Place’,
/28-iii-61. J. N. Belkin” (5, USNM). Larvae and pupa: label data:- “Paea, Public Garden, /
03-vii-81, D.A. Craig” (6 larvae, BMNH; 5 larvae, 1 pupa, MNHP).
Specific epithet. — Based on the Latin word “exaspero”, meaning “to irritate”, in reference
to the difficulty of associating the stages of this species.
Diagnosis. —
Pupa: thoracic tubercles absent; dorsal gill filaments long.
Larva: tahitiense-Yike, but anal papillae simple.
Recognition. — This species is very difficult to distinguish in all stages from both, S.
tahitiense and S. lotii. Consequently, further work is needed to clarify the limits of the taxa.
Adults of S. tahitiense , without the associated pupal exuviae, are virtually impossible to
separate from S. exasperans. For this reason alone, reared adults are absolutely necessary for
taxonomic studies on Tahitian simuliids.
Description. —
Adult Female. Body: generally blackish-brown; length 1. 5-2.1 mm. Head: very dark brown; width 0.5 mm, depth 0.4
mm. Eyes: dark red; interocular distance 0.50 width of clypeus; frontal angle, 75 degrees; ommatidia 0.02 mm in diameter,
ca. 24 and 26 respectively across and up eye in middle row. Post-ocular hairs black; extended to eye margin. Clypeus: 0.16
mm, lighter than frons, margins pilose, vestiture of sparse black hairs. Antenna: length 0.4 mm; scape, pedicel and anterior
region of first flagellomere yellow, remaining flagellomeres dark brown. Mouthparts: length 0.4 times head depth;
mandibles, as long as labrum, with ca. 21 teeth; lacinia with ca. 16 retrorse teeth; palpus, 0.4 mm long, distal article 0.13
mm long, proximal article concolourous with clypeus, remainder lighter, sensory vesicle occupying 0.50 width of third
article, opening 0.50 width of vesicle, eight to 10 sensillar sockets visible. Cibarial pump: space between proximal arms
flat, smooth, three times as wide as deep. Thorax: length 0.8-0. 9 mm; scutum evenly very dark brown; postpronotal lobes
with posterior margins lighter, sparse, even vestiture of silver-golden scales; scutellum cone-shaped, forming slightly obtuse
angle, lateral margins straight, vestiture of sparse, long golden and black hairs; postscutellum, concolourous with scutum,
pilose in some views; pleuron evenly dark brown. Wing: length 1. 9-2.1 mm, maximum width 0.8-0. 9 mm; stem vein hair
tuft substantial; basicostal vein with dense black hairs. Halter: pale. Legs: as for S. tahitiense. Abdomen: evenly very dark
brown; basal fringe of golden hairs extended back to posterior of abdominal segment II. Abdomen: tergites II— VII mottled
with lighter markings; vestiture of very sparse golden scales, black on tergites VIII and IX; pleural and sternal regions
pilose and greyish. Genitalia (Fig. 5): cercus pointed distally in ventral view; anal lobe curving smoothly medially;
hypogynial valves diverging distally, but converging apically, very blunt, extended just to anal lobes; stem of genital fork
thin, smooth, lateral sclerite with rounded angulate anterior apex and long thin posterior arm; sternite VIII with
pigmented median region narrow posteriorly, expanded anteriorly. Spermatheca: ovoid; no pattern; not heavily pigmented;
!
Quaest. Ent., 1987,23 (3)
394
Craig
clear area at junction of sperm duct normal.
Adult Male. Body: generally blackish-brown; length 1.6-1. 7 mm. Head: width 0.6 mm, depth 0.5 mm. Eyes: upper
ommatidia 0.09 mm in diameter, 16 and 18 per row respectively across and up the eye; lower ommatidia, 0.03 mm in
diameter. Clypeus: 0.20 as wide as head; vestiture of sparse black hairs. Antenna: length 0.4 mm; evenly dark brown.
Mouthparts: 0.33 length of head depth; mandibles very thin, no teeth, only fine apical hairs; lacinia with ca. 23 small
retrorse teeth; palpus 0.4 mm long, distal article 0.2 mm, sensory vesicle spherical, occupying 0.66 of width of third palpal
article, opening less than 0.50 width of vesicle, ca. 10 sensillar sockets visible. Thorax: length 0.6 mm; scutum evenly black;
postpronotal lobes slightly lighter; scutellum lighter, pale posteriorly, vestiture of long black hairs crossing in midline;
postscutellum dark brown, slightly pilose; pleuron brown. Wing: length 1.5-1. 7 mm, maximum width 0.7-0. 8 mm; stem
vein hair tuft small, but distinct. Halter: brown. Legs: as for S. tahitiense. Abdomen: generally blackish- brown; basal
fringe of black hairs extended to abdominal segment II; tergite II pale; III— VII mottled brown; VIII-X dark brown;
tergites III, VI, VII in some views pilose laterally. Genitalia (Fig. 12): gonocoxa 2.0 times as long as basal width;
gonostylus approximately 3.5 times longer than basal width, curved with one blunt terminal spine; ventral plate with
broadly rounded apex posteriorly, concave posterolaterally, anterior arms angulate and strongly pigmented, anteromedial
notch barely present; hairy region narrow in middle.
Pupa. Length 2. 0-2. 5 mm. Gill (Fig. 19): length 0. 9-1.0 mm; branching as for 5. tahitiense. Head and thoracic
cuticle: without tubercles; cuticle straw coloured and shiny. Abdomen: tergites essentially as for S. tahitiense ; tergite I
without tubercles, but with small scales; tergite II without hooks, but with anteriorly directed short hairs; tergites III and
IV hooks slender; those of tergites VI-IX small and in single rows; caudal spines essentially absent; sternites as for S.
tahitiense.
Larva. Virtually indistinguishable from 5". tahitiense. Head capsule: paler than S. tahitiense ; to very pale (similar to S.
lotii). Labral fans: with 29 rays (15s ray robust others finer), ca. 0.45 mm. in length; microtrichia of medial rays up to 1.6
times longer than ray width; no microtrichial pattern. Hypostoma: as for S. tahitiense (Fig. 44). Abdomen: posterior shape
not as pronounced as for S. tahitiense ; ventral tubercles not prominent; posterodorsal cuticle with laterally elongated and
sharply pointed sculpting; trichoid sensilla 16.0 jum. Anal sclerite: with posteroventral arms 2.0 times longer than
dorsolateral arms. Circlet of hooks: as for S. tahitiense. Anal papillae: simple.
Bionomics. — Larvae tend to be found on vegetation in well shaded, larger streams. In some
situations it forms almost the complete population, e.g., Aoua Stream, Paea, otherwise, it is
found with S. lotii, S. oviceps and S. tahitiense.
Phylogenetic Relationships. — It is interesting that no karyological evidence of S.
exasperans was detected by Rothfels {in Craig, 1983). However, this species may be
cytospecies “IIS” which Rothfels {loc. cit .) found only on Tahiti-iti, but S. exasperans is also
found extensively on Tahiti-nui. Association of stages of this species was particularly difficult.
The connection between larval and pupal stages was eventually made via some pupae retaining
their larval exuviae in the cocoon, and by partially ecdysed pharate adults.
Material Examined. — In addition to types, I have seen the following:
Alcohol material.
Papeete , Fautaua R., Bain Loti, 17° 33' 15" S. 149° 33' 00" W. 15-vii-74, (1 pupa, CNCI); l-vii-81, (10 pupae,
BMNH), D.A. & R.E.G. Craig.
Papenoo, Papenoo V., 22-X-28. 17° 36' 22" S. 149° 25' 08" W. A.M. Adamson (2 pupae, BPBM).
Tiarei, Onofea R„ 17° 33 ' 30" S. 149° 23 ' 47" W. alt. 1 10m. 14-vii-80, D.A. & R.E.G. Craig (3 pupae, UASM)
Mataiea, Vaihiria R„ 17° 41 ' 31" S. 149° 25' 02" W. alt. 250m. 17-vii-80, D.A. & R.E.G. Craig (4 pupae, BPBM).
Tautira, (Tahiti-iti), E. trib. Vaitepiha R., 17° 46' 30" S. 149° 15' 20" W. alt. 50m. 14-vii-80 (3 pupae, DSIR),
19-vii-80 (3 pupae, USNM), D.A. & R.E.G. Craig; E. trib. Vaitepiha R., 17° 46' 58" S. 149° 10' 39" W. alt. 50m.
02-vii-81. D.A. Craig (5 larvae, BPBM).
Teahupoo , (Tahiti-iti), Tiirahi R„ 17° 51 ' 05" S. 149° 14' 15" W. 20-vii-80, D.A. & R.E.G. Craig (4 pupae, UASM).
Tauemaui, (Tahiti-iti), Fanaa R., 15-ii-84, P. Schroder (1 pupa, MNHP).
Paea, Public Gardens, 03-vii-81, D.A. Craig (larvae, DAC).
Punaauia , Punaruu R., 17° 38' 20" S. 149° 35’ 05" W. alt. 120m. 27-iv-69, B.M. Hocking (1 pupa, UASM); 7-vi-80,
D.A. & R.E.G. Craie (2 pupae, UASM); 17-vii-80, P. Schroder (5 pupae, MNHP).
Simulium (Inseliellum) tahitiense Edwards
(Figs. 9, 16, 21, 23, 35, 44, 56, 65, 71-73)
Simulium tahitiense Edwards, 1927: 242. Lectotype female (by present designation), TAHITI. Edwards, 1935: 35. Smart,
1945: 242. Grenier and Rageau, 1960: 728. Grenier and Rageau, 1961b: 174. Rubtsov, 1974: 242, 244. Craig, 1975a:
468. Crosskey, 1981: 8. Craig, 1983: 534. Schroder, 1985: 17.
Society Islands Black Flies
395
Simulium (Inseliellum) tahitiense Edwards. Crosskey, 1987: 388.
Types. —
Lectotype. Female: pinned; label data:- “Lectotype”, “Syntype”, “Tahiti. Tautira /9.viii.25
/Miss Cheesman B.M. 1925. 392”, “Simulium tahitiense Edw”, “Simulium tahitiense Edw
/Syntype $,/det. R.W. Crosskey, 1979”, “Simulium (I) tahitiense Edw. / det./ D.A. Craig
1986” (BMNH).
Paralectotype. Female: pinned; label data:- as for lectotype (BMNH).
Diagnosis. —
Pupa: gill filaments long; thoracic tubercles rounded.
Larva: head spots positive; general body colour dark; anal papillae complex.
Recognition. — Larvae of this species are morphologically varied, particularly so in size,
colour and shape of body. Some of the apparent variation may be due to the difficulty of, as yet,
clearly separating the similar stages of S. exasperans.
Description. —
Adult Female. Body: generally dark brown; length 1.5-1. 9 mm. Head: dark brown, slightly pollinose; width 0.71 mm,
depth 0.55 mm. Eyes: interocular distance 0.50 width of clypeus; frontal angle, 90 degrees; ommatidia 0.012 mm in
diameter, ca. 31 and 43 respectively, across and up eye in middle row. Vertex: dark brown. Frons and clypeus pilose; latter
with sparse golden hairs. Clypeus: as long as wide; apex with small medial projection. Antenna: length 0.47 mm; scape,
pedicel and proximal portion of first flagellomere lighter in colour than remainder. Mouthparts: length less than 0.50 head
depth; mandibles as long as labrum, with 22 fine teeth; lacinia with ca. 21 retrorse teeth; palpus, 0.46 mm long, distal
article 0.23 mm long, sensory vesicle spherical, occupying 0.50 width of third article, opening 0.50 width of vesicle, ca. 13
sensillar sockets visible. Cibarial pump: space between proximal arms U-shaped, smooth, as wide as deep. Thorax: dark
brown, lightly pilose; length 1.1 mm; postpronotal lobe pollinose at junction with scutum (in dorsal view); scutum
uniformly dark brown, vestiture of fine silver scales; scutellum pale brown, angle of posterior apex obtuse, sides slightly
concave, vestiture of sparse black hairs; postnotum dark brown; pleuron medium brown. Wing: length 1.8-2. 5 mm,
maximum width 0.8-1. 2 mm; stem vein hair tuft distinct; hairs on basicosta not prominent. Halter: pale gray. Legs:
forecoxa concolourous with pleuron; trochanter, femur and most of tibia mottled yellow-brown; distal portion of tibia and
tarsus dark brown; mid- and hind leg as for fore leg; hind basitarsus yellow with distal portion and tarsomeres mottled
yellow and dark brown; pretarsal claw slender with sharp apex, basal tooth 0.50 length of claw, lying subparallel to claw.
Abdomen: generally dark brown; basal fringe of golden hairs extended to segment II; tergites slightly pollinose, mottled,
vestiture of dark hairs; posterior tergites subshining with sparse dark hairs; sternum pale gray. Genitalia (Fig. 9): cercus,
in ventral view, slightly curved laterally, curved posteromedially; stem of genital fork, thin, smooth, lateral sclerite
triangular with anterior apex forming acute angle, directed laterally, notched, or not; hypogynial valves slightly curved
medially, broadly rounded posteriorly, extended to anal lobes; sternite VIII with ovoid median pigmented region.
Spermatheca: slightly ovoid; not densely pigmented; no reticulation; unpigmented area at junction of sperm duct normal.
Adult Male. Body: generally dark brownish-black; length 1.6-1. 9 mm. Head: width 0.5 mm, depth 0.4 mm. Eyes:
upper ommatidia, 0.04 mm in diameter, 12 rows both up and across eye; lower ommatidia 0.1 mm in diameter. Clypeus:
0.20 as wide as head. Antenna: total length 0.3 mm. Mouthparts: 0.50 length of head depth; mandibles 0.33 length of
labrum; lacinia as long as labrum, no retrorse teeth, apex hairy; palpus 0.4 mm long, distal article 0.2 mm, sensory vesicle
spherical, occupying only 0.33 of width of third palpal article, opening 0.50 width of vesicle, ca. 10 sensillar sockets visible.
Thorax: length 0.7 mm; postpronotal lobe pale; scutum evenly dark brown to black, vestiture of moderately long, golden
scales; scutellum pale, slightly convex laterally, forming right angle at apex, vestiture of sparse, long black hairs;
postscutellum dark brown; pleuron brown. Wing: length 1.4-1. 6 mm, maximum width 0.6-0. 7 mm; stem vein hair tuft
small, but prominent. Halter: greyish brown. Legs: coxae brown, trochanter and portions of femur and tibia pale, banding
distinct, vestiture of dense black hairs; pretarsal claw with ca. 23 grappling hooks on each claw. Abdomen: brown; vestiture
of very sparse golden scales; basal fringe of pale hairs extended to abdominal segment II; tergites V-VII mottled. Genitalia
(Fig. 16): gonocoxa 1. 8 times as long as basal width; gonostylus approximately 2.5 times as long as basal width, single
blunt terminal spine; ventral plate with median posterior apex rounded, slightly convex sides expanded laterally to broadly
rounded angle, extended anteriorly to pigmented arms with rounded apices, anteromedially slightly domed, with broad
medial notch, median hairy patch, slightly expanded anteriorly.
Pupa. Length: 1.8-2. 4 mm male, 1.9-2. 3 mm female. Gill (Fig. 21): length 0.8-1. 2 mm male, 0. 7-1.1 mm female;
branching pattern typical, dorsal-most filament long, often reflexed posteriorly over thorax under cocoon. Thorax: cuticle
light brown; tubercles rounded (Fig. 71-73); scutellum in lateral view with distinct blunt peak. Abdomen: tergite I bare,
tergites 1 1— I V with anteriorly directed hooks, tergite V bare, tergites VI— VI 1 1 with fine spine combs; sternites IV and V
with hooks close to mid-line, those on sternites VI and VII more lateral, evenly spaced; caudal spines absent; terminal
caudal sclerites, distinct, angular.
Larva. Last instar with dark pharate pupal gills; length 3. 9-4. 9 mm. Body: greyish with anterior intersegmental
regions light, producing a banded appearance. Head capsule (Figs. 35, 65): generally dark brown to blackish; frontoclypeal
Quaest. Ent., 1987,23 (3)
396
Craig
apotome pale anteriorly, very dark posteromedially; spot pattern generally positive, or posteromedian and posterolateral
spot-two, negative, or concolorous with surrounding head cuticle; apotome with anterior pale area extended posteriorly
along ecdysial line to posterolateral spot-one; cervical sclerites distinct. Stemmata: surrounded by small clear area, with, or
without, curved anterodorsal mark. Head cuticle: corrugated transversely, bent, or not, around anteromedian head spots in
form of U- or V- shaped mark anteriorly; sensilla number normal. Antenna: as long as labral fan stem, distal article 0.1
mm, proximal articles 0.2 mm. Labral fans: with 30-33 rays, ca. 0.45 mm in length; microtrichia of medial rays up to 1.4
times longer than ray width, no microtrichial pattern. Hypostoma (Fig. 44): median tooth protruded, other teeth in line
sloped posterolaterally; five sublateral teeth per side; lateral tooth slightly higher and broader; one paralateral tooth; two to
three lateral serrations; six to eight hypostomal sensilla per side. Postgenal cleft: wider than deep, squarish, with either flat
or slightly bifurcate apex. Postgenal bridge: 2.3 times length of cleft (Fig. 35). Mandible: apex (Fig. 56) with nine to 11
spinous teeth; distance, between spinous teeth and mandibular serration less than 0.50 width of serration base; serration as
high as basal width; sensillum small and not on serration. Maxillary palpus: 3.0 times as long as width at base. Mandibular
phragma: extended .ventrad to 0.33 depth of maxilla base. Abdomen: segments increased gradually in size to segment V;
segment VI markedly enlarged ventrally; at segment VIII size reduced sharply (Fig. 23); ventral tubercles placed laterally;
posterodorsal cuticle with low, laterally ovoid tubercles and with trichoid sensilla, 9.2 am long. Anal sclerite: with
posteroventral arms 3.0 times longer than dorsolateral arms; lateral accessory sclerite absent, but in darker specimens
evident as lightly pigmented disc. Circlet of hooks: 123 rows of hooks, 21-23 hooks per row. Anal papillae: complex (Fig.
23).
Bionomics. — Simulium tahitiense is the most wide-spread species in Tahiti. Its larvae are
found mainly in the larger rivers, in dense aggregations on and under large boulders in strong
flow. If vegetation trails in high velocity water it usually supports large populations of S.
tahitiense also. In the Papanoo Valley and elsewhere, the adults form dense swarms around
humans and cause considerable nuisance although they do not bite. Larvae of the ephydrid
Apulvillus cheesmanae Edwards (W. N. Mathis, pers. comm., 1983) are found with many of
the larger aggregations of Simulium tahitiense. Although extremely slow moving there is a
possibility that A. cheesmanae larvae are predaceous on the simuliid larvae. S. tahitiense larvae
can be found in almost all the other simuliid habitats in Tahiti. In more shaded localities, such
as at Bain Loti, it is associated with S. lotii and S. exasperans larvae, but its larvae prefer hard
substrate to vegetation. Schroder (1985) provides details of the feeding behaviour of larvae of
this species.
Material Examined. — In addition to types, I have seen the following:
Papeete , Fautaua R., Bain Loti. Tahiti, Baini (sic) Loti, April 14, 1961, J.N. Belkin #35 (4 2, USNM); 17° 33' 15" S.
149° 33' 00" W. alt. 92m, ll-vii-74 (larvae, DSIR; UASM), 15-vii-74 (larvae, UASM), 16-vii-74 (larvae, UASM),
07-vii-80 (larvae, CNCI), 08-vii-80 (1 3, CNCI; 2 2, DSIR), 1 l-vii-80 (5 2, 5 3, BMNH; 5 2, 5 3, BPBM; 1 2, 5 3, CNCI;
1 2, DSIR; 1 3, UASM; 6 2, 2 5, USNM; karyotyped larvae, BMNH, UASM; larvae, UASM) D.A. & R.E.G. Craig.
Haapape , Tuauru R., 17° 32' 10" S. 149° 29' 15" W. alt. 50m. 09-vii-80, D.A. & R.E.G. Craig (1 <3, BMNH; 1 <3,
BPBM; larvae, DSIR; 2 3, UASM; 2 3, USNM). Papenoo, Papenoo R„ 17° 33 ' 15" S. 149° 25 ' 50" W. 22-X-28, A.M.
Adamson (1 3, BMNH); 17° 36' 52" S. 149° 24' 51" W. 10-vii-80, D.A. & R.E.G. Craig (1 2, DSIR; 1 2, 3 3, MNHP; 4
3, UASM; karyotyped larvae, BMNH).
Tiarei, Fareteuira R., Cascades de Faarumai, 17° 32' 10" S. 149° 23 ' 48" W. 14-vii-80, 2 1 -vii-80, D.A. & R.E.G. Craig
(larvae, UASM). Onofea R„ 17° 33' 30" S. 149° 23' 47" W. alt. 110 m. 14-vii-80, D.A. & R.E.G. Craig (larvae,
UASM).
Mahaena, Mahape R., 17° 34' 00" S. 149° 20' 33" W. alt. 700 m 14-vii-80, D.A. & R.E.G. Craig (karyotyped larvae,
BMNH).
Faaone , Utuofai R„ 17° 39' 15" S. 149° 19' 00" W. 10-vii-73, D.A. & R.E.G. Craig (larvae, UASM).
Tautira, (Tahiti-iti), E. trib. Vaitepiha R„ 17° 46' 30" S. 149° 15 ' 20" W. alt. 50 m. 19-vii-80, D.A. & R.E.G. Craig (1
2, 3 3, DSIR; larvae & pupae BMNH); Vaitepiha R., 17° 46' 30" S. 149° 10' 21" W. alt. 50m. 19-vii-80, D.A. &
R.E.G. Craig. (3 2, CNCI; 1 3, DSIR; karyotyped larvae, BMNH; 1 3, larvae, UASM).
Teahupoo, (Tahiti-iti), Tiirahi R., 17° 51 ' 05" S. 149° 14' 15" W. 20-vii-80, D.A. & R.E.G. Craig (karyotyped larvae,
BMNH; 3 2, 1 3, larvae, pupae, CNCI; 2 2, 2 3, larvae, pupae, DSIR; larvae and pupae UASM).
Paea, Ofaipapa, Robinson‘s Place, 02-iv-61, 28-iv-61, J.N. Belkin (larvae, USNM). Vaitiu R., 17° 34' 00" S. 149° 41 '
22" W. 16-vii-74, D.A. & R.E.G. Craig (larvae, UASM).
Mataiea, Vaihiria R., (= Tahiria) 17° 44' 22" S. 149° 24' 15" W. alt. 80 m. 16-vii-80, (1 2, 1 3, DSIR; 1 2, larvae,
UASM); 17° 41' 31" S. 149° 25' 02" W. alt. 250m. 17-vii-80, (1 2, larvae, pupae, BMNH; 1 2, 2 3, CNCI; 1 3, DSIR; 1
2, USNM; larvae, UASM), D.A. & R.E.G. Craig.
Mahaiatea, Taharuu R., 17° 42 ' 45" S. 149° 29 ’ 1^" W. alt. 135 m. 15-vii-80, D.A. & R.E.G. Craig (larvae, CNCI).
Punaauia , Punaruu R., 27-iv-1969, B. Hocking (larvae, UASM); 17° 38' 20" S. 149° 35' 40" W. 21-vi-73, 02-vii-73,
Society Islands Black Flies
397
03-vii-73, 04-vii-73, (larvae, UASM); 17° 38' 20" S. 149° 35' 05" W. alt. 120m. 05-vii-73 (larvae, UASM), 06-vii-73
(larvae, UASM), 09-vii-73 (larvae, UASM), 05-vii-74 (larvae, UASM), 06-vii-74 (larvae, UASM), 08-vii-74 (larvae,
UASM), 22-vii-74 (larvae, UASM), 07-vii-80 (2 <3, BMNH; 1 2, MNHP; 1 2, 1 <3, UASM) D.A. & R.E.G. Craig.
Simulium (Inseliellum.) “IIS“
Larvae of this cytospecies as identified by Rothfels {in Craig, 1983), cannot be
morphologically separated from larvae of S. tahitiense. Possibly these larvae will eventually be
shown to be those of S. exasperans (Rothfels, pers. comm., 1986), however, “IIS” is known
only from Tahiti-iti.
Material Examined. — Tautira, (Tahiti-iti), Vaitepiha R., 17° 46' 30" S. 149° 10' 21" W. alt. 50 m.
19-vii-80, D.A. & R.E.G. Craig (5 karyotyped early instar larvae as slide mounts, DAC).
Remarks. — The material was taken with S. tahitiense larvae.
oviceps - group
Diagnosis. —
Pupa : scutellum rounded dorsally; with or without tubercles, if present, pointed; gill with
dorsal filaments shorter, or subequal in length to others.
Larva: labral fans reduced; ray number small, rays short to almost absent; head spot pattern
partially or completely negative; head margins prominently convex; postgenal cleft small to
absent; head sensilla number normal or greatly increased; hypostomal teeth variable, from
medial and lateral teeth sharp and prominent, to all teeth rounded and subequal in length, to
median tooth with adjacent sublateral teeth short, or median tooth absent; posterior abdomen
expanded abruptly laterally at segment VI, decreased in size more gradually posteriorly, or
shape more normal. Anal sclerite with posteroventral arms extended partway, or virtually
completely around posterior proleg. Anal papillae simple, with at most, only small accessory
basal papillae.
Included taxa. — S. castaneum n. sp.; RAIATEA. S. admixtum n. sp., S. arlecchinum n.
sp., S. cataractarum n. sp., S. mesodontium n. sp., S. neoviceps n. sp., S. oviceps Edwards and
Simulium sp.; TAHITI.
Simulium ( Inseliellum ) admixtum n. sp.
(Figs. 36, 45)
Type.—
Holotype. Larva: penultimate instar; as slide mount; label data:-“Holotype”, “Simulium (I.)
admixtum Craig/ det D.A. Craig 1986”, “Tahiti, Tiarei, Fareteuira R.,/ Cascades de
Faarumai, /17° 32 ' 10” S. 149° 23' 48" W. /16-ii-83, D.A. Craig” (MNHP).
Specific epithet. — Named after the Greek word “admixtus” meaning “mixed”, in reference
to the unique suite of characters possessed by the single larva.
Diagnosis. —
Larva: head; sensilla numerous; spot pattern negative; median hypostomal tooth lacking;
abdomen; cuticle with numerous, low ovoid tubercles.
Description. —
Adult Female. Unknown.
Adult Male. Unknown.
Quaest. Ent., 1987,23 (3)
398
Craig
Pupa. Unknown.
Larva. Single penultimate instar larva; length 5.2 mm. Body: generally greyish with indistinct intersegmental regions.
Head capsule: margins strongly convex posteriorly, as for S. cataractarum (Fig. 29); background colour pale with apices of
corrugations dark brown; frontoclypeal apotome lighter anteriorly, darker posteriorly; anteromedian and posteromedian
head spots negative, but barely so, posterolateral spots concolourous with surrounding darker brown; cervical sclerites
fused to postocciput by narrow junction; stemmata surrounded by dark brown anterodorsally, narrow oblique clear
posterodorsal region; cuticle with numerous lateral corrugations dorsally and ventrally; sensilla numerous with raised and
strongly pigmented sockets, producing rugose appearance. Antenna: length 0.4 mm, just longer than labral fan stem, distal
article length 0.1 mm. Labral fans: with 26 rays, nine lateral rays less substantial, ca. 0.6 mm. in length; microtrichia of
median rays 0.50 times as long as ray width, pattern of long microtrichia interspersed between eight to 10 smaller
microtrichia. Hypostoma (Fig. 36): median tooth lacking, four sublateral teeth per side, increasing slightly in length
laterally, lateral teeth prominent, one paralateral tooth with steep base down to four lateral serrations, six hypostomal
sensilla in row per side. Postgenal cleft: very shallow, broadly U-shaped, 5.0 times wider than deep. Postgenal bridge: 4.0
times longer than cleft. Mandible: apex (Fig. 45) with seven to nine spinous teeth extended almost to mandibular serration;
serration as high as basal width, anterior edge longer than posterior, sensillum virtually absent from one mandible, double
on other. Maxillary palpus: 3.5 times as long as width at base. Mandibular phragma: heavily pigmented, extended ventrad
to 0.33 depth of maxilla base. Abdomen: segments I— IV of similar size, gradually increased in size to segment VIII;
posteroventral tubercles situated laterally and very small; posterodorsal cuticle with clear, low ovoid tubercles and with
trichoid sensilla, 16.0 nm long. Anal sclerite: with posteroventral arms 2.3 times longer than dorsolateral arms; pigment
more distinct along anterior edges, sensillar sockets numerous and obvious. Circlet of hooks: 91 rows of hooks, 18-20 hooks
per row. Anal papillae: simple, but each with very small basal papilla.
Bionomics. — The single larva was collected from the sheet of water flowing down the
vertical rock face on the western side of the Cascade de Vaimahuta (Cascades de Faarumai),
along with larvae of S. cataractarum , S. oviceps and S. neoviceps. Water temperature was 17°
C. This temperature is relatively cool for simuliid localities in Tahiti.
Phylogenetic Relationships. — This species is similar in many respects to S. cataractarum
in possessing strongly convex head margins, negative spots, lateral teeth of hypostoma longer
than remainder of teeth and fused cervical sclerites. Eventual cytological examination of this
species will probably show a close relationship. However, lack of the median hypostomal tooth
(Fig. 36), and possession of two mandibular serrations on one mandible (Fig. 45), are unique.
Simulium (Inseliellum) arlecchinum n. sp.
(Figs. 27, 46)
Types. —
Holotype. Larva: mature last instar as slide mount; thorax used for scanning electron
microscopy; hypostoma damaged; label data:- “Holotype”, “Simulium (I.) arlecchinum Craig/
det/ D.A. Craig 1986”, “Tahiti, Mataia, Vaihiria R.,/ 21-ii-84, P. Schroder” (MNHP).
Paratypes. Larvae: penultimate instars; as slide mounts; label data:- as for holotype (1,
BMNH; 1, MNHP).
Specific epithet. — Based on the Italian word “arlecchino”, meaning “harlequin” - in
reference to the distinctive colour pattern on the ventral surface of the abdomen of the larvae.
Diagnosis. —
Larva: colour pattern on frontoclypeal apotome H-shaped; lateral hypostomal teeth
prominent; abdomen with triangular pattern ventrally.
Description. —
Adult Female. Unknown.
Adult Male. Unknown.
Pupa. (Based on pharate material from holotype larva). Gill: length 0.9 mm; branching pattern normal, dorsal-most
filament more than half as long as other filaments. Thorax: tubercles absent.
Larva. Last instar with dark pharate pupal gills; length 4.2 mm. Body: greyish-brown with pale narrow intersegmental
regions. Head capsule (Fig. 27): margins generally parallel-sided; generally light brown; frontoclypeal apotome with broad
Society Islands Black Flies
399
pale bands along ecdysial sutures; anteromedian and posteromedian head spots negative, H-shaped brown region
surrounding muscle spots, extreme posteromedian region darker; cervical sclerites distinct; cuticle essentially smooth,
corrugations normal; sensilla number normal. Antenna: slightly longer than labral fan stem, distal article length 0.8 mm,
proximal article length 0.19 mm. Labral fans: with ca. 18 normal rays, 0.56 mm. in length, plus 13 smaller, less robust
rays; microtrichia of medial rays up to 1.3 times longer than ray width, no microtrichial pattern. Hypostoma: essentially as
for S. tahitiense (Fig. 44), but sublateral teeth slightly more prominent; six to eight hypostomal sensilla in row per side.
Postgenal cleft: twice as wide as deep, sloping to slightly rounded vertex. Postgenal bridge: 3.0 times longer than cleft.
Mandible: apex (Fig. 46) with ten spinous teeth extended essentially to mandibular serration; serration as high as basal
width; proximal sensillum 0.50 height of serration. Maxillary palpus: 2.5 times as long as width at base. Mandibular
phragma: extended ventrad to dorsal edge of maxillary base. Abdomen: segments I— III with triangular and diagonal
pattern on sterna, less distinct on terga; posterior abdominal shape as for S. cataractarum ; posteroventral tubercles placed
laterally and not obvious; posterodorsal cuticle with ovoid, scale-like tubercles and with trichoid sensilla, 4.0 /an long. Anal
sclerite: with posteroventral arms 1 .4 times longer than dorsolateral arms; no lateral accessory sclerites. Circlet of hooks:
108 rows of hooks, 18-20 hooks per row. Anal papillae simple.
Bionomics. — Collected from grass trailing in the main stream of the unshaded, upper
Vaihiria River, in fast, 5 cm deep water (P. Schroder, pers. comm., 1985).
Phylogenetic Relationships. — This species is placed in the oviceps - group on the basis of
abdominal shape and negative head-spot pattern; however, head capsule and mouthpart
structure are very similar to those of species in the tahitiense - group.
Simulium (Inseliellum) castaneum n. sp.
(Figs. 2, 28,37,47,58,60,61)
Types. —
Holotype. Larva: slide mount; label data:- “Holotype”, “Simulium (I.)/ castaneum Craig”,
“Society Islands, Raiatea, Temehani R., / 1 800’, l-ix-77, S.L. Montgomery,/ Bishop Museum
Acc. #1977.361”, “det. D.A. Craig 1986” (BPBM).
Paratypes. Adult: female; cleared and in vial; label data:- “Simulium (I.)/ castaneum Craig
/det./ D. A. Craig 1986”, “SOCIETY IS/ Raiatea, Uturoa,/ 0-100 m. III. 1971.”, “N.H.L.
Krauss/ Collector/ BISHOP MUSEUM” (BPBM). Material in alcohol: one pupa, five last
instar larvae, 10 penultimate larvae, twenty( + ) earlier instar larvae; label data:- as for
holotype (BPBM).
Specific epithet. — Based on the Greek word “castanea”, meaning “brown”, in reference to
the generally even dark-brown of the larval head, labral fans, pupa and its cocoon.
Diagnosis. —
Larva: head uniformly dark brown; labral fan rays brown; hypostomal teeth very sharp.
Description. —
Adult Female. (From single specimen and immature pharate specimens). Body: generally black. Head: width 0.7 mm,
depth 0.5 mm. Eyes: interocular distance 0.12 head width; frontal angle 86 degrees; ommatidia 0.01 mm in diameter, ca.
28 respectively across and up eye in middle row. Vertex: dark brown; vestiture of sparse dark brown hairs. Antennae: dark.
Mouthparts: length, 0.33 head depth; mandibles, shorter than labrum, with ca. 14 minute teeth; lacinia with ca. 15-16
retrorse teeth; palpus, sensory vesicle 0.33 width of third article, opening 0.33 width of vesicle, ca. 10 sensillar sockets
visible. Cibarial pump: space between proximal arms U-shaped, as wide as deep, smooth. Thorax: length 0.85 mm; scutum
dark brown, shiny; scutellum lighter. Wing: length 1.7 mm, maximum width 0.8 mm. Legs: hind basitarsus parallel-sided,
7.5 times longer than wide; pretarsal claw smoothly curved with pointed apex, basal tooth 0.20 length of claw. Genitalia
(Fig. 2): cercus bluntly cone- shaped in ventral view; hypogynial valves broadly rounded apically, extended to anal lobes,
strongly diverged medially; stem of genital fork narrow, slightly swollen anteriorly, lateral triangular sclerites with blunt
anterior apices; sternite VIII sclerotized medially, narrow posteriorly, widening anteriorly. Spermatheca: ovoid;
pigmented; no pattern; clear area at junction of sperm duct normal.
Adult Male. Unknown.
Pupa. Length: 2.8 mm, female. Gill: length 1.3 mm; branching pattern typical, dorsal-most filament 0.50-0.66 length
of others. Thorax: tubercles absent, cuticle shiny. Abdomen: tergite I without hooks or spines; tergites 1 1— IV with
pronounced recurved hooks, those of tergite III smaller; tergites V and VI bare; tergites VII and VIII with very fine spine
fields, those of tergite VIII stronger; sternite III clear; sternites IV-VII with anteriorly directed hooks, those of tergites IV
Quaest. Ent., 1987,23 (3)
400
Craig
and V grouped close to midline, those of sternites VI and VII widely spaced; caudal spines very small. Cocoon:
slipper-shaped, of medium weave; darkly coloured; covering pupa to base of gills,; anterior edge loosely woven, with spaces
between threads.
Larva. Last instar larva with dark pharate pupal gills; length 6.2 mm. Body: evenly dark greyish-brown with thin
lighter intersegmental regions. Head capsule (Fig. 28): relatively uniformly dark brown; posterolateral margins strongly
convex; frontoclypeal apotome slightly lighter anteriorly, darker posteriorly; posteromedian and anterolateral head
spot-two, negative, others positive; stemmata with darker area anteriorly, distinct lighter diagonal mark; cuticle with
corrugations running laterally; sensilla numerous, with distinct raised sockets (Fig. 60); cervical sclerites fused to
postocciput. Antenna: longer than labral fan stem, distal article length 0.14 mm, proximal articles length 0.34 mm, basal
article very dark brown and longitudinally striated, medial article with irregular lighter patches, distal article uniformly
light brown. Labral fans: with 29 dark brown rays, ca. 0.88 mm. in length, lateral four rays less substantial; microtrichia of
all rays short, only 0.6 times as long as ray width, pattern of long microtrichia interspersed with five smaller microtrichia,
very distinct (Fig. 58). Hypostoma (Fig. 37): teeth sharply pointed, median tooth prominent. Five sublateral teeth per side,
subequal in length, line of teeth tips sloping anteriorly to prominent lateral tooth longer than median tooth, one paralateral
tooth, with steep base down to five or six lateral serrations; five or six hypostomal sensilla in row per side. Postgenal cleft:
small, wider than deep, flat anteriorly. Postgenal bridge: 2.0 times longer than cleft. Mandible: apex (Fig. 47) with nine to
11 spinous teeth extended almost to mandibular serration; serration as high as basal width, length of sides subequal;
proximal sensillum cone-shaped, on base of serration. Maxillary palpus: 3.1 times as long as width at base. Mandibular
phragma: extended ventrad to 0.50 depth of maxilla base. Abdomen: segments increased gradually in size posteriorly to
segment IV, then smoothly increased to maximum size at segment VII; posteroventral tubercles very small; posterodorsal
cuticle with regular hemispherical tubercles and with trichoid sensilla, 15.0 /an long (Fig. 61). Anal sclerite: with
posteroventral arms broad and continuous with lateral accessory sclerite extended to ventral surface, but not fused
ventrally. Circlet of hooks: 106 rows of hooks, 14-16 hooks per row. Anal papillae: essentially simple; lateral papillae with
one or two very small pointed basal papillae.
Bionomics. — Little is known about the habitat of this species. Larvae and pupae were taken
from a cascade; a piece of grass was included with the material, but it is not known if the
specimens were on vegetation.
Material Examined. — In addition to types I have seen the following: Two pupal cocoons
and pieces of pupal exuviae, in alcohol, data as for holotype (BPBM).
Simulium (Inseliellum) cataractarum n. sp.
(Figs. 3, 10, 24, 29, 38,48,64)
Simulium “IS". Craig, 1983: 534. Schroder, 1985: 18.
Types. —
Holotype. Male: pinned; reared; emerging from pupa, with larval abdominal exuviae still
attached; label data:- “Holotype”, “<$”, “Simulium (I.)/ cataractarum Craig/ det./ D.A. Craig
1986”, “TAHITI, Mataiea, E. Lac Vaihiria,/ 17° 40f 58" S. 149° 24' 51" W. / alt. 480m.
22-vii-80, / D.A. Craig. Stat. #16.” (MNHP).
Paratypes. Female: pinned; reared; pupal exuviae and cocoon as subsidiary material; head
and genitalia in vial; label data as for holotype (MNHP). Females: pinned; reared; pupal
exuviae and cocoons as subsidiary material; label data:- “TAHITI, Tiarei, Fareteuira R.,/
Cascade de Faarumai,/ 17° 32’ 10" S. 149° 23’ 28" W./ 17-ii-87, D.A. Craig. Stat. #15” (1
each, BPBM; BMNH; CNCI; DSIR; USNM; UASM). Male: pinned; reared; with pupal
exuviae and cocoon as susidiary material; complete specimen cleared and in vial; label data as
for holotype (MNHP). Male: pinned; reared; with pupal exuviae and cocoon as subsidiary
material; head and genitalia in vial; label data as for holotype (BMNH). Males: pinned; reared;
pupal exuviae and cocoons as subsidiary material; label data:- “TAHITI, Tiarei, Fareteuira
R.,/ Cascade de Faarumai,/ 17° 32’ 10" S. 149° 23’ 28" W./ 17-ii-87, D.A. Craig. Stat.
#15” (1 each, BPBM; BMNH; CNCI; DSIR; USNM; UASM). Pupa: (in alcohol), posterior
larval exuviae still attached; label data as for holotype (BMNH). Larvae: in alcohol; label data
as for holotype (9 penultimate, BMNH). Larvae: karyotyped; label data as for holotype (4,
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401
BMNH; 3, BPBM; 4, MNHP). Larvae: in alcohol; locality data:- “Tiarei, Cascades de
Faarumai, Fareteuira R., 17° 32’ 10" S. 149° 24’ 00" W. 20-vii-80, D.A.C.” (8 last,
MNHP). Larvae: penultimate; label data:-“Mataiea, E. Lac Vaihiria, /17° 40’ 58" S. 149°
24' 51" W./ alt. 430m. 17-vii-80, /D.A. & R.E.G. Craig”. (14, CNCI; 19, DSIR). Larvae:
label data:- “Tiarei, immediately west of Trou du Souffleur, 17° 31 ’ 30" S. 149° 23 ' 32" W.
alt. 10m. 17-ii-87,/ D.A.C.” (9 penultimate, DAC).
Specific epithet. — Named after the Latin word “cataracta” for “water fall”, in reference to
the cascades where larvae of this species are usually found.
Diagnosis. —
Pupa: gill with short dorsal filament; thoracic tubercles absent.
Larva: oviceps -type; head spot pattern negative; labral fan ray number reduced, rays
pigmented medially; anal sclerite extended ventrally around anal proleg.
Recognition. — The holotype was chosen because the larval abdominal exuviae and the
pupal exuviae provide definitive association to all stages. Larvae of S. cataractarum are very
distinctive and it is quite suprising that they were not discovered previously. Included here for
the present is the single karyotyped larval specimen designated Simulium “IS-complex” by
Rothfels {in Craig, 1983). That specimen (DAC) is morphologically indistinguishable from
larvae of S. cataractarum.
Description. —
Adult Female. Body: generally dark brown-black; length 1.7 mm. Head: width 0.6 mm, depth 0.3 mm. Eyes:
interocular distance 0.6 times width of clypeus; frontal angle, 105 degrees; ommatidia 0.016 mm in diameter, ca. 33 and 37
respectively across and up eye in middle row. Post-ocular hairs: extended just to eye margin. Clypeus: pale, concolourous
with ventral frons, as wide as deep; vestiture of substantial pale hairs. Antenna: total length 0.36 mm; scape, pedicel and
base of first flagellomere yellowish, remaining flagellomeres light brown. Mouthparts: length, 0.33 head depth; mandibles,
shorter than labrum with ca. 14 fine teeth; lacinia with 14-16 retrorse teeth; palpus, 0.5 mm long, distal article 0.2 mm
long, third article with pronounced distomedial angle; sensory vesicle occupying 0.50 width of third article, opening 0.33
width of vesicle, ca. 20+ sensillar sockets visible. Cibarial pump: space between proximal arms smooth, U - shaped, as
wide as deep. Thorax: length 0.9 mm; postpronotal lobe with tuft of yellow hair extended laterally beyond head margin;
scutum uniformly blackish brown, anterolateral margins lighter, vestiture of substantial pale hairs; scutellum with
distinctly concave posterolateral edges, vestiture of pale golden and dark hairs meeting medially; postscutellum
concolourous with abdomen; pleuron brown; mesepimeral tuft of hairs golden. Wing: length 1.6 mm, maximum width 0.8
mm: veins pale yellow; stem vein hair tuft sparse; basicosta with darker hairs, remainder of costa with spines and hairs.
Halter: pale. Legs: foreleg light brown; fore-femur 0.63 mm long; fore-basitarsus 6.0 times as long as wide, tarsomeres
dark brown: meso- and metatrochanter dark brown, distal quarters of meso-femur and tibia darker brown; basitarsus
darker brown on distal 0.33; hind basitarsus parallel-sided, 6.0 times longer than wide, distal tarsomere squarish; pretarsal
claw slender, gently curved with sharp apex, basal tooth 0.50-0.66 length of claw. Abdomen: generally dark brown; basal
fringe of pale yellow hairs extended to abdominal segment II; tergites not distinct, vestiture of tergites I-VI of pale shiny
hairs, those of tergites VII-IX darker; pleural regions concolourous with tergum; sternites paler. Genitalia (Fig. 3): cercus,
bluntly rounded in ventral view; hypogynial valves extended to anal lobe, medial margins slightly concave, converging
posteriorly, broadly rounded posteriorly; stem of genital fork smooth, lateral arms broad, lateral sclerite large, with apex
anteriorly-directed, sharply angulate; sternite VIII pigmented region broadly dome-shaped. Spermatheca: ovoid; slight
reticulate pattern; not heavily pigmented; clear area at junction of sperm duct large.
Adult Male. Body: generally blackish-brown; length 1.7 mm. Head: width 0.5 mm, depth 0.4 mm. Eyes: upper
ommatidia 0.03 mm in diameter, 19 and 22 respectively up and across middle rows; lower ommatidia 0.01 mm in diameter,
eight and 12 respectively up and across middle rows. Labrum: pale. Clypeus: narrow dorsally, widening ventrally, 0.20 as
wide as head. Antenna: length 0.4 mm; pedicel and scape slightly lighter in colour than flagellum; pilose. Mouthparts: 0.25
length of head depth; mandibles insubstantial, three very small apical teeth; lacinia hairy, without retrorse teeth; palpus
0.29 mm long, distal article 0.27 mm, sensory vesicle occupying only 0.50 of width of third palpal article, opening less than
0.33 width of vesicle, eight to 10 sensillar sockets visible. Thorax: length 0.24 mm; postpronotal lobe with small tuft of
golden hairs; scutum black, vestiture of golden hairs, longer posteriorly; scutellum pale yellow, posterior edges slightly
concave in dorsal view, vestiture of lateral golden hairs; postscutellum dark, dull brown. Wing: length 1.5 mm, maximum
width 0.7 mm; veins pale; stem vein dark, hair tuft black, substantial. Halter: yellowish. Legs: coxae shiny brown;
fore-femur with yellow medial area; middle and hind femurs similar, vestiture of long dark hairs; tibiae dark on posterior
margin, fore basitarsus parallel sided, but expanded slightly distally, 5.0 times longer than wide; fore tibia 7.0 times as long
as wide; pretarsal claw smoothly curved, ca. 22 grappling hooks. Abdomen: generally blackish-brown; basal fringe of
yellow hairs extended to abdominal segment III; tergite II yellow anteriorly; tergites I-VI with vestiture of golden hairs,
Quaest. Ent .. 1987,23 (3)
402
Craig
that on tergites VII and VIII darker; pleuron mottled brown; sternites III pale yellow laterally, squarish. Genitalia (Fig.
10): gonocoxa ca. 2.0 times as long as basal width; gonostylus ca. 0.33 as wide as long, strongly curved with one blunt
terminal spine, as long as wide; ventral plate with posterior apex broadly rounded, hairy, posterolaterally slightly concave,
broadly angulate, anterior arms heavily sclerotized and angulate, anteromedian notch distinct.
Pupa. Length 1.9-2. 6 mm. Gill: length 1. 4-2.0 mm; branching pattern typical, dorsal-most filament 0.33-0.50 length
of other filament, occasionally reflexed posteriorly over thorax. Thorax: cuticle clear, brown; tubercles absent. Abdomen:
tergite I— 1 1 1 bare; tergites IV and V with hooks; caudal spines very short, but distinct. Cocoon: slipper-shaped.
Larva. Last instar with dark pharate pupal gills; length 4. 3-5. 3 mm. Body: greyish-brown, with slightly paler
intersegmental regions. Head capsule (Fig. 29): generally rich brown; margins convex, widest at 0.33 length from posterior
of capsule (Fig. 64); frontoclypeal apotome brown anteriorly; head spots negative, surrounded by darker area; cuticle
slightly rugose, with corrugations; sensilla numerous; cervical sclerites distinct, but fused to occiput; Antenna: slightly
shorter than labral fan stem; length 0.38 mm; distal article 0.13 mm long, proximal article darker than head capsule, other
articles lighter; base pigmented. Labral fans: with 12-15 robust brown rays ca. 0.8 mm. in length, plus three to five
thinner, smaller medial rays; each! ray with median dark brown patch; microtrichia of rays 0.50 as long as ray width, no
microtrichial pattern. Hypostoma (Fig. 38): flexed dorsally; median tooth rounded, not protruding beyond prominent
lateral teeth; five sublateral teeth per side increasing in size slightly to prominent lateral teeth; one small paralateral tooth;
six lateral serrations; five to six hypostomal sensilla in row per side. Postgenal cleft: small, U-shaped, twice as wide as deep.
Postgenal bridge: 2.5 times as long as cleft (Fig. 29). Mandible: apex (Fig. 48) with 16 to 20 spinous teeth, decreased
rapidly in size posteriorly, extended almost to mandibular serration; serration prominent, as high as basal width, anterior
edge twice as long as posterior edge, basal sensillum minute and on serration base, heavily pigmented. Maxillary palpus:
4.1 times as long as width at base. Mandibular phragma: heavily pigmented and substantial, extended ventrad to 0.50
depth of maxillary base. Abdomen: segments increased gradually in size to segment V where a sudden lateral increase in
size occurs (not as prominent as in 5. oviceps ); posteroventral tubercles prominent (Fig. 24); posterodorsal cuticle smooth,
with numerous trichoid sensilla 1 1.0-22.0 /am long. Anal sclerite: with median portion broad, deeply pigmented: sensillar
sockets showing clearly; anterodorsal arms with small lateral extension; posteroventral arms continuous with lateral
accessory sclerites, extended, or almost, completely around posterior proleg. Circlet of hooks: 108 rows of hooks, 15-17
hooks per row. Anal papillae: simple.
Bionomics. — A typical habitat for larvae of this species, such as at the Cascades de
Faarumai (= La Trois Cascades) and at Lac Vaihiria, consists of thin sheets and trickles of
water flowing over hard rock. At Cascades de Faarumai, larvae were on the rock face, on
filamentous algae, leaves of macrophytes and dead leaves blown onto the rock face. The
Cascades de Faarumai (Vaimahuta Cascade) site is of spring origin, arising from a horizontal
crack two-thirds of the way up, and to the right of the main falls. On the 1 7-ii- 1 987 suspended
particulate material larger than 0.45 pm in the water was 6.9 mg/litre. That in the main
cascade was 3.0 mg/litre. This difference is suprising, because the spring-fed flow over the rock
was expected to have less particulate material. Examination of the filters, showed that diatoms
were the dominant material in the spring-fed flow. Whether this is significant for evolution of
the species which occupy this habitat is not known. The pH was 8.2. Water temperatures
ranged from 17-23° C. The origin of the Lac Vaihiria cascade is not known, but water
temperature was 20° C.
Simulium cataractarum is clearly a torrenticolous species. Schroder (1985) has collected S.
cataractarum larvae from cascades in the Maruia, Taharuu and Tuauru Valleys, and in the
Potiai and Vaihiria Rivers. A particulary important site for this species is the small cascade
west of le Trou du Souffleur. Here, S. cataractarum larvae only are found in a small cascade
above the circum-island road, but immediately above that, where the stream flattens out, only
S. malardei larvae are found. This unique situation may allow determination of habitat
requirements for the two species. Schroder (1985, 1988) notes that larvae of S. cataractarum
have lower selectivity for small algae than do larvae of S. tahitiense.
Larvae of S. cataractarum , along with those of S. neoviceps and some of those of S. oviceps ,
have countercoloured body pigmentation, i.e., with the dorsal surface much darker than the
pale ventral surface. Presumably this is to blend into the dark basaltic rock substrate. Whether
this is an adaptation to avoid predation, or for protection from intense solar radiation is not
known.
Society Islands Black Flies
403
Material Examined. — In addition to types, I have seen the following:
Tiarei , Fareteuira R., Cascades de Faarumai, 17° 32' 10" S. 149° 23' 28" W. 30-vi-80, 14-vii-80, 20-vii-80, 2 1 -vii-80,
02-vii-81, 1 l-ii-83, 16-H-83, D. A. & R.E.G. Craig (larvae, DAC).
Teahupoo, (Tahiti-iti), cascade below Mt. Arope, alt. 30 m. 22-iii-84, P. Schroder (small, early instar larvae, DAC).
Paea, Grotte de Mara, 17° 44' 48" S. 149° 34' 10" W. 00-00-74, D.A. Craig (2 larvae, DAC).
Simulium ( Inseliellum ) mesodontium n. sp.
(Figs. 41,52, 63)
Type.—
Holotype. Larva: mature last instar; as slide mount; label data:- “Holotype”, “Tahiti, Tiarei,
Cascades de Faarumai, /Fareteuira R., 17° 32' 10M S. 149° 23 T 48M W./ 14-vii-80, D.A.
Craig.” (MNHP).
Specific epithet. — Named after the Greek words “mesos” for “middle”, and “odontos” for
“tooth” in reference to the prominent median tooth of the hypostoma.
Diagnosis. —
Larva : hypostoma with sublateral teeth laterad of median tooth small; abdomen covered
with ovoid tubercles.
Description. —
Adult Female. Unknown.
Adult Male. Unknown.
Pupa. Based on pharate material from last instar larva. Thorax: tubercles absent. Gill: length 1.2 mm; branching
pattern normal, dorsal-most filaments equal in length, slightly longer than 0.50 length of longest other filament.
Larva. (Single carcase remaining from karyotyping). Last instar with dark pharate pupal gills; length 4.0 mm. Body:
generally light brownish-grey with slightly lighter intersegmental regions. Head capsule: pattern as for S. oviceps-, sensilla
numerous; cervical sclerites fused to postocciput. Hypostoma (Fig. 41): with two main lateral lobes, separated by large
median tooth and two small sublateral teeth; other sublateral teeth increased smoothly in size out to lateral tooth and
single paralateral tooth; lateral serrations absent; five hypostomal sensilla grouped per side. Postgenal cleft: very small,
V-shaped. Postgenal bridge: 5.0 times longer than cleft. Mandible: apex (Fig. 52) with 13 fine spinous teeth extended to
base of mandibular serration, apical teeth rounded; serration wider at base than high; sensillum small and closely applied
to serration. Abdomen: as for S. oviceps-, posteroventral tubercles absent; posterodorsal cuticle with raised, closely packed,
lightly pigmented, ovoid tubercles, 18 nm long, tubercles evenly distributed over dorsal surface of abdomen, smaller on
prothoracic dorsum, absent from ventral cuticle; and with trichoid sensilla, 22.0 long (Fig. 63). Anal sclerite: with
anterodorsal arms longer than normal; posteroventral arms thin, fused to accessory sclerite, extended around anal proleg,
not fused ventrally. Circlet of hooks: 96 rows of hooks, 15-17 hooks per row. Anal papillae: simple; each with small basal
papilla.
Bionomics. — Gut contents appear normal as for typical filter feeding larvae, so no special
function can be assigned to the aberrant median tooth of the hypostoma.
Phylogenetic relationships. — Head, labral fan structure and body shape of the larva place
this species firmly in the oviceps - group. Karyotypic examination of the specimen provisionally
identified it as S. oviceps (Rothfels, pers. comm. 1984), but morphologically it is clearly a
separate species.
Simulium ( Inseliellum ) neoviceps n. sp.
(Figs. 32, 53, 67, 68)
Simulium “neooviceps”. Craig, 1983: 534.
Types. —
Holotype. Larva: penultimate instar; in alcohol; label data:- “Holotype”, “Tiarei, Fareteuira
R., Cascade de Faarumai,/ 17° 32' 10" S. 149° 24’ 00" W. 18-ii-87”, “Simulium (I.)
neoviceps Craig/ det/ D. A. Craig 1987” (MNHP).
Quaest. Ent., 1987, 23 (3)
404
Craig
Paratypes. Larva: penultimate instar; as slide mount; label data:- “Tahiti, Mataiea, E. Lac
Vaihiria,/ 17° 40’ 58” S. 149° 24' 51” W. alt. 480m. /16-vii-80. D.A. & R.E.G. Craig.”
(MNHP).
Larva: penultimate instar; as slide mount; label data:- “Paea, Grotte de Mara, /17° 44’ 48"
S. 149° 33’ 48” W. /alt. 10m. 1974, D.A. Craig,” (MNHP). Larvae: in alcohol; label data as
for holotype (1, BMNH; 2, BPBM; 2, DAC; 1, MNHP).
Specific epithet. — Named after the Greek word “neos” meaning “new” and “oviceps”, from
S. oviceps, in reference to this species’ similarity to S. oviceps.
Diagnosis. —
Larva : labral fans highly reduced, virtually absent; hypostoma protruding anteriorly; anal
sclerite extended ventrally around posterior proleg.
Recognition. — Although S. neoviceps is quite distinct morphologically and cytologically
from S. oviceps (Rothfels in Craig, 1983) by virtual absence of labral fans, extent of the
posteroventral arms of the anal sclerite around the anal proleg and number of head sensilla (or
hairiness), some larvae presently assigned to S. oviceps have very small labral fans and may be
mistaken for those of S. neoviceps.
Description. —
Adult Female. Unknown.
Adult Male. Unknown.
Pupa. Unknown.
Larva. Similar to S', oviceps, but differing in the following characters:- Head: spot colour pattern negative, but not
distinctly so medially (Figs. 32, 67); cuticle essentially smooth; sensillar number normal. Antenna: much longer than
cephalic fan stem, length 0.18 mm; distal article length 0.07 mm; proximal articles darker, concolourous. Labrum:
cone-shaped; labral fans (Figs. 67, 68) highly reduced with only four longer rays; ca. 0.04 mm in length, plus two very
short rays; microtrichia as long as ray width. Hypostoma: extending anterior of head capsule lateral margins; teeth as for
S. oviceps, six to seven hypostomal sensilla per side, tightly grouped. Mandible: apex (Fig. 53.) with seven spinous teeth
extended to serration; serration small, as high as wide; sensillum separate from serration. Abdomen: shape as for 5.
oviceps', posteroventral tubercles distinct. Anal sclerite: extended completely around anal proleg; fused ventrally. Circlet of
hooks: 86 rows of hooks, 16-18 hooks per row. Anal papillae: simple.
Bionomics. — The habitat for larvae of this species is perhaps the most extreme of all
Tahitian simuliid habitats. In the Cascades de Faarumai, larvae were taken from the right of
the main falls where water was flowing in a thin sheet down vertical bare rock. Larvae probably
do not filter feed, since the labral fans are so small. Browsing feeding behaviour is suggested
from examination of gut contents which showed many pieces of fine inorganic material, and by
the rounded and broken hypostomal teeth in specimens ready to moult. Recently ecdysed
individuals had sharp hypostomal teeth, as do S. oviceps (Craig, 1975a). No other simuliid
species were taken from that particular site. A few meters closer to the main falls where
filamentous algae grew on the rocks, both S. cataractarum and S. oviceps were collected,
similar to the situation at the Lac Vaihiria site.
Phylogenetic relationships. — The morphological convergences in S. neoviceps and S.
oviceps larvae to those of the browsing simuliid Gymnopais is startling. The body shape, to
allow the abdomen to bend so that the mouthparts can be applied to the substrate (Craig,
1977); the reduction of the labral fan with concomitant reduction in size of the labrum (Craig,
1974); the reduction in the size of postgenal cleft, probably for strengthening the head; the
shortening of the mandible and reduction of apical teeth for scraping the substrate; and
flattening of the hypostomal teeth and extension of the hypostoma anteriorly for the “pan and
broom” technique of feeding (Currie and Craig, 1987), have been independently developed in
both groups.
Society Islands Black Flies
405
Material Examined. — In addition to types I have seen the following: Tiarei, Fareteuira R.,
Cascade de Faarumai, 17° 32' 10" S. 149° 23 f 48" W. 16-ii-83, D.A. Craig (SEM specimen,
DAC). Larvae: two destroyed during karyotyping.
Simulium (Inseliellum) oviceps Edwards
(Figs. 8, 15, 20, 25, 34, 43, 55, 66, 70)
Simulium oviceps Edwards 1933: 37, 1935: 37. Lectotype larva (by present designation), TAHITI. Smart, 1945: 511.
Grenier and Rageau, 1960: 734. Grenier and Rageau, 1961b: 174. Dumbleton, 1962: 77. Davies, 1965: 172. Rubtsov,
1974: 242, 244. Craig, 1975a: 466. Craig, 1975b: 299. Crosskey, 1981: 8, 61. Craig, 1983: 533. Schroder, 1985: 17.
Inseliellum oviceps (Edwards). Rubtsov, 1974: 275.
Simulium ( Inseliellum ) oviceps Edwards. Crosskey, 1987: 388.
Types. —
Lectotype. By designation from syntype series. Larva: slide mount; label data:- “625 TYPE
625,/ Simulium oviceps Edw./ Papara River, Tahiti,/ Mumford and Adamson” (BPBM).
Paralectotypes. Adults: pinned; on acetate; label data:- “Paralectotype”, “Syntype”,
“Simulium (I.) oviceps Edw./ det. /D.A. Craig 1986”, “Tahiti, Tautira. /9.viii.l925. /Miss
Cheesman./ B.M. 1925. 392”, “Syntype” (2 2, BMNH). Adult: pinned; on acetate; label data:-
“Paralectotype”, “Syntype”, “Simulium (I.) oviceps Edw./ det. /D.A. Craig 1986”, “Society Is.
Tahiti. /16.3.25 L. E. Cheesman. /B. M. 1925 - 464”, “Syntype” (1 2, BMNH). Adult: pinned;
label data:- “Paralectotype”, “Syntype”, “Simulium (I.) oviceps Edw./ det./ D.A. Craig 1986”,
“Tahiti”, “Society Is. 500’. /Tahiti I.”, “Faraura Val. / 11-17-28”, “Hitiaa 11-17-28”, “5 miles
from sea”, “A.M. Adamson Collector”, “Pacific Entomological Survey”, “Brit. Mus.
1931.234”, “? oviceps” (1 2, BMNH). Adult: pinned; label data:- “Society Is. Tahiti I”,
“Faraura Val. 12-20-28”, “Hitiaa 500”, “5 miles from sea”, “Mumford & Adamson”, “Pacific
Entomological Survey”, “Brit. Mus. 1931-234”, “Syntype”, “Paralectotype”, “Simulium (I.)
oviceps Edw./ det. /D.A. Craig 1986” (12, BMNH). Adult: pinned; label data:- “Society Is. /
1000’ Tahiti I”, “Fautaua Val. / 8-23-28”, “NR VANILLA PLANT”, “SIMULIUM”,
“A.M. Adamson collector”, “Pacific Entomological Survey”, “Brit. Mus. 1931-234”, “Syntype”,
“Paralectotype”, “Simulium (I.) oviceps Edw./ det./ D.A. Craig 1986” (1 2, BMNH). Adult:
pinned; label data:- “Society Is. / 1000' Tahiti I”, “Fautaua Val. / viii-22-28”, “NR
VANILLA PLANT”, “SIMULIUM”, “A.M. Adamson collector”, “Pacific Entomological
Survey”, “Brit. Mus. 1931-234”, “Syntype”, “Paralectotype”, “Simulium (I.) oviceps Edw./
det. /D.A. Craig 1986” (1 2, BMNH). Adult: pinned; label data:- “Society Is. /150m Tahiti I”,
“Papenoo Val /xi-9-28”, “A.M. Adamson Collector”, “Pacific Entomological Survey”, “Brit.
Mus. 1931-234”, “Syntype”, “Paralectotype”, “Simulium (I.) oviceps Edw./ det./ D.A. Craig
1986” (1 2, BMNH). Adult: pinned; label data:- “Society Is. /150m Tahiti I”, “Papenoo Val
/ 10-23-28”, “10 kilo from sea”, “A.M. Adamson Collector”, “Pacific Entomological Survey”,
“Syntype”, “Paralectotype”, “Simulium (I.) oviceps Edw./ det. /D.A. Craig 1986” (1 2,
BMNH). Adult: pinned; label data:- “Society Is. /150m Tahiti I”, “Tipaerui Val /ix- 12-28”,
“3 miles from sea”, “A.M. Adamson Collector”, “Pacific Entomological Survey”, “Syntype”,
“Paralectotype”, “Simulium (I.) oviceps Edw./ det. /D.A. Craig 1986” (1 2, BMNH).
Diagnosis. —
Pupa : dorsal gill filament short or long; thoracic tubercles pointed.
Larva: reduced labral fans; head sensilla numerous; laterally expanded posterior abdomen.
Recognition. — Included for the present in this species are larvae which have even smaller
labral fans with fewer rays. These larvae have the arms of the anal sclerite further extended
around the anal proleg.
Quaest. Ent., 1987,23 (3)
406
Craig
Description. —
Adult Female. Body: colour variable, generally black to light brown; length 1.2-1. 5 mm. Head: black-brown; width
0.36-0.41 mm, depth 0.25-0.36 mm. Eyes: interocular distance 0.12 width of head.; frontal angle 85 degrees; ommatidia
0.013 mm in diameter, ca. 28 and 30 respectively across and up eye in middle row. Vertex: black with vestiture of
concolorous hairs. Frons: black-brown. Clypeus: concolorous with frons, pale ventrally; finely pilose; slight medial
projection. Antenna: length 0.25 mm; evenly dark brown. Mouthparts: length less than 0.50 head depth; mandibles as long
as labrum, ca. 15 small teeth; lacinia with ca. 15 retrorse teeth; palpus, 0.88 mm long, distal article 0.19 mm long,
proximal article concolorous with clypeus, sensory vesicle occupying 0.50 width of third article, opening 0.33 width of
vesicle, ca. 10 sensillar sockets visible. Cibarial pump: space between proximal arms U-shaped, smooth, as wide as deep.
Thorax: length 0.52-0.63 mm; postpronotal lobe with tuft of hairs paler than scutum; scutum evenly dark brown, slightly
shiny, vestiture of short, slightly golden hairs; scutellum apex sharp, posterolateral sides concave, forming generally obtuse
angle, pale to concolorous with scutum, vestiture of sparse black hairs; postscutellum, concolorous with scutellum; pleuron
dark brown. Wing: length 1.1-1. 4 mm, maximum width 0.53-0.58 mm; stem vein hair tuft variable, from sparse to almost
absent; basicostal vein with few hairs. Halter: pale. Legs: generally evenly brown: pretarsal claw smoothly curved with
sharp apex; basal tooth 0.50 length of claw, diverging from claw. Abdomen: light brown to dark brown; basal fringe of
sparse hairs extended to posterior of segment II. Abdomen: tergites I and II as wide as abdomen; tergites 1 1 1— VI narrower
anteriorly, expanded posteriorly, occasionally subshining; pleural regions pilose; sternum slightly mottled; sternites I— IV
not apparent; sternites VI and VII narrow anteriorly, expanded posteriorly. Genitalia (Fig. 8.): cercus in lateral view
slightly curved dorsally and ventrally with sharp apex; hypogynial valves with dark edges medially, pale apically, broadly
rounded posteriorly extended just beyond base of anal lobe; stem of genital fork smooth, long, lateral sclerites triangular
with anterior apex sharply angulate, laterally concave. Spermatheca: ovoid; slight pattern of longitudinal striations; not
heavily pigmented; clear area at junction of sperm duct normal.
Adult Male. Body: generally black; length 1.1-1. 4 mm. Head: width 0.44-0.50 mm, depth 0.34-0.39 mm. Eyes: upper
ommatidia 0.02 mm in diameter, 16 and 14, respectively up and across eye; lower ommatidia, 0.01 mm in diameter. Frons:
shiny black-brown, 0.33 head width. Clypeus: 0.25 as wide as head. Antenna: length 0.24 mm, uniformly dark brown.
Mouthparts: 0.20 length of head depth; mandibles with fine hair-like teeth apically; lacinia lacking retrorse teeth, but with
fine hairs apically; palpus 0.19 mm long, sensory vesicle spherical, occupying only 0.33 of width of third palpal article,
opening less than 0.33 width of vesicle, ca. nine sensillar sockets visible. Thorax: length 0.41-0.55 mm; scutum velvety
black brown, vestiture of short golden hairs; postpronotal lobe lighter with small tuft of hairs; scutellum cone-shaped in
dorsal view, apex pronounced and pale with vestiture of sparse, black hairs; postscutellum dark brown, medial black area;
pleuron brown. Wing: length 1.1-1. 2 mm, maximum width 0.5-0. 6 mm, stem vein hair tuft of few, but substantial hairs.
Halter: greyish. Legs: generally brown, with black vestiture; pedisulcus distinct; pretarsal claw slightly curved; ca. 22
grappling hooks. Abdomen: jet black; vestiture of sparse black hairs; basal fringe of dark hairs extended to segment III.
Genitalia (Fig. 15): gonocoxa 1.0-1. 2 times as long as basal width; gonostylus approximately 2.0 times as long as basal
width, strongly curved with one large blunt terminal spine; ventral plate hairy medially, broadly rounded posteriorly,
slightly concave laterally, anterior sclerotized arms pointed, anteromedian notch very distinct and rounded.
Pupa. Length; 1.3-1. 6 mm male; 1.3-1. 9 mm female. Gill (Fig. 20): length, 0. 8-0.9 mm male, 0.56-1.0 mm female;
branching pattern typical, dorsal filament 0.25-0.75 length of other filament, occasionally as long, or very short, or absent,
variable even on same specimen. Thorax: tubercles pointed (Fig. 70); sternites and tergites as for S. exasperans.
Larva. Last instar with dark pharate pupal gills; length 2. 7-3. 8 mm. Body: greyish-brown with narrow pale anterior
intersegmental regions; ventrally pale, dorsal colour occasionally extended completely around abdominal segment I. Head
capsule (Figs. 34, 66): variable, generally rich brown, but may be pale ventrally; frontoclypeal apotome pale anteriorly and
laterally, posteriorly to stemmata, light brown medially, brown posteriorly; shape, narrow anteriorly, widest opposite
stemmata, narrowing rapidly to parallel sided (variable - may be narrowest just posterior to stemmata, widening slightly
posteriorly); spot pattern negative; stemmata with clear area posterolaterally, occasionally with narrow curved mark
dorsally; cervical sclerites fused to head capsule; sensillae numerous (Fig. 66). Antenna: 3.0 times longer than labral fan
stem, length 0.14 mm, distal article 0.06 mm; antennal phragma extensive and heavily pigmented. Labral fans: reduced;
stem small, but distinct; eight to 12 short rays, ca. 0.17 mm in length; microtrichia of medial rays as long as ray width,
pattern of longer microtrichia interspersed with seven or eight smaller microtrichia. Hypostoma (Fig. 43): all teeth similar
and forming slightly convex line, median tooth protruding slightly; depending on age of larva, wear may be apparent on
sublateral teeth; two paralateral teeth; two or three small lateral serrations; four or five grouped hypostomal sensilla per
side; hypostoma produced anteriorly from head capsule. Postgenal cleft: rounded, as wide as deep, irregular anteriorly.
Postgenal bridge: 3.0 times length of cleft. Mandible: short and curved, apex (Fig. 55) with blunt apical teeth of similar
length (depending on age of larva), seven or eight spinous teeth extended to mandibular serration; serration as high as
basal width, sharply pointed and curved posteriorly; sensillum more than 0.50 height of serration, similarly shaped.
Maxillary palpus: 2.0 times as long as width at base. Mandibular phragma: strongly pigmented, extended ventrad to 0.50
depth of maxilla base. Abdomen: segments I to IV of similar size, segment V increased in size rapidly laterally to
maximum size at segment VI; segments I— IV, constricted at intersegmental region, giving corrugated appearance to
abdomen; posteroventral tubercles prominent; posterodorsal cuticle essentially smooth, with bifurcate sensilla, 8.4 jum long.
Anal sclerite; with posteroventral arms 2.2 times longer than dorsolateral arms, expanded ventrally; separate lateral
accessory sclerites absent. Circlet of hooks; 91 rows of hooks, 13-14 hooks per row. Anal papillae: variable; two or three
basal tubercles on each papilla, ranging in size from small to as long as papilla (Fig. 25).
Society Islands Black Flies
407
Bionomics. — Simulium oviceps is the second most common simuliid on Tahiti, occurring
sympatrically with S. tahitiense in most of its habitats. Simulium oviceps has always
commanded the most interest of all the Tahitian simuliids, because of the highly reduced labral
fans of the larvae (Dumbleton, 1962; Davies, 1974; Craig, 1974; 1975 a, b). I had suggested
(Craig, 1975a), that S. oviceps larvae were browsers because of the particles of volcanic glass
in their guts. That was corroborated by observations on feeding behaviour (Craig, 1977).
Schroder (1985) found a higher proportion of ingested detritus particles in S. oviceps larvae
than in associated S. tahitiense larvae.
In the larger rivers, larvae of S. oviceps tend to be found around the outside of the dense
aggregations of S. tahitiense which are normally found under large rocks in strong flow.
Elsewhere, S. oviceps larvae and pupae can be found in deep depressions in the volcanic
boulders, or on the smooth rock of cascades, or occasionally on filamentous algae. Although
nowhere nearly as abundant as larvae of S. tahitiense , those of S. oviceps can be numerous and
form monospecific populations. At Bain Loti ( 1 9-ii- 1 987), a full range of larval instars and
pupae, were collected from dead leaves, and in particular, from plastic bags (garbage) in the
water. Particulate matter larger than 0.45 pm, in the water at that time, was 8.3 mg/litre. The
pH was 8.3 and the temperature 23° C.
Phylogenetic Relationships. — S. oviceps as presently described is probably a complex of
species. Larvae taken with S. tahitiense from the larger rivers are morphologically very
consistent. However, some larvae from the Cascades de Faarumai and the Vaihiria River
possess labral fans intermediate in size between S. oviceps and S. neoviceps, and the anal
sclerite posteroventral arms extend well around the anal proleg. Further material and
karyological examination of that form is needed.
Material Examined. — In addition to types, I have seen the following:
Adults : pinned:
Tahiti , Faraura Val. 1 1-17-28, 1 1-17-28, 5 miles from sea, A.M. Adamson Collector, Pacific Entomological Survey, Brit.
Mus. 1931.234, Simulium (I) oviceps /det./ D.A. Craig 1986 (1 2, BMNH).
Papeete , Fautaua R., Bain Loti, 17° 33 ' 15" S. 149° 33 ' 00" W. 19-ii-87, D.A. Craig. (2 3, DSIR; 2 3, 1 2, MNHP; 4 <3,
BPBM; 2 3, 2 2, USNM)
Haapape, Tuauru R., 17° 32' 10" S. 149° 29’ 15" W. alt. 50 m. 09-vii-80, D.A. & R.E.G. Craig. Stat. #3 (2 3, BMNH;
2 3, MNHP).
Papenoo, Society Is. 150 m. Tahiti I., Papenoo Val. 10-27-28, 10 kilo, from sea, in cop., A.M. Adamson Collector, Pacific
Entomological Survey, Brit. Mus. 1931.231, ? oviceps, (specimens now cleared and in vial) (1 <3, 1 2, in cop., BMNH).
Papenoo R., 17° 36’ 52" S. 149° 24’ 51" W. 10-vii-80, D.A. & R.E.G. Craig (1 5, BPBM; 1 <3, 1 2, CNCI; 4 3, DSIR; 2
3, UASM).
Tautira, (Tahiti-iti), Vaitepiha R„ 17° 46’ 30" S. 149° 10’ 21" W. alt. 50 m. 19-vii-80, D.A. & R.E.G. Craig. Stat.# 13
(1 2, 1 3, BPBM).
Teahupoo, (Tahiti-iti), Tiirahi R., 17° 51 ’ 05" S. 149° 14’ 15" W. 20-vii-80, D.A. & R.E.G. Craig (2 2, 1 3, DAC; 2 3,
DSIR; 1 3, UASM).
Mataiea, Aug. 1928. A. Tonnoir. BM. 1930.513, ?oviceps (1 3, BMNH). Vaihiria R., 17° 44’ 22" S. 149° 24’ 15" W.
alt. 80m. 16-vii-80, D.A. & R.E.G. Craig (1 3, CNCI).
Punaauia, Punaruu R., 17° 38’ 20" S. 149° 35’ 05" W. alt. 120m. 07-vii-80, D.A. & R.E.G. Craig (1 2, cleared in vial,
BMNH; 1 2, MNHP; 1 2, UASM).
Alcohol material:-
Papeete, Fautaua R., Bain Loti, 17° 33’ 15" S. 149° 33* 00" W. 08-vii-74 (larvae, DSIR), 15-vii-74 (larvae, pupa,
BMNH); 16-vii-74 (larvae, BMNH); 08-vii-80, (larvae CNCI; larvae, pupae, USNM), 01-vii-81 (larvae, pupa, CNCI;
larvae, DAC), D.A. & R.E.G. Craig.
Haapape, Tuauru R„ 17° 32’ 10" S. 149° 29’ 15" W. alt. 50m. 09-vii-80, D.A. & R.E.G. Craig (pupa, DAC; larvae,
MNHP).
Papenoo, Papenoo R., 17° 36’ 52" S. 149° 24’ 51" W. 10-vii-80, D.A. & R.E.G. Craig (larvae, CNCI).
Tiarei, Fareteuira R., Cascades de Faarumai, 17° 32’ 10" S. 149° 24’ 00" W. 16-ii-83, (larvae, MNHP); 17-ii-87
(larvae, DAC) D.A. Craig .
Mahaena, Mahape R„ 17° 33’ 00" S. 149° 20’ 33" W. 14-vii-80, alt. 700m. D.A. & R.E.G. Craig (karyotyped larvae,
BPBM; larvae, DAC).
Quaest. Ent., 1987, 23 (3)
408
Craig
Tautira, (Tahiti-iti), Vaitepiha R„ 17° 46' 30" S. 149° 10’ 21" W. alt. 50m. 19-vii-80, D A. & R.E.G. Craig (larvae,
CNCI; larvae, MNHP).
Teahupoo, (Tahiti-iti), Tiirahi R„ 17° 51' 05" S. 149° 14’ 15" W. 20-vii-80, D.A. & R.E.G. Craig (larvae, pupae,
BMNH).
Vairoa, (Tahiti-iti), Vavi R., 22-ii-84, P. Schroder (pupa, MNHP).
Mataiea, Vaihiria R., 17° 44' 22" S. 149° 24' 15" W. alt. 80m. 16-vii-80, D.A. & R.E.G. Craig (larvae, DAC); Vaihiria
R., 17° 41 ' 31" S. 149° 25' 02" W. alt. 250m. 17-vii-80, D.A. & R.E.G. Craig (larva, DAC). E. Lac Vaihiria, 17° 40'
58" S. 149° 24' 51" W. alt. 480m. 22-vii-80, D.A. Craig, (pupa, DAC).
Mahaiatea, Taharuu R., 17° 42' 45" S. 149° 29' 15" W. alt. 135m. 15-vii-80, D.A. & R.E.G. Craig (pupae, DSIR).
Paea, Public Gardens, 03-vii-81, D.A. Craig (larvae, DAC).
Punaauia, Punaruu R., 17° 38' 20" S. 149° 35' 05" W. 27-vi-69, B. Hocking (larvae, UASM); 03-V-73 (larvae, pupae,
DAC); 05-vii-73 (larvae, UASM), 6-vii-73 (larvae, MNHP), 04-vii-74 (larvae, DAC), 05-vii-74 (larvae, DAC), 08-vii-74
(larvae, DSIR), 18-vii-74 (larvae, CNCI), 07-vii-80 (larvae, DAC), D.A. & R.E.G. Craig.
Simulium (Inseliellum) sp.
(Fig. 17)
Description. —
Adult Female. Unknown.
Adult Male. (Based on pharate material). Head: width 0.6 mm, depth 0.5 mm. Eyes: upper, larger ommatidia 0.04
mm, 12 rows both up and across eye, lower, smaller ommatidia 0.01 mm. Mouthparts: less than 0.50 of head depth; lacinia
with hairs apically; palpal sensory vesicle spherical, 0.50 width of article, opening 0.33 width of vesicle, four or five
sensillar sockets visible. Legs: segments with pigmented distal regions; pretarsal claw with 17-20 grappling hooks.
Genitalia (Fig. 17): gonocoxa with pronounced posteromedial projection; gonostylus broad, roundly truncated distally,
single prominent apical spine; ventral plate hairy and broadly cone-shaped posteromedially, rounded laterally, anterior
arms heavily pigmented, anteromedian notch broad.
Pupa. Thorax: tubercles pointed. Gill: length 1.2 mm; as for S', oviceps ; dorsal-most filament 0.33 length of other
filaments.
Larva. Unknown.
Phylogenetic relationships. — This adult has been placed in the oviceps- group on the basis
of pupal characteristics, namely the short dorsal gill filament and the pointed thoracic
tubercles, both characteristic of S. oviceps. However, the genitalia (Fig. 17) with unique large
posteromedian projection on the gonocoxa, show that this male is not of S. oviceps. Because this
specimen may belong to one of the oviceps-group of species that are based only on larvae, it is
not given specific status at this time.
Material Examined. — Pupa: pharate adult; as slide mount; label data: Simulium (I.) sp.,
TAHITI, Mataiea, E. Lac Vaihiria, 17° 40’ 58” S. 149° 24’ 51" W. alt. 480 m. 22-vii-80,
D.A. Craig. Stat. # 16 (MNHP).
CONCLUDING REMARKS
I estimate that only 75% of simuliid species in Tahiti have been described, based on the rate
at which species were discovered once it was realized that the new species were from specialized
habitats; which had not been well collected. Further, morphological variation indicates that
species such as S. oviceps and perhaps S. neoviceps are probably complexes of species.
When description of the Marquesas (Sechan in Klein et al., 1983) and Huahine (Sechan
pers. comm.) simuliids are complete, preferably with karyological data, it should be possible to
attempt a cladistic analysis for Inseliellum. This could be particularly valuable, because it
would include almost a total fauna which, to judge from the ages of the islands involved
(Duncan and McDougall, 1974; 1976), and from karyological and morphological data, is
monophyletic. If so, it should then prove possible to test the zoogeographic hypotheses of Craig
(1983) and to make predictions regarding discovery of simuliids on other Polynesian islands.
Society Islands Black Flies
409
Such an analysis will require establishing the sister taxon to Inseliellum for outgroup
comparison. That may prove difficult because of the homogeneous morphological nature of
possible candidate subgenera.
Bionomically, Polynesian simuliids are poorly known, but Schroder (1985) has made a
valuable start with his studies on feeding behaviour and distribution of Tahitian simuliids. This
provides a basis for investigation of resource partitioning by the larvae in relation to their
morphological and presumably behavioural characteristics. Of particular interest, but probably
very difficult to do, will be examination of the reproductive behaviour which keeps the
sympatric species from hybridizing, and the mechanisms by which the very rare species manage
to locate mates.
Marquet and Lamarque (1987) report that there are 17 species of fresh water fish in Tahiti
and Moorea, and that waterfalls have a marked effect on their distribution. In the larger rivers
on Tahiti fish are very common and noticeable (pers. obs.). Fish are known predators of black
fly larvae and these can be an important food item (Davies, 1981; Allen, 1982). Further, there
are suggestions (Peckarsky, 1984) that such prey organisms may form large groups to reduce
chances of predation per individual.
At the Cascades de Faarumai, in particular at the Haamaremarerahi cascade, where the
wet rocks directly entered the pool at the base of the cascade, small gobiid sucker-fish
(Sicyopterus taeniurus ) were common on the rock faces. These fish could move with startling
agility up and across the wet rock. Upon disturbance, the fish would either swim rapidly up the
rock, or leap off into the pool below. Noticeable was that in these situations the rock was bare.
Where there was a rubble base to the rock face and the fish did not have direct access to a pool
beneath, they did not occur and algae and macrophytes grew in the water flowing over the rock.
Black fly larvae were found on the vegetation and patches of bare rock. A search of the bare
rock faces above the pool failed to reveal larvae of S. neoviceps or, indeed, of any black fly.
However, analysis of gut contents of six of the gobiids produced five larval carcasses of the
short labral fan-form of S. oviceps. The remainder of the gut contents was algae, of which a
high proportion was diatomaceous, and small pieces of volcanic glass. The gobiids are
apparently browsers, ingesting the black fly larvae incidentally along with the epilithic algae.
This observation, in addition to those on distribution of S. tahitiense and S. oviceps larvae in
the larger rivers, where larvae respectively, either form large aggregations, or are in depression
in the rocks, suggests that fish feeding behaviour has an effect on distribution of Tahitian
Simuliidae. This hypothesis could perhaps be tested by examining the distribution and
aggregation behaviour of larvae of black flies species above and below waterfalls high enough to
effect distribution of fish.
ACKNOWLEDGMENTS
I thank the Institut Territorial de Recherches Medicales Louis Malarde for providing
laboratory space, transportation and technical assistance over the years, and personnel of the
Institut and of Office de la Recherche Scientifique et Technique Outre-mer, for help in
collecting material. Also to my good friend D. Huston who provided accommodation and moral
support while I was in Papeete.
The British Museum (Natural History) loaned available Edwards syntypes. A large series of
pinned material was made available by the Bishop Museum, Honolulu and also the United
States National Museum, Washington, D.C. F. Rodhain, Institut Pasteur, Paris, loaned the S.
Quaest. Ent., 1987,23 (3)
410
Craig
lotii material described by Grenier and Rageau (1960). I extend thanks to D. Hollingdale for
her usual high quality line drawings, and for being such a delight to work with. J. S. Scott and
G. D. Braybrook provided technical assistance with scanning electron microscopy and
photography.
Particular thanks are extended to R. W. Crosskey for a great deal of help, and to G.
Marquet and J. Nelson for identifying the fish.
K. Rothfels was always pleased to receive larval simuliids from such places as Tahiti and he
provided me with invaluable karyological information. His sudden death in October. 1986 will
make work such as this much more difficult. I am pleased to dedicate this paper to his memory.
Financial assistance was provided by Natural Sciences and Engineering Research Council,
Canada, grant No. A5753 and by a grant from the Central Research Fund, University of
Alberta. I thank B.K. Mitchell, Chairman, Department of Entomology, for supplementary
funds for the illustrative work.
REFERENCES
Allen, J.D. 1982. The effects of reduction in trout density on the invertebrate community of a
mountain stream. Ecology 63: 1444-1455.
Bedo, D.G. 1977. Cytogenetics and evolution of Simulium ornatipes Skuse (Diptera:
Simuliidae) I. Sibling speciation. Chromosoma (Berlin) 64: 37-65.
Bedo, D.G. 1984. Sibling species and sex chromosome differentiation in Simulium neornatipes
(Diptera: Simuliidae). Canadian Journal of Genetics and Cytology 26: 318-325.
Cheesman, E. 1932. Hunting insects in the South Seas. Philip Allen, London. 243 pp.
Craig, D.A. 1974. The labrum and cephalic fans of larval Simuliidae (Diptera: Nematocera).
Canadian Journal of Zoology 52: 133-159.
Craig, D.A. 1975a. The larvae of Tahitian Simuliidae (Diptera: Nematocera). Journal of
Medical Entomology 12: 463-476.
Craig, D.A. 1975b. Origin and evolution of the Simuliidae (Diptera) of Tahiti. Abstracts, pp.
299-300. 13th Pacific Science Congress (1975) 1: 358 pp.
Craig, D.A. 1977. Mouthparts and feeding behaviour of Tahitian larval Simuliidae (Diptera:
Nematocera). Quaestiones Entomologicae 13: 195-218.
Craig, D.A. 1983. Phylogenetic problems in Polynesian Simuliidae (Diptera: Culicomorpha): A
progress report. GeoJournal 7: 533-541.
Craig, D.A. and A. Borkent. 1980. Intra- and inter-familial homologies of maxillary palpal
sensilla of larval Simuliidae (Diptera: Culicomorpha). Canadian Journal of Zoology 58:
2264-2279.
Craig, D.A. and R.E.G. Craig. 1986. Simuliidae (Diptera: Culicomorpha) of Rarotonga, Cook
Islands, South Pacific. New Zealand Journal of Zoology 13: 357-366.
Crosskey, R.W. 1967. The classification of Simulium Latreille (Diptera: Simuliidae) from
Australia, New Guinea and the western Pacific. Journal of Natural History 1: 23-51.
Crosskey, R.W. 1974. The identity and taxonomic status of the man-biting black-fly of Fiji
(Simuliidae: Diptera). Journal of Entomology (B) 43: 149-158.
Crosskey, R.W. 1981. Geographical distribution of Simuliidae. pp. 57-68. In Laird, M.
(Editor). Blackflies: the future for biological methods in integrated control. Academic Press,
London. 399 pp.
Crosskey, R.W. 1987. Annotated checklist of the world black flies (Diptera: Simuliidae). pp.
Society Islands Black Flies
411
358-440. In Kim, K.C. and R.W. Merritt (Editors). Black flies: ecology, population
management and annotated world list. Pennsylvania State University Press, University
Park. 500 pp.
Currie, D.C. 1986. An annotated list of and keys to the immature Black Flies of Alberta
(Diptera: Simuliidae). Memoirs of the Entomological Society of Canada 134: 1- 90.
Currie, D.C. and D.A. Craig. 1987. Larval feeding strategies, pp. 136-148. In Kim, K.C. and
R.W. Merritt (Editors). Black Flies: ecology, population management and annotated world
list. Pennsylvania State University Press, University Park. 500 pp.
Davies, D. M. 1981. Predators upon blackflies. pp. 139-159. In Laird, M. (Editor). Blackflies:
the future for biological methods in integrated control. Academic Press, Toronto. 399 pp.
Davies, L. 1965. The structure of certain atypical Simuliidae (Diptera) in relation to evolution
within the family, and the erection of a new genus for the Crozet Island black-fly.
Proceedings of the Linnean Society of London 176: 159-180.
Davies, L. 1974. Evolution of larval head-fans in Simuliidae (Diptera) as inferred from the
structure and biology of Crozetia crozetensis (Womersley) compared with other genera.
Zoological Journal of the Linnean Society 55: 193-224.
Doutch, H.F. 1981. Southwest Quadrant, Plate-Tectonic Map of the Circum-Pacific Region.
Circum-Pacific Council for Energy and Mineral Resources. American Association of
Petroleum Geologists, Tulsa, Oklahoma.
Dumbleton, L. J. 1962. Aberrant head-structure in larval Simuliidae (Diptera). Pacific Insects
4: 77-86.
Dumbleton, L. J. 1973a. Simulium laciniatum Edwards - a valid species (Simuliidae:
Diptera). Journal of the Royal Society of New Zealand 3: 453-456.
Dumbleton, L. J. 1973b. The genus Austrosimulium Tonnoir (Diptera: Simuliidae) with
particular reference to the New Zealand fauna. New Zealand Journal of Science 15 (1972):
480-584.
Duncan, R.A. and I. McDougall. 1974. Migration of volcanism with time in the Marquesas
Islands, French Polynesia. Earth and Planetary Science Letters 12: 414-420.
Duncan, R.A. and I. McDougall. 1976. Linear volcanism in French Polynesia. Journal of
Volcanology and Geothermal Research 1: 197-227.
Edwards, F.W. 1927. Diptera Nematocera from the South Pacific collected by the ‘St. George4
Expedition, 1925. The Annals and Magazine of Natural History 20(9): 236-244.
Edwards, F.W. 1932. Marquesan Simuliidae. Bulletin of the Bernice P. Bishop Museum 98:
103-109.
Edwards, F.W. 1933. Tahitian Simuliidae. Pacific Entomological Survey Publication 6: 35-38.
Edwards, F.W. 1935. Society Island insects. Tahitian Simuliidae. Bulletin of the Bernice P.
Bishop Museum 113: 35-38.
Freeman, P and B. De Mellion. 1953. Simuliidae of the Ethiopian Region. British Museum
(Natural History). 224pp.
Grenier, P. and J. Rageau. 1960. Simulies (Dipt., Simuliidae) de Tahiti. Remarques sur la
classification des Simuliidae. Bulletin de la Societe de Pathologie Exotique 4: 727-742.
Grenier, P. and J. Rageau. 1961a. Un nouveau genre de Simulies: Hebridosimulium N.G.
(Diptera, Simuliidae) des Nouvelles-Hebrides. Description des stades preimaginaux et du
male, redescription de la femelle de H. jolyi (Roubaud), 1906. Bulletin de la Societe de
Pathologie exotique 54: 95-102.
Grenier, P. and J. Rageau. 1961b. Simulies (Dipt. Simuliidae) de Tahiti. Remarques sur la
Quaest. Ent., 1987, 23 (3)
412
Craig
classification des Simuliidae. pp. 174-177. Verhandlungen XI Internationalen Kongress
Entomologie, Wien (1960). 803 pp.
Holloway, J. D. 1979. A survey of the Lepidoptera, biogeography and ecology of New
Caledonia. W. Junk. The Hague. 588 pp.
Klein, J. M. F. Riviere and Y. Sechan. 1983. Recherches d’entomologie medicale aux lies
Marqueses en 1982. Notes et documents d’hygiene et de Sante Publique Entomologie
Medicale, Office de la Recherche Scientifique et Technique Outre-mer, Papeete, Tahiti. 7:
1-81.
Lacey, L. A. and J. M. Lacey. 1983. Filter feeding of Simulium fulvinotum (Diptera:
Simuliidae) in the Central Amazon Basin. Quaestiones Entomologicae 19: 41-51.
Loti, P. 1887. Le mariage de Loti. Calmann-Levy, Paris. 313 pp.
McLea, M.C. and D.M. Lambert. 1983. Cytogenetics of New Zealand blackflies of the genus
Austrosimulium (Diptera: Simuliidae) 1. The cytogenetics of Austrosimulium australense.
New Zealand Journal of Zoology 10: 271-280.
Marquet, G. and P. Lamarque. 1987. Prospecting the fauna in the freshwaters of Tahiti and
Moorea (French Polynesia), p. 86. In Abstracts. XXII Congress of the International
Association of Limnology, Hamilton, New Zealand. 142 pp.
O’Hara, J. E. 1983. Classification, phylogeny and zoogeography of the North American species
of Siphona Meigen (Diptera: Tachinidae). Quaestiones Entomologicae 18: 261-380.
Peckarsky, B.L. 1982. Predator-prey interactions among aquatic insects, pp. 196-254. In Resh,
V.H. and D.M. Rosenberg (Editors). The ecology of aquatic insects. Praeger Scientific,
Toronto. 625 pp.
Roubaud, M.E. 1906. Sur deux types interessant de Simuliides de l’Afrique equatoriale et des
Nouvelle-Hebrides. Bulletin du Museum d’Histoire Naturelle. Paris. 12: 140-143.
Rubtsov, I.A. 1956. Simuliidae. Fauna SSSR No. 23, (6). Moscow. 532 pp.
Rubtsov, I.A. 1974. Evolution, phylogeny and classification of the family Simuliidae (Diptera).
Trudy Zoologischeskogo Instituta. Leningrad 53: 230-281. (In Russian).
Schroder, P. 1985. Feeding biology of Tahitian Blackfly larvae (Dipt. Simuliidae). Notes et
Documents, Hygiene et Sante Publique Entomologie Medicale, Office de la Recherche
Scientifique et Technique Outre-mer, Papeete, Tahiti. 1 1: 1-230.
Schroder, P. 1988. Gut-passage, particle selection and ingestion of filter-feeding blackfly
(Dipt., Simuliidae) larvae inhabiting a waterfall in Tahiti (French-Polynesia). Aquatic
Insects. (In Press).
Smart, J. 1945. The Classification of the Simuliidae (Diptera). Transactions of the Royal
Entomological Society of London 95: 463-532.
Smart, J. and E.A. Clifford. 1965. Simuliidae (Diptera) of the territory of Papua and New
Guinea. Pacific Insects 7: 505-619.
Society Islands Black Flies
413
Simulium cheesmanae
Figures 1-4. Female genitalia, ventral views. Fig. 1. Simulium anatolicum n. sp. Right side as true ventral view. Left side
with hypogynial valve transparent to show basal connection to anal lobe. Fig. 2. S. castaneum n. sp. Fig. 3. S.
cataractarum n. sp. Fig. 4. S', cheesmanae Edwards. Scale line = 0.1 mm; an lb = anal lobe; cere = cercus; gen fk =
genital fork; hyp valv = hypogynial valve; spmth = spermatheca.
Quaest. Ent., 1987,23 (3)
414
Craig
Simulium tahitiense
Figures 5-9. Female genitalia, ventral view. Fig. 5. Simulium exasperans n. sp. Fig. 6. S. lotii n. sp. Fig. 7. S.
opunohuense n. sp. Fig. 8. S. oviceps Edwards. Fig. 9. S. tahitiense Edwards. Scale line = 0.1 mm.
Society Islands Black Flies
415
Simulium cheesmanae
Simulium
exasperans
Simulium tahitiense Simulium sp
Figures 10-17. Male genitalia, ventral views. Fig. 10. Simulium cataractarum n. sp. Fig. 1 1. S', cheesmanae Edwards. Fig.
12. S. exasperans n. sp. Fig. 13. S. lotii n. sp. Fig. 14. S. malardei n. sp. Fig. 15. S. oviceps Edwards. Fig. 16. S. tahitiense
Edwards. Fig. 17. S. sp. Scale lines = 0.1 mm; goncx = gonocoxa; gonst = gonostylus; v pit = ventral plate.
Quaest. Ent., 1987, 23 (3)
416
Craig
i i
Simulium anatolicum
19
Simulium exasperans
Simulium tahitiense
Figures 18-21. Pupae, right lateral views. Fig. 18. Simulium anatolicum n. sp. Fig. 19. S. exasperans n. sp. Fig. 20. S.
oviceps Edwards. Fig. 21. S. tahitiense Edwards. Scale lines = 1.0 mm.
Society Islands Black Flies
417
i i
Simulium malardei
22
Simulium oviceps
Figures 22 and 23. Left lateral views of larval posterior abdomen. Fig. 22. Simulium malardei n. sp. Fig. 23. S. tahitiense
Edwards. Figures 24 and 25. Posterior views of larval abdomen. Fig. 24. Simulium cataractarum n. sp. Fig. 25. S. oviceps
Edwards. Scale lines = 1 .0 mm; acc scl = accessory sclerite; an ppl = anal papillae; an scl = anal sclerite; d tub = dorsal
tubercle; ex an scl = ventral extension of anal sclerite; v tub = ventral tubercle.
Quaest. Ent., 1987, 23 (3)
418
Craig
Simulium admixtum
Simulium arlecchinum
i l
Simulium castaneum
Simulium cataractarum
Figures 26-29. Dorsal and ventral views of head capsule (labral fan rays omitted). Fig. 26. Simulium admixtum n. sp.
Fig. 27. S. arlecchinum n. sp.. Fig. 28. 51. castaneum n. sp. Fig. 29. S. cataractarum n. sp. Scale lines = 0.25 mm.
Society Islands Black Flies
419
Simulium lotii
Simulium neoviceps
33
i j
Simulium opunoliuense
Figures 30-33. Dorsal and ventral views of head capsule (labral fan rays omitted). Fig. 30. Simulium lotii n. sp. Fig. 31.
S. malardei n. sp. Fig. 32. S. neoviceps n. sp. Fig. 33. S. opunohuense n. sp. Scale lines = 0.25 mm.
Quaest. Ent., 1987, 23 (3)
420
Craig
Simulium oviceps
i i
Simulium tahitiense
Figures 34 and 35. Dorsal and ventral views of head capsule (labral fan rays omitted). Fig. 34. Simulium oviceps Edwards.
Fig. 35. S. tahitiense Edwards. Scale lines = 0.25 mm.
Society Islands Black Flies
421
40
Simulium malardei
Figures 36-40. Ventral views of larval hypostoma. Fig. 36. Simulium admixtum n. sp. Fig. 37. S. castaneum n. sp. Fig.
38. S. cataractarum n. sp. Fig. 39. S. lotii n. sp. Fig. 40. S. malardei n. sp. Scale lines = 0.1 mm; 1 sr = lateral serration; 1
tth = lateral tooth; m tth = median tooth; pi tth = paralateral teeth; si tth = sublateral teeth.
Quaest. Ent., 1987,23 (3)
422
Craig
44
Simulium tahitiense
Figures 41-44. Ventral views of larval hypostoma. Fig. 41. Simulium mesodontium n. sp. Fig. 42. S. opunohuense n. sp.
Fig. 43. S. oviceps Edwards. Fig. 44. S. tahitiense Edwards. Scale lines = 0.1 mm; 1 tth = lateral tooth; m tth = median
tooth.
Society Islands Black Flies
423
Simulium arlecchinum
Figures 45-50. Adoral view of left, larval mandibles. Fig. 45. S. admixtum n. sp. Fig. 46. S. arlecchinum n. sp. Fig. 47. S.
castaneum n. sp. Fig. 48. 5. cataractarum n. sp. Fig. 49. S. exasperans n. sp. Fig. 50. S. lotii n. sp. Scale lines = 0.025
mm; mnd s = mandibular serration; s = sensillum; sp tth = spinose teeth.
Quaest. Ent., 1987, 23 (3)
424
Craig
Simulium mesodontium
Simulium opunohuense
Simulium oviceps
Figutes 51-56. Adoral view of left, larval mandibles. Fig. 51. Simulium malardei n. sp. Fig. 52. S. mesodontium n. sp.
Fig. 53. S. neoviceps n. sp. Fig. 54. S. opunohuense n. sp. Fig. 55. S. oviceps Edwards. Fig. 56. S. tahitiense Edwards.
Scale lines = 0.025 mm.
Society Islands Black Flies
425
Fig. 57. S. castaneum n. sp. Mandibular serration (mnd s), showing basal sensillum (s). Scanning electron micrograph
(SEM). Fig. 58. S. castaneum n. sp. Microtrichial pattern on labral fan ray (SEM). Fig. 59. S. malardei n. sp. As for Fig.
58 (SEM). Fig. 60. S. castaneum n. sp. Larval head cuticle, showing corrugations and raised sensillar sockets (SEM). Fig.
61. S. castaneum n. sp. Posterodorsal abdominal cuticle (SEM). Fig. 62. S. malardei n. sp. As for Fig. 61 (SEM). Fig. 63.
S. mesodontium n. sp. Photomicrograph of posterodorsal abdominal cuticle. Scale lines =10 /am.
Quaest. Ent., 1987,23 (3)
426
Craig
Figures 64-67. Dorsal views of last instar larval heads (SEM). Fig 64. S. cataractarum n. sp. Fig. 65. S. tahitiense
Edwards. Fig. 66. S. oviceps Edwards. Fig. 67. 5. neoviceps n. sp. Scale lines = 100 /m i.
I
Society Islands Black Flies
427
Fig. 68. Right labral fan of S. neoviceps n. sp. (SEM). Fig. 69. S. anatolicum n. sp. Pupal thoracic tubercles
(Photomicrograph). Fig. 70. S. oviceps Edwards. Pupal thoracic tubercles (SEM). Fig. 71. S', tahitiense Edwards. Pupal
thoracic tubercles (SEM). Fig. 72. S. tahitiense Edwards. Pharate pupal thoracic tubercles (SEM). Fig. 73. S. tahitiense
Edwards. Pharate pupal thoracic tubercles (Phase contrast photomicrograph). Fig. 74. S. exasperans n. sp. Pupal thoracic
cuticle (SEM). Fig. 75. S. lotii n. sp. Pharate pupal thoracic tubercles (SEM). Scale lines = 20 /am; ant = antenna; If =
labral fan; mnd = mandible; mxp = maxillary palpus.
Quaest. Ent., 1987, 23 (3)
428
Craig
admixtum n. sp., Simulium, 377-381,
397,418,421,423
Allen, J.D., 409
anatolicum n. sp., Simulium , 377,
379-380,383,413,416, 427
Apulvillus cheesmanae Edwards, 396
arlecchinum n. sp., Simulium , 377-381,
397-398,418,423
Bedo, D.G., 374
Borkent, A.,
see Craig, D.A., 376
bussoni Edwards, Simulium , 375
castaneum n. sp., Simulium , 377, 397,
399,413,418,421,423,425
cataractarum n. sp., Simulium , 375, 377,
379-381, 397-402, 404, 413, 415,
417-418,421,423,426
Cheesman, E., 386
cheesmanae Edwards, Apulvillus , 396
cheesmanae Edwards, Simulium , 374,
377-380,385-386,413,415
Clifford, E.A.,
see Smart, J., 373
Craig, D.A., , 374-375, 383-384,
387-388, 390, 394, 397, 401, 404,
407-408
see also Currie, D.C., 404
Craig, R.E.G.,
see Craig, D.A., 383-384, 387-388
Crosskey, R.W., 373, 381, 383-384
Currie, D.C., 376, 404
Davies, D.M., 409
Davies, L., 407
de Mellion, B.,
see Freeman, P., 384
Doutch, H.F, 383
Dumbleton, L.J., 373, 407
Duncan, R.A., 408
Edwards, F.W., 374-375, 384, 386
Eusimulium Roubards, 384
exasperans n. sp., Simulium, 374-375,
377, 379-381, 389-390, 392-397, 406,
414-416, 423,427
Freeman, P, 384
Gomphostibia Enderlein, 384
INDEX
Grenier, P., 374, 383-384, 386, 390
Hebridosimulium Grenier and Rageau,
381-383
Holloway, J.D., 374
Inseliellum oviceps (Edwards), 405
Inseliellum Rubtsov, 379, 383-385,
408-409
jolyi Roubaud, Simulium, 381, 383
Klein, J.M., 384, 388, 408
laciniatum Edwards, Simulium, 383
Lamarque, P.,
see Marquet, G., 409
Lambert, D.M.,
see McLea, M.C., 373
lotii n. sp., Simulium, 374, 376-377,
379-381, 383, 386, 388,' 390, 392-394,
396, 409, 414-415, 419, 421, 423, 427
malardei n. sp., Simulium, 374, 376-381,
387, 402, 415, 417, 419, 421, 424-425
Marquet, G., 409
McDougall, I,
see Duncan, R.A., 408
McLea, M.C., 373
mesodontium n. sp., Simulium, 377-381,
397, 403, 422, 424-425
neoviceps n. sp., Simulium, 377-381,
397-398, 402-404, 407-409, 419, 424,
426-427
opunohuense n. sp., Simulium, 377, 388,
392,414,419,422, 424
oviceps (Edwards), Inseliellum, 405
oviceps Edwards, Simulium, 373-377,
379-381, 383-384, 390, 394, 397-398,
402-405, 407-409, 414-417, 420, 422,
424, 426-427
Pomeroyellum Rubstov, 384
Rageau, J.,
see Grenier, P., 374, 383-384, 386,
390
Riviere, F.,
see Klein, J.M., 384, 388, 408
Rubtsov, I.A., 383-384
Schroder, P., 375, 384, 390, 392, 396, 402,
407, 409
Society Islands Black Flies
429
Sechan, Y,
see Klein, J.M., 384, 388, 408
Sicyopterus taeniurus, 409
Simuliidae, 374, 376, 378-380, 384, 409
Simulium admixtum n. sp., 377-381, 397,
418,421,423
Simulium anatolicum n. sp., 377,
379-380,383,413,416, 427
Simulium arlecchinum n. sp., 377-381,
397-398,418,423
Simulium bussoni Edwards, 375
Simulium castaneum n. sp., 377, 397, 399,
413,418,421,423,425
Simulium cataractarum n. sp., 375, 377,
379-381, 397-402, 404, 413, 415,
417-418,421,423,426
Simulium cheesmanae Edwards, 374,
377-380,385-386,413,415
Simulium exasperans n. sp., 374-375,
377. 379- 381, 389-390, 392-397, 406,
414-416, 423,427
Simulium jolyi Roubaud, 381, 383
Simulium laciniatum Edwards, 383
Simulium lotii n. sp., 374, 376-377,
379-381, 383, 386, 388, 390, 392-394,
396, 409, 414-415, 419, 421, 423, 427
Simulium malardei n. sp., 374, 376-381,
387, 402, 415, 417, 419, 421, 424-425
Simulium mesodontium n. sp., 377-381,
397, 403, 422, 424-425
Simulium neoviceps n. sp., 377-381,
397-398, 402-404, 407-409, 419, 424,
426-427
Simulium opunohuense n. sp., 377, 388,
392,414,419, 422, 424
Simulium oviceps Edwards, 373-377,
379-381, 383-384, 390, 394, 397-398,
402-405, 407-409, 414-417, 420, 422,
424, 426-427
Simulium tahitiense Edwards, 373-375,
377. 379- 381, 383-384, 386, 390-397,
399, 402, 407, 409, 414-417, 420, 422,
424, 426-427
Simulium teruamanga Craig and Craig,
387-388
Smart, J., 373
tahitiense Edwards, Simulium , 373-375,
377, 379-381, 383-384, 386, 390-397,
399, 402, 407, 409, 414-417, 420, 422,
424, 426-427
teruamanga Craig and Craig, Simulium ,
387-388
Quaest. Ent., 1987,23 (3)
/
Quaestiones
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 23
NUMBER 4
FALL 1987
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Issued February 1988
QUAESTIONES ENTOMOLOGICAE
ISSN 0033-5037
A periodical record of entomological investigation published at the Department of
Entomology, University of Alberta, Edmonton, Alberta.
Volume 23 Number 4 1987
CONTENTS
Anderson-Systematics, phylogeny and biogeography of New World weevils traditionally of the
tribe Cleonini (Coleoptera: Curculionidae; Cleoninae) 431
Book Notices 711
Editor’s Acknowledgements and Comments 713
t
)
SYSTEMATICS, PHYLOGENY AND BIOGEOGRAPHY OF NEW WORLD WEEVILS
TRADITIONALLY OF THE TRIBE CLEONINI (COLEOPTERA: CURCULIONIDAE;
CLEONINAE)
Robert Samuel Anderson
Department of Entomology
University of Alberta
Edmonton, Alberta, T6G 2E3 Quaestiones Entomologicae
CANADA 23:431-709 1987
ABSTRACT
A survey of representatives of generic-group names in Cleoninae and consideration of their
phylogenetic relationships indicates four genera are represented in the New World. These are:
Cleonis Dejean ( introduced ; 1 species ); Stephanocleonus Motschulsky (Holarctic, northern; 6
species including S. confusus n. sp. [type locality. Black Hills, South Dakota], S. immaculatus
n. sp. [type locality, Fort McMurray, Alberta], S. parshus n. sp. [type locality, Chalk River,
Ontario], and S. stenothorax n. sp. [type locality, Bluefish Caves, Yukon Territory]); Apleurus
Chevrolat (southwestern North America, arid lands; 8 species ); and, Cleonidius Casey ( eastern
and western North America, semi-arid and arid lands; 19 species including C. eustictorrhinus
n.sp. [type locality, Sacramento, California] , C. infrequens n. sp. [type locality, Scott City,
Kansas], and, C. notolomus n. sp. [type locality, Grant County, New Mexico]). Cleonidius is
hypothesized to be phylogenetically more closely related to traditional Lixini rather than
Cleonini and present tribal classification of Cleoninae is regarded as unsatisfactory. The
following new generic-group synonymies are presented: Lixus Fabricius ('=Epimeces Billberg
[type species Curculio filiformis Fabricius, here designated ]); Stephanocleonus Motschulsky
(^Coniocleonus Motschulsky); Apleurus Chevrolat (=Centrocleonus LeConte, Cleonopsis
LeConte, Cleonaspis LeConte, and Dinocleus Casey); and, Cleonidius Casey (=Lixestus
Reitter).
The following information is given in detail for each of Cleonis, Stephanocleonus, Apleurus
(Gibbostethus) n. subg. ( type species, Dinocleus hystrix Fall by monotypy), Apleurus
(Apleurus), and Cleonidius: synonymic list, notes about synonymy, diagnosis, description, list
of included species, and a discussion of phylogenetic relationships. Keys to species of North
American Stephanocleonus and Apleurus, and world Cleonidius, are presented. The following
information is given in detail for each species considered: synonymic list, notes about
synonymy, problems in recognition, description, distribution, natural history information, and
chorological relationships.
Lixus mixtus LeConte is a junior homonym and is replaced with Cleonidius longinasus, new
name. The following new species-group synonymies are presented: Apleurus lutulentus
(LeConte) (=Cleonus pulvereus LeConte, Apleurus fossus Chevrolat, and Dinocleus bryanti
Van Dyke); Apleurus porosus (LeConte)(= Dinocleus farctus Casey); Apleurus angularis
(LeConte) ( = Dinocleus denticollis Casey and Dinocleus porcatus Casey); Apleurus jacobinus
(Casey) (=Centrocleonus pilosus LeConte and Cleonus (Dinocleus) capillosus Csiki); Apleurus
Present address: Department of Entomology, Texas A & M University, College Station, TX 77843, U. S. A.
432
Anderson
albovestitus (Casey) (= Cleonus molitor LeConte, Dinocleus densus Casey, Dinocleus wickhami
Casey, Dinocleus interruptus Casey, Dinocleus mexicanus Casey, and Cleonus (Dinocleus)
structor Csiki); Apleurus saginatus (Casey) (= Dinocleus dentatus Champion ); Cleonidius
subcylindricus Casey (= Cleonus (Cleonidius) graniferus Casey); Cleonidius texanus (LeConte)
(= Cleonus (Cleonidius) circumductus Casey); Cleonidius frontalis (LeConte) (= Cleonus
sparsus LeConte and Cleonus (Cleonidius) stratus Csiki); Cleonidius poricollis (Mannerheim)
(= Cleonus (Cleonidius) lobigerinus Casey and Cleonus (Cleonidius) kirbyi Casey); Cleonidius
boucardi (Chevrolat) (= Cleonus carinicollis LeConte and Cleonus (Cleonidius) lecontellus
Csiki); and, Cleonidius trivittatus (Say) (= Cleonus inornatus LeConte and Cleonus
(Cleonidius) bicarinatus Casey). Lixus californicus Motschulsky is considered a nomen
dubium.
Only for species of Apleurus and Cleonidius are reconstructed phytogenies presented based
on analyses of 31 characters in Apleurus and 22 characters in Cleonidius. Apleurus is
hypothesized to be the monophyletic sister genus of the Palearctic Chromoderus. In Apleurus,
two monophyletic subgenera are recognized. These are Gibbostethus (1 species, A. hystrix) and
Apleurus s. s. (7 species). Within Apleurus s. s. included species and hypothesized phylogenetic
relationships are as follows: ((((A. albovestitus + A. saginatus) + ((A. jacobinus + A.
angularis) + A. porosus)) + A. lutulentus) + A. aztecus). Cleonidius is monophyletic and
hypothesized to comprise an unresolved trichotomy with the Palearctic Lixus (Lixoglyptus)
and Cylindropterus. Four monophyletic species groups are recognized in Cleonidius. Along
with constituent species and hypothesized phylogenetic relationships, these are: C. erysimi
group, (((C. texanus + C. longinasus) + C. subcylindricus) + C. pleuralis + (C. erysimi + C.
eustictorrhinus)); C. americanus group, ((((C. puberulus + C. infrequens) + (C. collaris + C.
notolomus)) + C. canescens) + (C. americanus + C. frontalis)); C. poricollis group, (C.
poricollis + C. calandroides); and C. boucardi group, (((C. placidus + C. quadrilineatus) + C.
trivittatus) + C. boucardi). Phylogenetic relationships of groups are (((C. boucardi group +
C. poricollis group) + C. americanus group) + C. erysimi group). Phylogenetic relationships
of the Nearctic species component of Cleonidius are unresolved but the latter is likely sister to
the Palearctic species component.
Both Apleurus and Cleonidius are hypothesized to have entered North America in savanna
or grassland habitats (or their precursors) across Beringia during late Eocene time. In North
America, primary diversification has taken place in the arid lands of the southwestern United
States although two species of Cleonidius are found in coastal regions of eastern North
America. In both genera there appears to be an initial vicariance of lineages, on Compositae,
in grassland or mesquite-grassland habitats between California, central Mexico, and the
southern Rocky Mountains. Further speciation events in Apleurus appear due primarily to
shifts to new, apotypic habitat types at differing altitudes brought about by increased aridity
during the later Tertiary; host plant shifts are not a factor because Apleurus species are all
associated with Compositae. Host plant shifts, however, primarily between
Leguminosae-Rosaceae and likely also Compositae-Cruciferae, and within-habitat-type
allopatric speciation appear to predominate as the causes of further speciation in Cleonidius,
most species of which remain associated with plesiotypic less-arid habitat types. Habitat
shifts promoting speciation in Cleonidius are evident, but are not as associated with formation
of increasingly arid habitats as in Apleurus.
Evolutionary trends in Apleurus and Cleonidius are discussed and patterns and predictions
of biogeographic or evolutionary interest summarized and presented for examination for
New World Cleonini
433
general applicability.
RESUME
Une diagnose des characlres de la sous-famille est aussi brievement discutee. L’examen de representants des
groupes-genres de Cleoninae et leurs relations phyletiques indiquent que quatre genres sont representes dans le
Nouveau-Monde. Ils sont: Cleonis Dejean (introduit; 1 esp'ece); Stephanocleonus Motschulsky (Holoartique, nordique; 6
especes incluant S. confusus n. esp. [localite-type, Black Hills, Dakota du SudJ, S. immaculatus n. esp. [localite-type,
Fort McMurray, Alberta], S. parshus n. esp. [localite-type, Chalk River, Ontario], et S. stenothorax n. esp. [localite-type,
Bluefish Caves, Territoire du Yukon]; Apleurus Chevrolat (sud-ouest de VAmerique du Nord, terres arides; 8 especes); et
Cleonidius Casey (est et ouest de VAmerique du Nord, terres arides et semi-arides; 19 especes incluant C. eustictorrhinus
n. esp [localite-type, Sacramento, Califournie] , C. infrequens n. esp. [localite-type, Scott City, Kansas], et C. notolomus
n. esp. [localite-type. Grant County, Nouveau Mexique]. Cleonidius est suggere comme etant phyletiquement plus
apparente aux traditionels Lixini plutot qu’aux Cleonini, et la classification actuelle de Cleoninae est consideree comme
etant insatisfaisante. De nouvelles synonymies de groupe-genres sont presentee: Lixus Fabricius (= Epimeces Billberg
[espece-type Curculio filiformis Fabricius, ici designee]); Stephanocleonus Motschulsky (^Coniocleonus Motschulsky);
Apleurus Chevrolat (=Centrocleonus LeConte, Cleonopsis LeConte, Cleonaspis LeConte, et Dinocleus Casey); et
Cleonidius Casey ('=Lixestus Reitter).
L’information suivante est presentee en detail pour chacun des Cleonis, Stephanocleonus, Apleurus (Gibbostethus)
nouveau sous-genre ( espece-type , Dinocleus hystrix Fall par monotypie), Apleurus (Apleurus), et Cleonidius: liste
synonymique et notes les concernant, diagnose, description, liste des especes incluses, et une discussion concernant les
relations phyletiques. Des clefs d’identification a Vespece pour les Stephanocleonus et Apleurus de I’Amerique du Nord,
et les Cleonidius Mondiaux sont produites. L’information suivante est aussi presentee en detail pour chacune des especes
considerees: liste synonymique et notes les concernant, problemes associes d l’ identification, description, distribution,
histoire naturelle, et relations chorologiques.
Lixus mixtus LeConte est considere comme homonyme plus recent et est remplace par Cleonidius longinasus, nouveau
nom. Les nouvelles synonymies suivantes des groupes-especes sont presentees: Apleurus lutulentus (LeConte), (=Cleonis
pulvereus LeConte, Apleurus fossus Chevrolat, et Dinocleus bryanti Van Dyke); Apleurus porosus (LeConte) (= Dinocleus
farctus Casey); Apleurus angularis (LeConte) (— Dinocleus denticollis Casey et Dinocleus porcatus Casey); Apleurus
jacobinus (Casey) (=Centrocleonus pilosus LeConte et Cleonus (Dinocleus) capillosus Csiki); Apleurus albovestitus
(Casey) (= Cleonus molitor LeConte, Dinocleus densus Casey, Dinocleus wickhami Casey, Dinocleus interruptus Casey,
Dinocleus mexicanus Casey, et Cleonus (Dinocleus) structor Csiki); Apleurus saginatus (Casey) (= Dinocleus dentatus
Champion); Cleonidius subcylindricus Casey (=Cleonus (Cleonidius) graniferus Casey); Cleonidius texanus (LeConte)
(= Cleonus (Cleonidius) circumductus Casey); Cleonidius frontalis (LeConte) (= Cleonus sparsus LeConte et Cleonus
(Cleonidius) stratus Csiki); Cleonidius poricollis (Mannerheim) (= Cleonus (Cleonidius) lobigerinus Casey et Cleonus
(Cleonidius) kirbyi Casey); Cleonidius boucardi (Chevrolat) ('=Cleonus carinicollis LeConte et Cleonus (Cleonidius)
lecontellus Csiki); et Cleonidius trivittatus (Say) (= Cleonus inornatus LeConte et Cleonus (Cleonidius) bicarinatus
Casey). Lixus californicus Motschulsky est consideree nomen dubium.
Des reconstitutions phyletiques sont presentees pour les especes r/’Apleurus et Cleonidius uniquement, et sont basees
sur I’analyse de 31 characteres pour Apleurus et 22 characteres pour Cleonidius. Apleurus est suggere comme etant le
genre-soeur monophyletique du Chromoderus paleartique. A I’interieur J’Alleurus, deux sous-genres monophyletiques
sont reconnus. Ils sont Gibbostethus (1 espece, A. hystrix) et Apleurus s. s. (7 especes). A I’interieur ^/’Apleurus s. s. les
especes incluses ainsi que les relations phyletiques suggerees sont les suivantes: ((((A. albovestitus + A. saginatus + ((A.
jacobinus + A. angularis) + A. porosus)) + A. lutulentus) + A. aztecus). Cleonidius est monophyletique et est propose
comme englobant une trichotomie non-resolue avec le Lixus (Lixoglypus) paleartique et Cylindropterus. Quatre
groupe-especes monophyletiques sont reconnus pour Cleonidius. De conserve avec les especes constituantes et les relations
phyletiques suggerees, ils sont: le groupe C. erysimi, (((C. texanus + C. longinasus) + C. subcylindricus) + C. pleuralis
+ (C. erysimi + C. eustictorrhinus)); le groupe C. americanus, ((((C. puberulus + C. infrequens) + (C. collaris + C.
notolomus)) + C. canescens) + (C. americanus + C. frontalis)); le groupe C. poricollis, (C. poricollis + C. calandroides);
et le groupe C. boucardi, (((C. placidus + C. quadrilineatus) + C. trivattatus) + C. boucardi). Les relations phyletiques
des groupes sont: (((C. boucardi groupe + C. poricollis groupe + C. americanus groupe + C. erysimi groupe. Les
relations phyletiques entre les especes de Cleonidius de la composante neartique sont non-resolues, cependent cette
derniere est vraisemblablement soeur de la composante paleartique.
L’hypothese gu’Apleurus et Cleonidius atteignirent I’Amerique du Nord via Beringia, aux travers des savanes ou des
prairies (ou de leurs precurseurs ) durant I’Eocene superieur, est avancee. En Amerique du Nord, une diversification
initiate aurait pris place dans les terres arides du sud-ouest des Etats-Unis, bien que deux especes de Cleonidius soient
retrouvees dans les regions cotieres de Vest de VAmerique du Nord. II semble, qu ’a Vinterieur de ces deux genres, les
lignees demontrent une vicariance initiate, sur Compositae, en prairies avec ou sans mesquites entre la Califournie, le
Mexique Central, et le sud des Montagnes Rocheuses. Plus ample evenements de speciation chez Apleurus sont apparus
dues principalement aux deplacements vers de nouveaux habitats apotypiques-types aux differantes altitudes, induits
par Varidite accrue raignant au Tertiaire superieur; le deplacement vers de nouvelles plantes-hotes nest pas un facteur
Quaest. Ent., 1987,23 (4)
434
Anderson
puisque toutes les especes J’Apleurus sont associees aux Compositae. Cependent, le deplacement vers de nouvelles
plantes-hotes, principalement entre Leguminosae-Rosacae et vraisemblablement Compositae-Cruciferae, de-meme
qu’une speciation intra-habitats de type allopatrique semblent dominer en tant qu’instigateurs de plus ample speciations
chez Cleonidius, la majorite de ces dernier demeurant associes h la formation d’habitats plesiotypiques moins arides. Les
deplacement vers de nouveaux habitats encourageant la speciation chez Cleonidius sont evidents, mais ne sont pas aussi
fortement associes 'a V apparition d’habitats de plus en plus arides comme chez Apleurus.
Les tendances evolutionaires chez Apleurus et Cleonidius sont discutees et les patrons de distribution ainsi que des
predictions d’interets biogeographiques ou evolutionaires sont resumes et presentes pour examen de leur applicability
globale.
TABLE OF CONTENTS
Introduction 434
Materials and methods 435
Natural history of Cleoninae 445
Supraspecific taxa and included species 446
Subfamily Cleoninae 446
Key to genera of new world Cleoninae 447
Genus Cleonis Dejean 449
Genus Stephanocleonus Motschulsky 453
Key to species of adult Stephanocleonus 455
Genus Apleurus Chevrolat 467
Subgenus Gibbostethus Anderson 468
Subgenus Apleurus Chevrolat 470
Key to species of adult Apleurus 472
Genus Cleonidius Casey 492
Key to species of adult Cleonidius 496
Evolutionary analysis: reconstructed phylogeny 547
Evolutionary analysis: biogeography 574
Evolutionary trends and adaptations 590
Conclusions and future research 594
Acknowledgements 595
Literature cited 595
Figures 607
Appendix 1: Tables 1-33 670
Appendix 2 703
Index 705
INTRODUCTION
J.B.S. Haldane, in an oft repeated quote, once stated that the creator must have held a
special fondness for beetles for he made so many of them. To this I would hasten to add that he
must also have had a special predilection towards weevils, for the Curculionidae are the largest
single family of organisms known. More than 44883 species had been described as of 1971
(O’Brien and Wibmer, 1978) and countless others no doubt await discovery and formal
scientific description. In so far as numerous species await description, some clearer
understanding has yet to be achieved of those taxa already formally recognized. Crowson
(1955), commenting on this very problem, noted that “A satisfactory resolution of the
Curculionidae into subfamilies and tribes is probably the largest and most important
New World Cleonini
435
outstanding problem in the higher classification of Coleoptera particularly as a great number of
the species are more or less seriously injurious to economically valuable crops.”
But to begin to make progress towards achieving this goal, we must start at some lower level,
come to some understanding of a subfamily, tribe or genus, and then proceed to the larger and
more complex problem. This I have chosen to do by conducting a study of the North American
species that have traditionally been placed in the tribe Cleonini of the Cleoninae. The group
had not been revised since LeConte and Horn (1876) and subsequently Casey (1891), and has
been recently widely recognized by curculionid systematists as problematical and in need of
work. Not only were species very difficult to reliably identify, but classification of the North
American fauna in a world perspective had not been attempted. Use of the group in applied
problems ( e.g as biological control agents of pest weeds), or to address theoretical problems of
general evolutionary or biogeographic importance, was not yet possible.
This study will alleviate those problems of identification and provide more detailed .and
reliable definitions of species and species-group taxa than those previously presented. It will
also address the classification of the North American fauna in a world perspective and
represent a start at a consistent scheme for use by scientists in all faunal regions. Phylogenetic
and biogeographic patterns can then be recognized and resultant process-hypotheses
formulated so as to be available for general consideration and applicability by students from a
wide array of disciplines, not just those interested in Curculionoidea.
MATERIALS AND METHODS
Materials
This study was based on examination of 9498 adult specimens of Cleonidius (3586);
Apleurus (5627); North American Stephanocleonus (224), and Cleonis pigra (61), borrowed
from various collections, both institutional and private, and including material I personally
collected in the western United States and Canada. The following codens represent these
collections, both individual and institutional, and their respective curators.
AMNH American Museum of Natural History, Central Park West at 79th Street,
New York, New York, 10024, U.S.A.; L.H. Herman.
ASUT Arizona State University, Tempe, Arizona, 85281, U.S.A.; F.F.
Hasbrouck.
BMNH British Museum (Natural History), Cromwell Road, London, SW7 5BD,
England; R.T. Thompson and C.H.C. Lyal.
CASC California Academy of Sciences, Department of Entomology, Golden Gate
Park, San Francisco, California, 94118, U.S.A.; D.H. Kavanaugh.
CDFA California Department of Food and Agriculture, 1220 N Street,
Sacramento, California, 95814, U.S.A.; T.N. Seeno.
CNCI Canadian National Collection of Insects, Arachnids and Nematodes,
Biosystematics Research Centre, Research Branch, Ottawa, Ontario, K1A
0C6, Canada; D.E. Bright.
CUIC Cornell University Insect Collections, Department of Entomology, Cornell
University, Ithaca, New York, 14853, U.S.A.; J.K. Liebherr.
CWOB Charles W. O’Brien Collection, Department of Entomology, Florida
Agricultural and Mechanical University, Tallahassee, Florida, 32307,
U.S.A.; C.W. O’Brien.
Quaest. Ent., 1987, 23 (4)
436 Anderson
DLCC Douglas L. Caldwell Collection, Chemical Lawn R & D Center, P.O. Box
395, Milford Center, Ohio, 43045, U.S.A.; D.L. Caldwell.
ELSC Elbert L. Sleeper Collection, Department of Biology, California State
University at Long Beach, Long Beach, California, 90801, U.S.A.; E.L.
Sleeper.
FMNH Field Museum of Natural History, Roosevelt Road and Lake Shore Drive,
Chicago, Illinois, 60605, U.S.A.; L.E. Watrous.
HAHC Henry F. and Anne T. Howden Collection, Department of Biology,
Carleton University, Ottawa, Ontario, K1S 5B6, Canada; A.T. Howden.
JLCC John L. and Alberta F. Carr Collection, 23 Dalrymple Green NW,
Calgary, Alberta, T3A 1Y2, Canada; J.L. Carr.
JVMC John V. Matthews Collection, Energy, Mines, and Resources Canada, 601
Booth Street, Ottawa, Ontario, K1 A 0E8, Canada; J.V. Matthews.
LACM Los Angeles County Museum of Natural History, 900 Exposition
Boulevard, Los Angeles, California, 90007, U.S.A.; R.R. Snelling.
MCZC Museum of Comparative Zoology, Harvard University, Cambridge,
MAssachusetts, 02138, U.S.A.; A.F. Newton, Jr.
MNHP Museum National d‘Histoire Naturelle, Entomologie, 45 Rue Buffon,
Paris, France; N. Berti.
MSUC Montana State University, Department of Biology , Bozeman, Montana,
59717, U.S.A.;S. Rose.
NMSU New Mexico State University, Department of Biology, Las Cruces, New
Mexico, 88003, U.S.A.; J.R. Zimmerman.
NRS Naturhistoriska Riksmuseet, Sektionen for entomologi, S-104 05,
Stockholm, Sweden; P.I. Persson.
NSDA Nevada State Department of Agriculture, 350 Capitol Hill Avenue, P.O.
Box 1 1 100, Reno, Nevada, 89510, U.S.A.; R.C. Bechtel.
OSDA Oregon State Department of Agriculture, 635 Capitol Street N.E., Salem,
Oregon, 97310, U.S.A.; R.L. Westcott.
OSUC Oregon State University, Department of Entomology, Corvallis, Oregon,
97331, U.S.A.;G.L. Peters.
PUL Purdue University, Department of Entomology, West Lafayette, Indiana,
47907, U.S.A.; A. Provonsha.
RSAN denotes specimens in my own personal collection.
SBMN Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road,
Santa Barbara, California, 93105, U.S.A.; S. Miller and L. Marx.
SMC Scott McCleve Collection, 2210 13th Street, Douglas, Arizona, 85607,
U.S.A.; S. McCleve.
TAMU Texas Agricultural and Mechanical University, Department of
Entomology, College Station, Texas, 77843, U.S.A.; H.R. Burke.
UASM University of Alberta, Strickland Museum, Department of Entomology,
Edmonton, Alberta, T6G 2E3, Canada; G.E. Ball and D. Shpeley.
UAT University of Arizona, Department of Entomology, Tucson, Arizona,
85721, U.S.A.; F.G. Werner.
UCBC University of California, Division of Entomology and Parasitology,
Berkeley, California, 94720, U.S.A.; J.A. Chemsak.
New World Cleonini
437
UCMC University of Colorado Museum, Campus Box 218, Boulder, Colorado;
U.N. Lanham.
UIM University of Idaho, Department of Entomology, Moscow, Idaho, 83843,
U.S.A.; W.F. Barr.
UNSM University of Nebraska State Museum, Research and Systematics
Collections, W-436 Nebraska Hall, Lincoln, Nebraska, 68588, U.S.A.;
B.C. Ratcliffe.
USNM United States National Museum (Natural History), Washington D.C.,
20560, U.S.A.; D.R. Whitehead.
WECC Wayne E. Clark Collection, Department of Entomology, Auburn
University, Auburn, Alabama, 36849, U.S.A.; W.E. Clark.
WSU Washington State University, Department of Entomology, Pullman,
Washington, 99164, U.S.A.; R.S. Zack.
ZMMU Zoological Museum of the Moscow Lomonosov State University, Herzen
Street 6, Moscow K-9, U.S.S.R.; N. Nikitsky.
Classification of the North American fauna in a world perspective required knowledge of
structural diversity of taxa from other regions of the world. In view of this, representatives of
type-species (where designated) of 50 of a total of 77 described genus-group names of the tribe
Cleonini (not including New World taxa or junior homonyms where replacement names are
available) from the British Museum (Natural History) were studied. In instances where a
type-species has not been designated for a given genus-group name, representatives of a species
initially placed in that taxon (and thus available for subsequent designation as type-species)
were selected for study. These taxa are listed in Appendix II.
Methods
Collection of specimens. — Most specimens of Cleoninae collected by me were acquired
simply by placing a standard entomological beating sheet underneath foliage of a particular
plant and then striking the plant sharply with a sturdy stick in a downward motion over the
sheet. Weevils then on the plant feign death and drop downward onto the white sheet where
they are easily seen and captured. Specimens were killed in vials containing fine ethyl
acetate-soaked wood chips. A few specimens were also collected by sweeping low vegetation or
were handpicked from bare soil. Deciduous shrubs in oak-pinyon-juniper woodlands and annual
or perennial woody shrubs along dry stream beds, desert washes, or various lotic waterways
were the most productive places to concentrate collecting efforts.
Specimen preparation and examination. — A Wild M5® stereo dissecting microscope was
used for routine examination of external structural features of dried adult specimens.
Dissections were made of mouthparts and internal genitalia and these were examined with both
a Wild M5® dissecting microscope and a Leitz SM-Lux® compound brightfield microscope.
Special preparation procedures for specific characters examined in detail are given following.
For such characters, note is made in the text of the number of specimens that were examined in
detail.
For each species, a variable number of male and female individuals were selected from
various localities for study of internal genitalic structure. Individuals of both sexes were first
softened by immersion in hot distilled water, to which a few drops of liquid soap had been
added. Dissection consisted of insertion of the apices of a pair of fine forceps into the sternal
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438
Anderson
suture separating the base of the abdomen from the metasternum. Careful application of
pressure resulted in separation of the abdomen which was then placed into hot 5-10% KOH
(aqueous) for a variable amount of time to remove soft tissues. Repeated examinations were
made of the abdomen during this process to determine the suitable degree of treatment. The
abdomen was then transferred to a 5-10% acetic acid-alcohol solution to neutralize KOH
activity, and then to distilled water. Genitalia were then removed from the abdomen and placed
in 95% ethanol and then into glycerine for subsequent examination. The abdomen was dried
and mounted on a card placed on the specimen pin.
No special techniques were used in the examination of female genitalia; however, for the
internal sac of the aedeagus of males to be adequately examined, it was necessary that it be
everted and inflated. This was best accomplished using an unpublished technique devised by
Mr. Richard T. Thompson of the British Museum (Natural History).
Most practical for large specimens, this technique consists of eversion of the internal sac
following softening and treatment with KOH by careful insertion of a hooked minuten pin
mounted on the end of a wooden probe into the apex of the aedeagus. Slow and numerous
repeated tugs on the sac, which is visible through the wall of the aedeagus, worked best and
could be continued until the sac was completely everted, the apical sclerite complex exposed
and apical. Attempts to extract the sac rapidly and using only one or a few insertions of the
minuten pin frequently damaged the sac such that inflation was not possible. Following
eversion, the aedeagus was placed in a liquid (water was used most frequently but a liquid of
higher viscosity, such as glycerine, is recommended). The base of the aedeagus was placed at
the end of a narrow piece of flexible rubber tubing and the rubber adjacent to the base of the
aedeagus pinched with a pair of flat-tipped forceps placed firmly in a hole in a rubber stopper
attached to a sturdy metal stand. A continuous force was maintained on the forceps by placing
a small elastic band around the forceps near their apices such that they needed to be forced
open using another pair of forceps. It was important that the aedeagus was held firmly in place
in the apex of the tubing, but that the base of the aedeagus was not sealed by the forceps. The
other end of the piece of tubing was attached to a syringe filled with the liquid in which the
aedeagus was immersed. The syringe was also placed in a hole in a rubber stopper attached to a
separate sturdy metal stand. Gentle pressure on the syringe forced the liquid out the apex of the
tubing but ideally only through the aedeagus and thus the internal sac. As liquid passed into the
sac it became inflated and remained so for short periods of time while the pressure exerted by
the syringe continued to exceed the force limiting inflation of the sac. Drawings were thus made
of the sac by repeated application of slight pressure on the syringe. Various views of the sac
were obtained by manipulation of the aedeagus in the apex of the tubing and repetition of entire
procedure. Proper placement of the base of the aedeagus in the tubing was often difficult
because more often than not, pressure on the syringe did not inflate the sac because either the
forceps were sealing the base of the aedeagus or, too much liquid was passing out the tubing
adjacent to the base of the aedeagus and not through it. In this latter instance the aedeagus was
expelled from the apex of the tubing because it was not held firmly in place by the pressure
exerted onto the tubing by the forceps. Repeated attempts and manipulation of the location of
pressure of the forceps were required for good results.
Following examination, internal genitalia were placed in glycerine in a microvial attached to
the specimen pin. To examine mouthparts, heads were removed from individuals softened as
noted above, and placed in 5-10% KOH (aqueous) for a variable amount of time as required
for removal of soft tissues. The head was then transferred to a 10% hydrogen peroxide-95%
New World Cleonini
439
ethanol solution to bleach cuticle to facilitate examination of surface features with a brightfield
microscope. Following removal to a 95% ethanol-distilled water solution, maxillary palpi and
the prementum were carefully removed from the head by severing connections to the apex of
the rostrum with a hooked minuten pin mounted on a wooden probe. These structures were then
transferred to glycerine on a depression slide for detailed examination using a brightfield
microscope. The head was air dried and mounted on a card placed on the specimen pin.
Following examination, mouthparts were placed in glycerine in a microvial attached to the
specimen pin. Detailed mouthpart examination was only carried out for species of Apleurus.
Initial surveys of diversity of mouthpart structure in Cleonidius and Stephanocleonus species
did not prove promising and were discontinued.
Structural terms. — Structural terms follow those of Kissinger (1970) except as noted in
Anderson (1984b). Structure of the internal sac of the aedeagus of males in Cleoninae has not
been previously studied and an informal nomenclature has been developed (Fig. 115). Future
detailed studies on other Cleoninae should result in a more formal nomenclature based on
apparent homology between structures in a more comprehensive survey of structural diversity.
Ambiguity in descriptors used in association with various structural terms is a problem
worthy of comment. Objectively delimited character states ( e.g ., presence/absence, red/black,
three setae/one seta, etc.) are ideal for taxonomic and phylogenetic purposes because there is
no ambiguity in assigning a state to a particular taxon. Most character states however, are
subjectively expressed (e.g., elongate-narrow/robust, large/small, deep/shallow, etc.). For
these latter characters to be maximally useful in a taxonomic sense as criteria for identification
or otherwise, ambiguity must be reduced in assigning a state to a particular taxon without
requiring reference to other taxa. I have attempted to do this for those character states that are
taxonomically important in this study but which are not augmented in the text in a quantitative
manner or accompanied with figures. Characters and their states in which there is little
ambiguity involved are not considered.
Vestiture. I term all body vestiture of apparent non-sensory primary function as scales. Body
vestiture is important both phylogenetically and taxonomically. Pronotal scale patterns serve to
distinguish genera, elytral scale patterns species of Stephanocleonus , and species groups and
species of Cleonidius. Abdominal scale patterns are important in separating some species of
Apleurus.
Scales vary from flat or appressed, to recumbent, sub-erect and erect. I use the terms flat or
appressed for scales at an angle of 0°-5° relative to body surface; recumbent, 6°-45°; sub-erect,
46°-85°; and erect, 86°-90°.
Shape and size of the scale is important in determining macroscopic patterns of surface
vestiture. Small, fine scales obscure less of the underlying dark cuticle and thus the ground
color of the cuticle predominates in the pattern; larger, robust scales obscure more cuticle and
impart a predominantly whitish color to the pattern. Density of the scales is not as important as
size and shape in determining the macroscopic pattern.
More elongate, sub-erect or erect scales are termed “hair-like scales”, a name that would
correspond to most concepts of “hair”. Length of “hair-like scales” is important both
taxonomically and phylogenetically. Members of some species clearly lack sub-erect or erect
surface vestiture; members of most species have short sub-erect or erect vestiture that is not
more than approximately twice the length of the largest appressed scales on that particular
body part. Members of few species have moderately long sub-erect or erect vestiture between
twice and three times the length of the largest appressed scale. Finally, members of few species
have long vestiture, greater than three times the length of the largest appressed scale. Sub-erect
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440
Anderson
or erect vestiture is abraded to various degrees on some specimens causing problems in
identification.
Density of all vestiture and size of scales are indicated qualitatively with reference to other
taxa.
Punctation. Punctation is important both phylogenetically and taxonomically. Three aspects
of punctation are described: size, density and depth. States of all three aspects are part of a
continuum from small, sparse or shallow, to large, dense and deep.
Density of punctation is defined on the basis of the relationship between diameter of a
puncture and the distance between it and adjacent punctures. Sparse punctures are those with
diameters less than the distance between punctures; moderately dense are those with diameters
more or less equal to the distance between punctures; and dense are those with diameters
exceeding the distance between punctures.
Size and depth of punctation are more difficult to assess in an objective manner. Shallow
punctures are those with the walls of the puncture at about a 45° angle or less; deep are those
with walls between 45° and 90°. Moderately deep are those with the state difficult to assign as
either deep or shallow. Small punctures are those where a very rough estimate of the number of
punctures, if lined side by side spanning that body part, would number more than 50; large are
those where the number is less than 25. Estimates between 25 and 50 are considered
moderately large.
Punctation of the head, rostrum and pronotum are most important and most frequently
referred to in keys and descriptions.
Illustrations. — With the specimen placed in glycerine, all drawings of genitalia and
mouthparts were made on paper with the aid of a camera lucida attached to Wild M5® or Leitz
SM-Lux® microscopes at various magnifications and later traced, then inked, onto mylar
drafting film for reproduction. Drawings of other structural features were similarly made of
dried pinned specimens with the aid of a camera lucida attached to a Wild M5® microscope.
Details of surface vestiture and sculpture, unless taxonomically or phylogenetically valuable,
are not shown on illustrations. Homologous structures are drawn to the same scale (in most
instances) and orientation to facilitate comparisons. Only taxonomically or phylogenetically
valuable structures are illustrated; thus, comparisons of certain attributes of the structures for
the purposes of identification may be misleading. Such attributes although illustrated for
aesthetic completeness are not discussed in the text.
Distribution maps are presented for all species with the exception of C. vibex. Special maps
and illustrations were prepared to illustrate general ecological and historical biogeographic
patterns, evolutionary trends or adaptations, etc., or as accompanying visual simplifications of
various discussions in the text.
Hubbs-Hubbs diagrams representing intraspecific and interspecific variation in certain
mensural attributes were prepared separately for each sex. Range, mean, 1.5 standard
deviations (SD) on either side of the mean, and two standard errors (SE) on either side of the
mean are plotted. Choice of 1.5 SD and 2 SE follows Whitehead (1971) as noted in the section
on descriptive statistics.
Features illustrated are as follows: LEI, WRA/LR, WRA/WF, WE1M/LE1, LP/LE1 for
Stephanocleonus (Figs. 49-53) and Cleonidius (Figs. 197-201); and, LEI, WRA/LR,
WRA/WF, WE1M/LE1, LP/LE1, WPT/WPB for Apleurus (Figs. 105-110). Features
illustrated were selected on the basis of their taxonomic and/or phylogenetic value.
Hubbs-Hubbs diagrams were also prepared to illustrate the altitudinal distributions of
New World Cleonini
441
Apleurus (Fig. 232) and Cleonidius (Fig. 233) species. Samples used for preparation of
Hubbs-Hubbs diagrams minimally include those samples used in preparation of descriptive
statistics tables. In those instances where measurements were taken of individuals from more
than the sample used in the descriptive statistics tables, these other samples constituted part of
an examination for possible intraspecific variation in mensural features.
Measurements and descriptive statistics.-— Measurements were made on pinned specimens
at various magnifications with the aid of an ocular micrometer in a Wild M5® microscope.
Measurements selected for detailed study and for presentation were those that upon initial
survey revealed differences between species of potential taxonomic or phylogenetic value. For
this reason not all measurements or ratios presented were taken on species of all genera.
These measurements and their abbreviations are:
LR, length of rostrum measured in lateral aspect from anterior margin of eye to point of
articulation of mandible; WF, minimum width of frons between eyes in dorsal aspect; WRA,
maximum width of rostrum measured across apex in dorsal aspect; LP, length of pronotum
measured along midline from anterior to posterior margins in dorsal aspect; WPB, width of
pronotum measured across base in dorsal aspect; WPT, width of pronotum measured across
apical one-third or one-quarter at tubercles (if present) or immediately posterior to sub-apical
constriction in dorsal aspect; WE1H, width of elytra measured across base at humeri with elytra
contiguous throughout length at suture in dorsal aspect; WE1M, width of elytra measured
across midlength with elytra contiguous throughout length at suture in dorsal aspect; LEI,
length of elytra measured along suture from apex of scutellum to apex of elytra in dorsal
aspect; LEy, maximum length of eye measured in lateral aspect; WEy, maximum width of eye
measured in lateral aspect.
Some of these measurements were used to obtain ratios which serve to express the form of a
particular part of the body. These ratios are; WEy/LEy, shape of eye; WPB/LP, form of
pronotum; WRA/LR, relative length of rostrum; WF/WRA, relative separation of eyes;
WE1M/LE1, form of elytra; WP/LE1, relative length of elytra; and, WPT/ WPB, position of
maximum width of pronotum.
A comprehensive set of descriptive statistics is given for one sample of each species treated.
(See Appendix 1, Tables 1-33). Statistical analysis is less comprehensive for those species
where there was not adequate material. Sexes are treated separately and ideally each sample
includes eight members of each sex collected at the same locality and time. This ideal is not met
for all species and some samples have fewer than eitht specimens of one of the sexes or, for the
purposes of more comprehensive statistical analysis, if markedly less than eight specimens are
represented, are composed of a number of specimens collected from a more extensive
geographic area. Least preferred samples are composed of all specimens of the species available
for study. Mensural data treated in the descriptive statistics section include only those
measurements and ratios given previously. Number of specimens, geographic locality of the
specimens, range, mean, 1.5 standard deviations (SD), and 2 standard errors (SE) are given
where number of specimens permits. In instances where too few specimens were available for
meaningful statistical treatment, values for SD and SE are not given. Number of standard
deviations and standard errors on either side of the mean were chosen following Whitehead
(1971). Non-overlap of two standard errors from the mean is equivalent to a t-test at 0.05
probability, and indicates samples to be statistically significantly different; non-overlap of 1.5
standard deviations from the mean indicates that 90% or more of the specimens from one
sample can be distinguished from 90% or more from the other sample, and indicates samples to
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Anderson
be taxonomically significantly different (Whitehead, 1971).
For most species these mensural features serve primarily as aids in identification and as
characters valuable for phylogenetic analysis. For some species, mensural features are the
primary, although not sole, criteria for species recognition. In these latter instances, the
mensural features have been verified as reliable by further measurements taken on a larger
number of specimens than that indicated in the descriptive statistics tables or on the
Hubbs-Hubbs diagrams ( e.g rostrum length in C. texanus-C. longinasus, form of elytra in C.
trivittatus-C. quadrilineatus).
Analysis of geographic variation Where problems in species recognition were apparent
in terms of geographic variation in structural features, population samples from sites
throughout the species range were selected and examined to determine the taxonomic
significance of this variation. If more or less extensive differences were found between samples,
geographically intermediate samples (if available) were examined to infer the degree of
reproductive isolation of the geographically more distant samples. Should geographically
intermediate populations prove structurally intermediate between the two more distant
samples, gene flow is inferred and all samples are considered conspecific. Divergence from both
of the more distant samples or close similarity to any one of the two distant samples by the
intermediate sample may indicate barriers to gene flow and may result in separate species
status for the two most distant samples, particularly if geographically correlated with other
variable structural features or aspects of natural history.
Descriptive format. — For each nomenclaturally valid taxon considered, a complete list of
synonymy is given. Type material examined and depository are noted following each of the
names of the nominal taxa. Lectotypes and neotypes are designated where appropriate;
paralectotypes have not been designated. Complete label data for primary type material
examined, type locality, gender, and type species are given where appropriate for species,
subgenera and genera. All literature citations of names of North American species, Cleonidius
vibex (Pallas), and all citations pertaining to Cleonis pigra (Scopoli) in North America are
given. A concerted attempt has been made to make these listings as complete as possible. Page
numbers have not been given for all citations, but only for those wherein new species are
described. Published misidentifications (“misident.” in synonymy list) are indicated under both
the incorrect and correct names. Notes in parentheses following most literature references
indicate the nature of that reference as follows: biol., biology; catal., catalog; check., checklist;
desc., description; distn., distribution; and redesc., redescription. Lack of any notation indicates
use of that name with little or no added information about the taxon.
For each species, subgenus or genus, a “Problems in recognition” section is presented which
I feel to be more desirable than a diagnosis because the latter, in most instances, simply gives
characters and their states already presented in the keys. In this section I discuss particular
species that I had problems separating from, and perhaps at one time misidentified or confused
with, the species in question. In this manner I point out the species with which others are likely
to make errors. This section is meant to be consulted after a tentative identification has been
made using the keys.
A section with notes about synonymy and type material follows and deals in various detail
with specific criteria for newly proposed synonymy and with other details about type material
as required. For all newly described species, label data for holotype and allotype are given in
full with data from each label enclosed in separate quotation marks starting from the top label
down. Separate lines on each label are indicated by a A specific type locality is designated.
New World Cleonini
443
Paratypes are treated less rigorously and label information is given in a simple standard format
of: locality, date, collector, sex, location of deposition. Specimens are ordered alphabetically
according to locality. Derivation of the specific epithet is also given.
Following this section is a detailed description of the taxon including structural variation.
Complementing this verbal description are various illustrations of important structural features
and for all species, a set of variously comprehensive descriptive statistics from one population
for each of males and females. Only attributes that vary between species are described.
Character states are described in consistent order to facilitate comparisons. Subjective terms
such as “large” and “small” have been avoided as much as possible but if by necessity used, are
augmented with illustrations. All structural variation is incorporated in the appropriate places
in the species description.
Geographic distribution of each species is presented in the form of dot maps illustrating
positions of localities of collections. State records and questionable localities of collection are
noted in the text but not placed on maps. Literature records are not placed on maps nor noted
in text unless they fall outside the known range of the species based on other specimens
examined. Complete lists of specimens examined for each species are not presented but have
been placed in the Archives of the Department of Entomology, University of Alberta.
Natural history information from publications, specimen label data, and personal collecting
experiences are summarized for each species. For the purposes of biogeographic and
evolutionary interpretations, habitat associations were recognized using, as appropriate,
schemes by Rzedowski (1978), Brown (1982), or Livingston and Shreve (1921). Elevations of
collection localities are either taken directly from specimen labels (if noted), or, if site of
collection is a major geographic feature, are taken from a gazetteer or one of a number of maps.
Hubbs-Hubbs diagrams illustrate variation in elevational distribution within and among species
of Cleonidius and Apleurus but no statistical tests are performed on these data. A
Hubbs-Hubbs diagram is used only for the purpose of visual presentation of the altitudinal
ranges. Seasonal data are taken from labels. Supra-generic classification of plants follows
Cronquist (1968). Generic names of plants follow Kearney and Peebles (1960); species names
follow Kearny and Peebles (1960), Munz and Keck (1968) and Correll and Johnston (1970).
Patterns of allopatry and sympatry for each species are summarized in the chorological
relationships section. They are summarized for Apleurus and Stephanocleonus species, but
presented in a table format for Cleonidius species. Consultation of this section not only reduces
possible misidentifications in instances where specimen locality is known, but also indicates
retention of species identity under variable conditions of species packing and species
occurrence. For this reason special attention is given to chorological relationships with close
phylogenetic relatives.
A section about phylogenetic relationships concludes the descriptive format and verbally
states the immediate phylogenetic relationships of the taxon. For species of Apleurus and
Cleonidius , further details are given in the “Phylogeny” section. Phylogenetic relationships
have not been examined for species of Stephanocleonus.
For species groups of Cleonidius, only a short diagnosis and a statement of evidence for
monophyly and phylogenetic relationships are presented. Structural features not variable
within a species group but variable within Cleonidius are repeated in the species group
diagnosis and all species descriptions.
Criteria for recognition of species-group taxa. — As with most systematists, my primary
criterion for species recognition is the biological species concept (Mayr, 1963). I find the
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Anderson
evolutionary species concept of Wiley (1981), although theoretically desirable, so vague as to
be of little practical applicability. Since experimental evidence of reproductive isolation is
lacking for most insect species, Cleoninae being no exception, such isolation is inferred from
differences primarily in structural features, but also in features of natural history, especially
host plant associations.
In instances of sympatry, individuals are regarded as belonging to separate biological species
if variable numbers of structural features are substantially and consistently different. Differing
host plant associations alone are not evidence of separate species status, but in combination
with differences in structural features are used herein to indicate separate species status. This
consideration is especially important should the degree of structural difference be less than is
found in sympatric species where host plant associations are broader or not consistently
different yet structural differences are relatively extensive.
For allopatric individuals the situation is more complex and difficult to resolve. Allopatric
individuals are regarded as conspecific if they exhibit structural features that do not differ, or if
differing, do so to a lesser degree than differences noted between separate sympatric species
and in a geographically based clinal manner. If the allopatric forms differ in various features of
structure and/or natural history, and these features cannot be shown to have a consistent
geographically determined pattern of continuous variation, the forms are considered separate
species. Obviously, the more characters that differ in this manner, the more marked the
evidence for separate species status. It is very important to note that marked phenetic
differences are not the primary consideration, rather it is the pattern of geographic variation
(assumed indicative of degree of gene flow) that determines the taxonomic status of the various
populations. Phenetic differences may simply be the result of locally different selection
pressures and not reproductive isolation.
These are my general criteria. Especially complicated and/or problematical patterns of
structural variation must be considered individually and as such are discussed in detail in the
text where appropriate.
The subspecific category has not been used. As expressed elsewhere (Anderson and Peck,
1986), I feel that this category should only be used for allopatric populations where lack of
intermediate populations precludes analysis of geographic variation and inference of degree of
reproductive isolation. In no instances in this study was such a consideration warranted;
geographically intermediate populations and natural history information were consistently
available to permit assessment of extent of gene flow and recognition of status at the species
level.
Criteria for recognition of genus-group taxa with notes about classification. — Some
consideration must be here given to criteria for recognition and ranking of genus-group taxa for
this study is concerned with a regional fauna and classification of its constituents in terms of
the world fauna. All genus-group taxa should preferably be monophyletic (holophyletic)
although paraphyletic taxa are acceptable in certain special situations; polyphyletic taxa are
not acceptable. Genus-group taxa should contain an assemblage of related species sharing
structural features and/or features of natural history, and separated from other assemblages by
a gap in such features. Aside from this requirement, naming and ranking of taxa are always
debatable because there are no objective criteria for formal recognition of taxa, and if
recognized, at what hierarchical level they should be placed. Although this has been attempted
(Hennig, 1966), it has not met with widespread acceptance or success.
New World Cleonini
445
All decisions about naming and ranking must be made with reference to related taxa with
genealogical and phenetic relationships of the taxa as the prime considerations. Genealogical
relationships (including some consideration of confidence in them), should take priority over
phenetic relationships because only classifications based on evolutionary parameters can
logically contain more information, due to inheritance, than was initially used in their creation
(Whitehead, 1972). Classifications of this nature may be maximally predictive and highest in
information content, but may prove difficult to employ as information retrieval systems
depending upon their complexity and intricacy. Many recent books (e.g., Eldredge and
Cracraft, 1980; Wiley, 1981; and references cited therein) have dealt with this problem at
length and simple compromise between information content and facility for use is perhaps the
most important general consideration in constructing classifications.
Four genera are recognized in this study as occurring in North America. Cleonis is
monobasic; Stephanocleonus is represented in North America by only 6 species but in the
Palearctic region by numerous species, so no intrageneric classification has been attempted
(species in North America are herein arranged alphabetically); Cleonidius is represented by 20
species; 19 Nearctic, 1 Palearctic (at least); Apleurus is represented by 8 Nearctic species.
In Cleonidius only informal species groups, including the incertae sedis category, are
recognized primarily because of low confidence in genealogical relationships and thus possible
instability of the resultant classification, and little variation in structural features. By using
only species groups, the possible predictive value of the classification is enhanced and can be
tested without the problems associated with increased nomenclatural complexity, not desirable
for such a small number of species of a relatively restricted geographic distribution.
In Apleurus , two subgenera have been recognized rather than species groups. Although
Apleurus contains only eight species, the genealogically most primitive member of Apleurus
differs from other species of Apleurus in a number of structural features, some not yet found
elsewhere in Cleoninae. Confidence in this primitive phylogenetic position is high and
structural differences are marked, with remaining Apleurus species forming a structurally
cohesive grouping. For these reasons, recognition of subgenera was deemed desirable.
Furthermore, desire for recognition of Gibbostethus as a subgenus is enhanced by the largely
unknown natural history, restricted distribution, and possible endangered status of this
phylogenetically most primitive species of Apleurus.
Classification above the generic level is not attempted, although some statements concerning
relatives of genera occurring in North America and their shared character states are made,
forming part of the basis for a future reclassification of the subfamily.
NATURAE HISTORY OF CFEONINAE
Members of the subfamily Cleoninae are found from mesic to very arid habitats. Taxa
traditionally placed in the tribe Fixini are those usually associated with more mesic habitats
whereas most traditional Cleonini are found in arid habitats. Exceptions, especially of some
Lixus species associated with arid habitats, are known.
Most species breed in herbs or shrubs of a wide variety of plant taxa but most notably
Chenopodiaceae, Compositae, Feguminosae, Rosaceae, Cruciferae and Polygonaceae. Farvae
of most species mine stems or roots of these plants but those of some feed in flower heads or on
seeds (e.g., Rhinocyllus conicus, Larinus planus, Microlarinus spp.). Natural history of
Microlarinus spp. (Kirkland and Goeden, 1977; 1978a, b) and of Rhinocyllus conicus (Zwolfer
Quaest. Ent., 1987,23 (4)
446
Anderson
and Harris, 1984) have been particularly well studied; however, most other information about
natural history of Cleoninae, especially in North America, is brief and fragmentary ( e.g .,
Bargagli, 1884; Pierce, 1907).
Adults of most species of traditional Cleonini are brachypterous and even those that are
macropterous have not been observed to fly. Most traditional Lixini are macropterous.
Adults of various species have been collected as prey of Bufo sp. (Amphibia), Geococcyx
californicus (Aves), and various predatory Hymenoptera. Tachinidae (Diptera) (Arnaud,
1978) and various parasitic Hymenoptera (Krombein et al., 1979) have been recorded as
parasites of members of the genus Lixus in North America.
Various species of Cleoninae have been employed as, or are being considered as, biological
control agents of weeds including thistle ( Cirsium , Compositae), knapweed ( Centaurea ,
Compositae), and puncturevine ( Tribulus , Zygophyllaceae) (Anderson, 1984a; Kirkland and
Goeden, 1978a, b; Maddox, 1976; Wheeler and Whitehead, 1985; Zwolfer and Harris, 1984).
None of the species are considered to be economic pests although some are occasionally found
on cultivated Chenopodiaceae or Cruciferae.
SUPRASPECIFIC TAXA AND INCLUDED SPECIES
SUBFAMILY CLEONINAE
Historical review
As of 1978, 1445 species of Cleoninae had been described (O’Brien and Wibmer 1978).
However, the number of genera stated is perhaps on the low side for many taxa given
subgeneric rank under Cleonus likely require separate generic status. The subfamily name
Cleoninae (first used as the vernacular “Cleonides”) and generic name Cleonus were first
proposed by Schoenherr (1826) but three years earlier, Schoenherr (1823) had also used the
vernacular name “Geomorides” and the generic name Geomorus for the same taxa. Earlier
still, Dejean (1821) had also proposed the name Cleonis to include species, some of which were
subsequently placed by Schoenherr in Geomorus and Cleonus. Since Schoenherr (1823) used
the vernacular name “Geomorides” and this was never latinized as Geomorinae, under Article
1 If of the International Code of Zoological Nomenclature it is not a valid family-group name.
Under Article 40, Cleoninae, even though the name of the type genus Cleonus , is a junior
synonym, should not be replaced and is maintained as the valid family-group name. The name
Cleoninae has been in widespread and constant use since its introduction (e.g., Schoenherr,
1834; Chevrolat, 1873; Faust, 1904; Csiki, 1934; O’Brien and Wibmer, 1982).
North American species have traditionally been included in the tribe Cleonini (Casey, 1891;
LeConte 1876a; LeConte and Horn, 1883) which is comparable to Cleoninae of other authors
(e.g., Csiki, 1934). There has been little confusion regarding included world taxa with the
exception of the Holarctic genus Lepyrus now generally regarded not to be a cleonine.
Recognition of Cleoninae
Adults of the subfamily Cleoninae are recognized by two features not known in other
Curculionidae. These are: 1), short, telescoping, ventrally situated, three-articled labial palpi;
and 2), presence of paired symbiont sacs attached to the vagina near the base of gonocoxite II
in females. A more detailed diagnosis describing other features of the subfamily is not
presented, pending examination of further taxa.
New World Cleonini
447
Sex differentiation
Sexes are differentiated in Cleoninae as in most other weevils. Males are easily
distinguished by flat to concave abdominal sterna III and IV and by possession of two clearly
recognizable sclerotized terga at the abdominal apex, usually only visible upon displacement of
the elytral apices. Females possess more evenly convex abdominal sterna III and IV and possess
only a single sclerotized tergum at the abdominal apex. In addition, females tend to have a
longer rostrum than do males (especially in Cleonidius) and are generally larger in body size.
Key to genera of New World Cleoninae
1 Rostrum, in dorsal view, from apex of epistoma to anterior margin of eye,
more or less as long as greatest width (Figs. 1-2) 2
Y Rostrum, in dorsal view, from apex of epistoma to anterior margin of eye,
much longer than greatest width (Figs. 3-4, 10, 25-30, 81-88, 137-156) 4
2 (1) Prosternum with a pair of prominent ridges which form a deep ventral
channel Bangasternus Gozis
(two species, one deliberately introduced, one under consideration for
deliberate introduction)
2' Prosternum lacking ridges, no channel evident 3
3 (20 Elytron with intervals each with row of conspicuous erect setae.
Anterolateral margin of pronotum straight, with long postocular vibrissae
immediately behind eye (Fig. 1). Eye more or less round (Fig.
1) Microlarinus Hochhuth
(two species, deliberately introduced)
y Elytron with intervals without row of erect setae. Anterolateral margin of
pronotum with rounded postocular lobe and short postocular vibrissae
behind eye (Fig. 2). Eye distinctly elongate-oval (Fig. 2)
Rhinocyllus Germar
(one species, deliberately introduced)
4 (T) Pronotum dorsally and laterally with numerous shiny glabrous tubercles,
lacking distinct punctures 5
4 ' Pronotum dorsally and laterally distinctly punctate, with at most the outer
margins of punctures of lateral margins swollen, glabrous and shiny 6
5 (4) Rostrum dorsomedially sulcate throughout length (Fig. 10). Pronotum
with anterolateral margins with rounded postocular lobe (Fig. 9)
Cleonis Dejean, p. 449
(one species, adventive)
5' Rostrum dorsomedially carinate throughout length; lateral margins raised
basally, appearing as lateral carinae. Pronotum with anterolateral margins
straight, lacking rounded postocular lobe .... Cyphocleonus Motschulsky
(one species, under consideration for deliberate introduction)
6 (4') Mesosternum with mesosternal process markedly tumescent (Fig. 58).
Male with aedeagus markedly expanded laterally from midlength to apical
one-third (Fig. 97e) . . Apleurus subgenus Gibbostethus Anderson, p. 468
(one species, native)
6' Mesosternum with mesosternal process flat or at most only slightly convex,
Quaest. Ent., 1987,23 (4)
448
Anderson
not tumescent (Fig. 59). Male with aedeagus more or less uniform in width
throughout median portion of length 7
7 (6') Tibia with corbel ridge rounded (Fig. 5). Elytron with all intervals equally
flat or with at most only humerus and very base of interval 3 swollen and
convex. Pronotum with disk with scale pattern various. Prosternum with or
without swellings; swellings, if present, situated immediately anterior to
each procoxal cavity (Fig. 8) 8
7' Tibia with corbel ridge sharp (Fig. 6). Elytron with all intervals equally flat
or with humerus and variously, sutural interval and intervals 3, 5, 7, and 9
elevated and convex throughout the greater part of their lengths. Pronotum
with di-sk with large white scales in lateral stripe of various width, small
and fine in moderately broad to very broad apically narrowed median
stripe; median area largely black in color, underlying dark cuticle not
obscured by overlying scales. Prosternum with or without swellings;
swellings, if present, situated immediately anterior to each prosternal
impression (Fig. 7) 10
8 (7) Antenna with article 2 of funiculus distinctly longer than wide, distinctly
longer than each of articles 3 to 6, slightly shorter than to distinctly longer
than article 1 (Fig. 4). Pronotum with anterolateral margins straight,
slightly sinuate, or with at most variously developed (usually small) acute
postocular projection; postocular vibrissae unequal in length, greatest
length (more or less equal to or greater than one-half width of eye) behind
base of eye (Fig. 4). Ventral surface of femur dentate or not. (Some South
American species of Lixus have slightly developed rounded postocular
lobes and moderately long postocular vibrissae of more or less equal length,
but have dentate femora) Lixus Fabricius
(numerous species, native; worldwide)
8' Antenna with article 2 of funiculus more or less as long as wide, more or
less subequal in length to each of articles 3 to 6, shorter than article 1
(Figs. 3, 137-156). Pronotum with anterolateral margins straight or with
slightly to well-developed, rounded postocular lobe; postocular vibrissae
uniformly short (less than one-half width of eye in length) to unequal in
length, greatest length (more or less equal to or greater than one-half width
of eye) behind base of eye (Figs. 3, 137-156). Femur not dentate 9
9 (80 Elytra elongate-narrow (width at midlength less than 0.65 times length)
(Figs. 118-136). Pronotal disk with distinct white scales of various sizes
(Figs. 118-136). Elytra with white scales, various in size, but more or less
obscuring view of underlying cuticle over large part of elytral surface (Figs.
118-136) Cleonidius Casey, p.492
(19 species, native; also in Palearctic Region)
9' Elytra more robust (width at midlength greater than 0.65 times length).
Pronotal disk with at most only very short indistinct setae, distinct scales
absent. Elytra with scattered patches of elongate fine scales in addition to
very short setae; underlying cuticle not obscured by scale cover
Larinus Schoenherr
(one species, introduced)
New World Cleonini
449
10 (7') Epistoma with anterior margin emarginate (Figs 25-30). Pronotum with
well-developed, rounded postocular lobes; postocular vibrissae indistinct,
uniformly short (Figs. 25-30). Eye elongate “kidney shaped” (Figs.
25-30); flat, or only slightly convex in dorsal view
Stephanoeleonus Motschulsky, p. 453
(six species, native; also in Palearctic Region)
10' Epistoma with anterior margin rounded (Fig. 82-88). Pronotum with
anterior margin straight behind eyes or with small acute postocular
projection immediately behind base of eye; postocular vibrissae distinct and
long, longest immediately behind base of eye (Figs. 82-88). Eye “teardrop
shaped” (Figs. 82-88); protruding and moderately to markedly convex in
dorsal view Apleurus subgenus Apleurus Chevrolat, p. 470
(seven species, native)
Ileomus Schoenherr is not given separate generic status in this key. Champion (1902-1906),
although regarding it as a separate genus, states that “it is scarcely distinct from Lixus
Accordingly, individuals of species that are assigned to Ileomus by O’Brien and Wibmer
(1982) key out in the above key to Lixus. Detailed study of phylogenetic relationships of the
various groupings of Lixus species is needed to assess their validity as nominal subgeneric or
generic taxa.
Genus Cleonis Dejean
Curculio-, Scopoli 1763 (in part; sp. desc.). Linnaeus 1767 (in part; sp. desc.). Goeze 1777 (in part; sp. desc.). Knoch 1781
(in part; sp. desc.). Villers 1789 (in part; sp. desc.). Gmelin 1790 (in part; sp. desc.).
Cleonis Dejean 1821:96. Gender, feminine. Type species Curculio sulcirostris Linnaeus by subsequent designation
(O’Brien and Wibmer 1982:73). Silfverberg 1979 (in part; check.). O’Brien and Wibmer 1982 (in part; catal., distn.,
misident. Apleurus, Cleonidius, Stephanoeleonus). O’Brien and Wibmer 1984 (in part; misident. Apleurus,
Cleonidius, Stephanoeleonus ).
Geomorus Schoenherr 1823: column 1141. Gender, masculine. Type species Curculio sulcirostris Linnaeus by original
designation.
Cleonus Schoenherr 1826:145. Unjustified replacement name for Geomorus Schoenherr. Type species Curculio
sulcirostris Linnaeus according to Article 67(h) of International Code of Zoological Nomenclature. Say 1831
(misident. Cleonidius ). Kirby 1837 (misident. Cleonidius). LeConte 1850 (misident. Stephanoeleonus). Melsheimer
1853 (misident. Cleonidius). LeConte 1858 (misident. Apleurus). LeConte 1859 (misident. Apleurus). Gemminger
and von Harold 1871 (in part; misident. Apleurus, Cleonidius). LeConte 1876a (misident. Cleonidius). Henshaw
1881-1882 (misident. Cleonidius). LeConte and Horn 1883 (misident. Cleonidius). Henshaw 1885 (misident.
Cleonidius). Wickham 1889 (misident. Cleonidius). Wickham 1896 (misident. Cleonidius). Fall 1897 (misident.
Cleonidius). Fall 1901 (misident. Cleonidius). Wickham 1902 (misident. Cleonidius). Fletcher 1906 (misident.
Cleonidius). Fall and Cockerell 1907 (misident. Cleonidius). Pierce 1907 (misident. Cleonidius). Ely 1913 (misident.
Cleonidius). Gibson 1914 (misident. Cleonidius). Anderson 1914 (misident. Cleonidius). Blatchley and Leng 1916
(misident. Cleonidius). Yothers 1916 (misident. Cleonidius). Leng 1920 (misident. Stephanoeleonus, Apleurus,
Cleonidius). Leonard 1926 (misident. Stephanoeleonus , Cleonidius). Boving 1927 (in part; larval key). Bradley 1930
(misident. Apleurus, Cleonidius). Crosby and Blauvelt 1930 (biol.). Leng and Mutchler 1933 (catal.). Wilcox et al.
1934 (misident. Cleonidius). Bleasdell 1937 (misident. Stephanoeleonus, Cleonidius). Brimley 1938 (misident.
Cleonidius). Brown 1940 (distn.). Blackwelder 1947 (misident. Apleurus, Cleonidius). Bruhn 1947 (misident.
Cleonidius). Hicks 1947 (distn.). Hicks 1949 (distn.). Anderson 1956 (biol., distn.). Essig 1958 (misident.
Cleonidius). Sanders 1960 (misident. Cleonidius). Kissinger 1964 (in part; biol.). Tanner 1966 (misident. Apleurus,
Cleonidius). Kingsolver 1972 (misident. Cleonidius). Burke and Anderson 1976 (biblio.). Kumar et al. 1976
(misident. Cleonidius). Krombein 1979 (misident. Stephanoeleonus, Apleurus). Arnett et al. 1980 (misident.
Cleonidius). Batra et al. 1981 (distn.). Peschken 1984 (biol.).
Cleonus s.s.\ Csiki 1934 (catal.).
Cleonus ( Cleonidius ); Arnett 1960-1962 (in part; misident. Apleurus).
Quaest. Ent., 1987, 23 (4)
450
Anderson
Notes about synonymy. — Epimeces Billberg, 1820, p. 45, predates Cleonis Dejean, 1821
and originally included various Cleoninae (among others, Curculio sulcirostris Linnaeus, the
type species of Cleonis Dejean, and, Curculio filiformis Fabricius, presently assigned to Lixus
Fabricius [Csiki 1934]). The name Epimeces , to my knowledge, has not subsequently been used
but because no type species has been designated for the genus, the name represents a threat to
the priority of Cleonis Dejean. I therefore here designate Curculio filiformis Fabricius, 1781,
p. 172, as the type species of Epimeces Billberg, 1820, p. 45. Epimeces therefore becomes a new
junior subjective synonym of Lixus Fabricius, 1801, p. 498, type species Curculio paraplecticus
Linnaeus, 1758, p. 380, by subsequent designation (Latreille, 1810, p. 430).
Schoenherr (1823), seeking consistency in gender of generic names, changed all feminine
generic names in Curculionidae to a masculine name (not always the masculine form of the
former feminine name). As a result of this, and according to Schoenherr (1823), inadequate
characterization to recognize a type species, use of the name Cleonis Dejean, 1821 was
discontinued, and the species initially placed in Cleonis were assigned to one of four genera by
Schoenherr (1826). One of these was Geomorus Schoenherr, 1823. As such, Geomorus cannot
be regarded as an express replacement name for Cleonis. Subsequently, Schoenherr (1826)
proposed the genus Cleonus expressly as a replacement name (although unjustified) for
Geomorus as indicated by his designation of Curculio sulcirostris Linnaeus, 1767 as the type
species. This same species had already been designated by him as the type species of Geomorus.
Schoenherr (1826) also placed all species initially included in Geomorus in Cleonus. The
generic name Cleonus has since been in widespread use. However the name Cleonis was
recently resurrected by Silfverberg (1979) and used by O’Brien and Wibmer (1982), Lohse
(1983), and Wheeler and Whitehead (1985).
Cleonis Dejean is here used in a very restricted sense (subgenus Cleonis of Csiki [1934])
including only C. pigra (Scopoli), C. japonicus (Faust), and C. sardous (Chevrolat), the latter
two species not examined by me. Cleonis appears closely related to Cyphocleonus Motschulsky
and Adosomus Faust, and subsequent phylogenetic analysis may reveal the three to be
congeneric.
Diagnosis. — Adult Cleoninae with moderately robust body form (Fig. 19). Rostrum with
broad low median carina longitudinally sulcate throughout length (Fig. 10). Pronotum with
small glabrous shiny tubercles, not distinctly punctate; pronotal postocular lobes slightly to
moderately well-developed, rounded. Tibia with corbel ridge sharp (as in Fig. 6). Elytra with
strial punctures individually indistinct, dorsal surface of elytra with scattered, small, irregularly
shaped (usually transverse), glabrous, shiny swellings.
Description. — Size. Moderately large, moderately robust in form. Mouthparts. Prementum flat, with single large
seta on each side. Maxillary palpus with palpifer and stipes each with large seta. Basal articles of labial palpi separated by
distance subequal to width of a palpus. Rostrum. Moderately robust; with broad low median carina longitudinally
moderately deeply sulcate throughout length from base of epistoma to base of frons; not medially tumescent (Figs. 9-10).
Epistoma slightly swollen, with apical margin emarginate medially. Antenna with funiculus with article 1 very slightly
longer than article 2 (Fig. 9); apical three articles of club with placoidal sensillae. Head. Eye elongate-quadrate, slightly
wider at top than bottom; flat (Fig. 9). Vestiture. Dorsum lacking erect or suberect vestiture, with only simple
elongate-narrow appressed white scales. Procoxae, mesocoxae, metasternum and base of abdominal sternum III of both
sexes with moderately long suberect hair-like scales. Prothorax. Dorsal surface of pronotum with small glabrous shiny
tubercles, not distinctly punctate. Median basal area shallowly impressed. Disk with elongate, moderately large,
moderately dense, white scales present in broad lateral stripe and narrow median line, with scales small and fine in pair of
paramedian apically narrowed stripes; median area largely black in color, underlying dark cuticle not obscured by
overlying scales (Fig. 19). Pronotum widest at base, lateral margins slightly rounded and convergent from base to apex;
apical constriction very slight (Fig. 19). Pronotal postocular lobes rounded, slightly to moderately well-developed (Fig. 9).
Prosternum with slight impression anterolaterad of each procoxal cavity and with slight swelling immediately anterior to
each prosternal impression. Legs. Tarsi broad, articles 2 and 3 more or less as broad as long, subequal in length; article 1
New World Cleonini
451
only slightly longer than articles 2 or 3; article 3 deeply bilobed (Fig. 18). Ventral tarsal pilosity extensive, covering entire
ventral surface of articles 1 to 3 (Fig. 18). Claws connate in basal one-third, not to slightly divergent. Foretibia with small
to moderately well-developed second spur; inner margin with moderately large denticles throughout greater portion of
length. Tibia with corbel ridge sharp. Metatibia of male with uncus with ventral margin slightly sinuate. Wings. Present.
Elytra. Base of interval 3 and humerus very slightly swollen and convex, otherwise intervals uniformly flat. Humeri
distinct. Strial punctures individually indistinct, dorsal surface of elytra with scattered small irregularly shaped (usually
transverse) glabrous shiny swellings. Abdomen. Ventral surface with small shiny glabrous patches, each with single small
appressed scale. Genitalia. Female. Abdominal sternum VIII lacking basal arm (Fig. 17). Gonocoxite II
elongate-triangular, apex not prolonged into marked lobe; stylus moderately large, apical in position (Fig. 15).
Spermathecal gland slightly elongate-oval (Fig. 16). Male. Aedeagus elongate, moderately robust; in lateral view,
moderately arcuate near base then only slightly arcuate throughout rest of length (Fig. 13). Apex slightly spatulate.
Internal sac with median dorsal pocket high; various lobes present; apical and dorsal median pockets individually distinct;
apical sclerite complex present, individual sclerites distinctly scythe-like (Figs. 11, 12, 14).
Comparisons. — As noted, Cleonis species are very similar to those of Adosomus and
Cyphocleonus in that all possess a pronotum and elytra that are distinctly tuberculate and not
punctate. Adosomus and Cleonis species possess moderately well-developed, rounded,
postocular lobes, whereas postocular lobe are lacking from Cyphocleonus species. The only
feature which distinguishes Cleonis from both of these two genera is the medially longitudinally
sulcate rostrum of the former. In Adosomus the rostrum is medially tumescent and not carinate
or sulcate, whereas in Cyphocleonus species, the rostrum is variously medially tumescent and
carinate. Cyphocleonus trisulcatus (Herbst) has a low broad median carina but which is only
apicomedially sulcate.
Checklist of included species. — Following Csiki (1934), three species are recognized. They
are C. japonicus (Faust), C. sardous (Chevrolat), and C. pigra (Scopoli). Cleonis pigra is the
only one of the three species that I have examined and is the only one that occurs in the
Nearctic Region.
Faust (1904) distinguishes C. pigra and C. japonicus from C. sardous by the latter having
metatarsal article 2 slightly longer than article 3 and by differences in the pronotal scale
patterns. Cleonis pigra and C. japonicus are separated on the basis of extent of distribution and
development of the pronotal and elytral tubercles, the distribution of the ventral abdominal
glabrous patches, and body shape. Based upon examination of only Nearctic representatives of
C. pigra , I find all of these characters vary somewhat, and suspect that the three included
species may prove conspecific.
Phylogenetic relationships. — Cleonis, Cyphocleonus and Adosomus appear to represent a
monophyletic group based on the presence of a tuberculate rather than punctate pronotum and
elytra. Beyond this, affinities of the group are uncertain.
Cleonis pigra (Scopoli)
(Figs. 9-19,213)
Curculio piger Scopoli 1763:23.
Curculio sulcirostris Linnaeus 1767:617.
Curculio transversofasciatus Goeze 1777:409.
Curculio nebulosus Knoch 1781:87 [not Linnaeus 1758:385].
Curculio fasciatus Villers 1789:216 [not Muller 1776:86].
Curculio fasciatus Gmelin 1 790: 1 804.
Cleonis sulcirostris ; Dejean 1 821 .
Cleonus sulcirostris ; Gyllenhal 1834.
Cleonus indicus Fahraeus 1842:55.
Cleonus piger. Everts 1903.
Quaest. Ent., 1987, 23 (4)
452
Anderson
Cleonis piger, Silfverberg 1979 (check.)- O’Brien and Wibmer 1982 (catal., distn.).
Cleonis pigra\ O’Brien and Wibmer 1984.
Notes about synonymy. — The synonymical list follows Csiki (1934); I have not examined
type material of this species or of any of its conspecific forms. This is the species frequently
referred to in previous publications as C. sulcirostris.
Description. — Specimens examined. 61 unsexed. Size. Length, male, 7.5-13.4 mm; female, 11.4-14.8 mm.
Width, male, 3. 7-6. 6 mm; female, 5. 8-7.0 mm. Rostrum. Lateral margins sharp, indicated by low rounded carina from
above point of antennal insertion to frons (Fig. 10). Side portion of rostrum immediately ventrad of lateral carina
moderately deeply sulcate from immediately anterior to eye to point of antennal insertion; with moderately deep
elongate-narrow punctures, most linearly confluent or nearly so. Prothorax. Surface dorsally and laterally with numerous
small shiny glabrous tubercles; medially with small to large elongate-oval shiny glabrous swelling. Prosternum in lateral
view with apical portion flat, slightly shorter than adjacent posterior steeply declivous portion. Elytra. Sutural interval in
basal one-half, intervals 2 to 5 basally, and two obliquely posteromedially directed patches (at basal one-third from
intervals 2 to 5 and at apical one-third from intervals 2 to 4) with small irregular slightly elevated shiny glabrous patches
(Fig. 19). Wings. Long (greater than elytra in length). Bases of 2A joined but very lightly sclerotized. Abdomen. Ventral
surface with small shiny glabrous patches distinct and dense on abdominal sterna III and IV, less distinct on sterna V to
VII. Genitalia. Female (2 specimens). Abdominal sternum VIII with lateral arms narrow, evenly and slightly inwardly
arcuate throughout length, expanded apically (Fig. 17). Male (2 specimens). Aedeagus robust; in lateral view narrowed
from approximately apical one-third to apex (Fig. 13). Internal sac with single, large, broad, dorsally directed, lobe; with
moderately large, paired, laterally directed, lobes and small dorsally directed median lobe on dorsal surface at midlength;
with small dorsolaterally directed paired lobes at midlength and midheight (Figs. 1 1-12). Eversible apical sclerite complex
with paired scythe-like sclerites, each with median elongate-narrow projection long, visible in lateral view (Figs. 11, 14);
adjacent ventral surface and basal portion of sides of apex of sac with pair of large transverse dark sclerites (Fig. 11).
Geographic distribution. — This species was accidentally introduced into North America
from Europe prior to 1919, the year of the earliest known North American record (Anderson,
1956). It occurs throughout New York and southern Ontario, west to Michigan and east to
eastern Quebec and New Brunswick (Fig. 213). It is widespread in the Palearctic Region
(Csiki, 1934).
Variation. — As part of a program investigating the potential for use of C. pigra as a
biological control agent for the introduced Centaurea diffusa Lam. (diffuse knapweed) in the
prairie provinces of Canada, a study of variation based on 15 mensural features was carried out
on adult individuals of C. pigra reared from species of Cirsium , Carduus and Centaurea (all
Compositae) to determine if structurally distinct host races exist (Anderson, 1984a).
Measurements were made of 15 characters and a linear discriminant analysis was performed.
Separation using the generated discriminant function proved unreliable and because absolute
size was the principal component upon which the function was based, it was concluded that
only a single polyphagous morphotype was present. Differences in absolute size are likely a
result of structural attributes of the different larval feeding and pupation sites in the different
host plants and are doubtfully heritable.
Natural history. — Anderson (1956) summarizes information about the natural history of
this species in North America as follows. Adults are the overwintering stage and are first found
on Canada thistle, Cirsium arvense (L.) Scop., or bull thistle, Cirsium vulgare (Savi) Tenore,
in early June. Copulation and oviposition take place through mid-July. Eggs are laid singly in
the lower portions of the stems in cavities chewed by the females and subsequently plugged
with frass following deposition of the egg. Larvae bore downward into the primary root where
they feed for approximately 30 days. Pupation takes place in the root. Adults emerge in August
or September and overwinter under ground debris. Although a wide range of Compositae serve
as host plants in Europe (Anderson, 1956; Peschken, 1984, and references cited therein), only
two plants, Cirsium arvense and Cirsium vulgare , both adventive, are known to serve as hosts
in North America (Anderson 1956). Native North American Cirsium species or other
Compositae are not attacked by the weevils.
New World Cleonini
453
A similar range of host plant associations to those of C. pigra in Europe is also found in
another species of Cleoninae, Rhinocyllus conicus Froelich, which was recently shown to
consist of various host specific races (Zwolfer and Preiss, 1983). As such, host plant races are
also suspected in C. pigra although, as noted, an examination of the structural features of
adults reared from various Compositae did not reveal differences between individuals reared
from different plant taxa.
Immature stages were described by La Ferla (1939).
Genus Stephanocleonus Motschulsky
Cleonus ; LeConte 1850 (in part). Leng 1920 (in part; catal.). Leonard 1926 (in part; check.). Bleasdell 1937 (in part).
Krombein 1979 (in part; as prey).
Stephanocleonus Motschulsky 1860:540 (in key). Gender, masculine. Type species Curculio flaviceps Palliser by original
designation. LeConte 1876a (key, sp. descs.). Henshaw 1881-1882 (in part; check.). LeConte and Horn 1883 (key).
Henshaw 1885 (check.). Blatchley and Leng 1916 (key, sp. redesc.).
Coniocleonus Motschulsky 1860:540 (in key). NEW SYNONYMY. Gender, masculine. Type species Cleonus
carinirostris Gyllenhal by original designation.
Plagiographus Chevrolat 1873:21. Gender, masculine. Type species not designated.
Cleonus ( Stephanocleonus ); Casey 1891 (in part; key). Fa 11 and Cockerell 1907 (in part; check.). Csiki 1934 (in part;
catal.). Arnett 1960-1962 (in part; catal., key). Kissinger 1964 (in part; key).
Cleonis\ O’Brien and Wibmer 1982 (in part; catal., distn.). O’Brien and Wibmer 1984 (in part).
Notes about synonymy. — The genera Stephanocleonus and Coniocleonus were both
originally proposed by Motschulsky (1860). They were distinguished by the rostrum having
only a single median carina in the latter, whereas in Stephanocleonus , the rostrum, in addition
to the median carina, also had a smaller oblique carina on each side. Stephanocleonus is also
noted as being apterous but nothing is said of the state of the wings in Coniocleonus. I have
examined specimens of Stephanocleonus flaviceps Palliser, the type species of
Stephanocleonus , and, in the single male available, did not see small oblique lateral carinae,
but only a single uniformly steeply declivous median carina. The two females examined have
the median carina similarly steeply declivous basally but decreasingly declivous apically. A
very slight oblique carina is present where this more gradual apical declivity meets the flat
surface of the rostrum on each side of the median carina. All three individuals are wingless.
I have not seen specimens of Coniocleonus carinirostris Gyllenhal, the type species of that
genus, but I have examined a male and a female of each of C. excoriatus Gyllenhal and C.
glaucus Fabricius, a female of C. cineritius Gyllenhal, and a male of each of C. cinerascens
Hochhuth and C. ferrugineus Fahraeus. Individuals of C. excoriatus and C. glaucus are
macropterous and the rostrum has a single median carina which is gradually declivous
throughout its length. There are no small lateral carinae evident. However, the specimens of C.
cineritius , C. cinerascens and C. ferrugineus each have a rostrum with slightly to moderately
well-developed oblique lateral carinae in the same position as in the females of S. flaviceps, and
have wings of variable length.
In view of this variation and the lack of other distinguishing characters, I cannot accept
other than that Stephanocleonus is simply an apterous Coniocleonus. I do not think the rostral
differences between the type species warrant separate generic status. I therefore consider
Stephanocleonus and Coniocleonus to be new subjective synonyms, a decision anticipated by
Casey (1891:188) who stated, regarding the Nearctic species, that “It is a question whether our
species should be placed in Stephanocleonus or Plagiographus [a junior synonym of
Coniocleonus according to Faust, 1904 and Csiki, 1934], but this is a matter of but slight
importance as the differences between the subgenera appear to be very inconsiderable”.
Quaest. Ent., 1987, 23 (4)
454
Anderson
I choose to give the name Stephanocleonus priority over Coniocleonus because the former
has been the only name used in reference to the Nearctic species here placed in the genus.
Furthermore, no major works have recently appeared dealing with Coniocleonus , however, a
recent review by Ter-Minasyan (1979) of the Stephanocleonus of the Palearctic Region
includes species that in my opinion are congeneric with species in North America.
Diagnosis. — Adult Cleoninae with moderately robust to robust body form (Figs. 20-23).
Eye kidney-shaped, flat (Figs. 25-30). Rostrum with well-developed sharp median carina
(Figs. 25-30). Pronotum punctate; pronotal postocular lobes well-developed, rounded;
postocular vibrissae uniformly short (Figs. 25-30). Meso- and especially metatarsus
elongate-narrow, with all articles markedly longer than broad; article 2 distinctly longer than
article 3; article 1 markedly longer than articles 2 or 3. Tibia with corbel ridge sharp (as in Fig.
6). Procoxae, mesocoxae, metasternum, metacoxae and base of abdominal sternum III of male
with moderately long to very long erect hair-like scales; hair-like scales sparse to lacking in
female. Pronotal disk with elongate, white, moderately large, moderately dense scales present
in lateral stripe of various width, small and fine in moderately broad to very broad, apically
narrowed, median stripe, and variously small and fine to moderately large and elongate along
lateral margins; median area largely black in color, underlying dark cuticle not obscured by
overlying scales (Figs. 20-23). Prosternum with or without swellings, swellings, if present,
immediately anterior to each prosternal impression (as in Fig. 7). Metauncus of male with
ventral margin slightly to markedly sinuate. Female with abdominal sternum VIII with basal
arm very long (Figs. 37-42); gonocoxite II rounded basally, with apex prolonged into a marked
lobe (Fig. 54). Male with apex of aedeagus spatulate or not (Figs. 31-36); internal sac with or
without large paired sclerite at midlength (Figs. 31-36).
Description. — Size. Moderately large, moderately robust to robust in body form (Figs. 20-23). Mouthparts.
Prementum ventrally longitudinally carinate, slightly swollen to flat, with one or more large setae on each side. Maxillary
palpus with palpifer and stipes each with at least one large seta. Bases of labial palpi separated by distance subequal to
width of basal article of labial palpus. Rostrum. Moderately to markedly robust, not medially tumescent; with low to high,
sharp median carina (Figs. 25-30). Epistoma markedly swollen or not, with apical margin emarginate medially (Figs.
25-30). Antenna with funiculus with article 1 slightly to markedly longer than article 2 (Figs. 25-30); apical three articles
of club with placoidal sensilla. Head. Eye kidney-shaped, flat (Figs. 25-30). Upper margin of eye rounded to sharp, frons
convex to markedly concave. Vestiture. Dorsum lacking or with at most only very short indistinct suberect or erect
vestiture, with simple elongate-narrow appressed white scales of various size and density. Procoxae, mesocoxae,
metasternum, metacoxae and base of abdominal sternum III of male with moderately long to very long erect hair-like
scales; hair-like scales sparse to lacking from female. Prothorax. Dorsal surface of pronotum punctate. Pronotum with
median basal area shallowly to deeply impressed, disk with or without various other impressions. Disk with elongate white
moderately large moderately dense scales present in lateral stripe of various width, small and fine in moderately broad to
very broad apically narrowed median stripe, and variously small and fine to moderately large and elongate along lateral
margins; median area largely black in color, underlying dark cuticle not obscured by overlying scales (Figs. 20-23).
Pronotum widest at base to subequal in width from base to apical one-quarter, then constricted to various extent and
convergent to apex (Figs. 20-23). Pronotal postocular lobes moderately to well-developed, postocular vibrissae uniformly
short (length less than one-half maximum width of eye) (Figs.; 25-30). Prosternum with slight impression anterolaterad of
each procoxal cavity; with or without slight swelling immediately anterior to each prosternal impression (as in Fig. 7).
Legs. Foretarsus moderately broad, articles 2 and 3 more or less subequal in length, at most slightly longer than broad;
article 1 only slightly longer than articles 2 or 3; article 3 moderately deeply bilobed. Meso- and especially metatarsus
elongate-narrow; all articles markedly longer than broad, article 2 distinctly longer than article 3; article 1 markedly
longer than articles 2 or 3, article three slightly bilobed. Ventral tarsal pilosity various in extent, lacking entirely on at least
more basal articles in most species. Claws connate in basal one-third, not to slightly divergent. Foretibia with inner margin
near apex with moderately-developed subapical tooth, inner margin with at most only small denticles in apical one-half.
Metatibia of male with uncus with ventral margin slightly to markedly sinuate. Tibia with corbel ridge sharp. Wings.
Absent or present (various in length). Elytra. Intervals flat to variously slightly swollen and convex; striae distinctly
punctate. Humeri acute to rounded. Dorsal surface of most species with variously developed, posteromedially directed
oblique patches of small and fine scales at each of basal one-third and apical one-third to midlength (Figs. 20, 22-23).
Abdomen. Ventral surface with small shiny glabrous patches, each with single small appressed scale. Sternum VII of male
with apex variously medially emarginate in most species. Genitalia. Female. Abdominal sternum VIII with long basal arm
New World Cleonini
455
(Figs. 37-42). Gonocoxite II rounded basally, with apex prolonged into marked lobe; stylus moderately large, apical in
position (Fig. 54). Spermathecal gland round (Fig. 48). Male. Aedeagus robust and short to elongate-narrow, various in
shape in lateral view; apex not to markedly spatulate (Figs. 31-36). Internal sac various; median pocket low, apical and
dorsal median pockets individually distinct or not, various lobes present; apical sclerite complex with individual sclerites
distinctly scythe-like, large paired sclerite at midlength present or absent (Figs. 43-48).
Comparisons. — Stephanocleonus is very similar to Pleurocleonus however, the latter lacks
pronotal postocular lobes, has an elongate-teardrop-shaped eye, tarsal claws that are widely
divergent, and a low broad longitudinally sulcate median rostral carina. Species of
Stephanocleonus may also be confused with Nomimonyx and Epirhynchus because of
similarities in head and rostral form, and eye shape. Epirhynchus however, has a single tarsal
claw, and both have short broad tarsal articles, not the markedly elongate-narrow articles of
Stephanocleonus species.
Although unlikely, some Conorhynchus, Bothynoderes and Chromonotus may also be
confused with Stephanocleonus species. These taxa however have article 2 of the antennal
funiculus longer than article 1; in Stephanocleonus species, article 1 is longer than article 2.
Checklist of included species. — In the Nearctic Region, six species are placed in
Stephanocleonus. Numerous species occur in the Palearctic Region (Csiki 1934; Ter-Minasyan
1979) but are not listed here. The six Nearctic species are as follows:
1. S. confusus Anderson, n. sp.
2. S. cristaticollis Csiki
3. S. immaculatus Anderson, n. sp.
4. S. parshus Anderson, n. sp.
5. S. plumbeus (LeConte)
6. S. stenothorax Anderson, n. sp.
Phylogenetic relationships. — Distribution of the apotypic character state of female
gonocoxite II (rounded basally and with a marked apical lobe [Fig. 54]) suggests that
Stephanocleonus , Pleurocleonus Motschulsky, Menecleonus Faust, Xanthochelus Chevrolat,
and Conorhynchus Motschulsky ( =Temnorhinus Chevrolat) form a monophyletic group.
Menecleonus, Xanthochelus and Pleurocleonus appear to further comprise another
monophyletic group (based upon presence of an unpaired sclerite near the confluence of the
apical pocket and the dorsal median pocket in the internal sac of the aedeagus of males) that is
the sister-group of Stephanocleonus. This proposed sister-group relationship is based of article
1 of the antennal funiculus being longer than article 2. Conorhynchus has article 2 of the
funiculus longer than article 1 and is likely the sister-group to the
Stephanocleonus-Pleurocleonus, Menecleonus, Xanthochelus lineage. Bothynoderes
Schoenherr, Chromonotus Motschulsky and also Chromosomus Motschulsky, have the
apotypic state of elongate-narrow meso- and metatarsi and together with the above taxa likely
comprise another monophyletic group based upon shared possession of this character state.
They also have a very similar “elongate-teardrop” eye-shape to Conorhynchus and similarly all
have article 2 of the antennal funiculus longer than article 1.
Key to species of adult Stephanocleonus
1 Dorsal surface of elytra uniformly covered with small white scales, without
distinct maculations; with or without small shiny glabrous area at
confluence of intervals 4 to 6 (Fig. 21). Elytra with humeri distinct; lateral
margins more or less straight in basal one-half (Fig. 21)
S. immaculatus Anderson, p. 460
Quaest. Ent., 1987,23 (4)
456
Anderson
r Dorsal surface of elytra faintly to distinctly maculate, with small white
scales less dense in two posteriorly directed oblique patches (one at basal
one-third from intervals 2 to 6; the other at apical one-third from intervals
2 to 4), and with small triangular shiny glabrous area at confluence of
intervals 4 to 6 (Figs. 20, 22-23). Elytra with humeri less distinct or
rounded; lateral margins more or less arcuate in basal one-half (Figs. 20,
22-23) 2
2 (F) Elytra fused together; thoracic wings short, approximately equal to
one-half length of elytra 3
2' Elytra not fused together; thoracic wings long, approximately equal to or
greater than length of elytra 4
3 (2) Elytra with scattered small recurved hair-like scales, most evident on
declivity from sutural interval to interval 3; with greater part of length of
sutural interval and intervals 3 and 5 elevated and convex. Female with
sternum VII flat, lacking paired tumescence. Male with median lobe
elongate; apex slightly spatulate; in lateral view narrowed from apical
one-third to apex; with at most small basal ventral tubercle (Fig. 31)
S. confusus Anderson, p. 457
3' Elytra lacking erect hair-like scales, with only the extreme base of interval
3 convex, otherwise with all intervals flat throughout their length. Female
with ventral surface of sternum VII with paired tumescence. Male with
median lobe less elongate; apex markedly spatulate; in lateral view,
narrowed from approximately midlength to apex; with large basal ventral
tubercle (Fig. 32) S. cristaticollis Csiki, p. 459
4 (2") Pronotum with subapical constriction well-defined dorsally and laterally,
with distinct moderately deep impressions laterad of median carina at
apical one-quarter laterally continuous with subapical constriction; with
apical margin broadly and moderately deeply emarginate at middle. Elytra
with tubercle at confluence of intervals 4 to 6 markedly elevated. Rostrum
robust (width at apex greater than 0.90 times length); approximately
uniform in width throughout length (Fig. 30)
S. stenothorax Anderson, p. 465
4' Pronotum with subapical constriction only laterally well-defined, lacking or
with at most only very shallow indistinct impressions laterad of median
carina; with apical margin not or only very slightly emarginate at middle.
Elytra with tubercle at confluence of intervals 4 to 6 only slightly elevated.
Rostrum more elongate-narrow (width at apex less than 0.90 times length);
slightly narrowed at midlength (Figs. 28-29) 5
5 (4') Pronotum with punctures large, deep and dense, distance between
punctures distinctly less than diameter of a puncture (Fig. 28). Female
metatarsal article 3 with large ventral pilose pads. Male with median lobe
in ventral view with apex symmetrical (Fig. 34b); internal sac lacking
median dorsal pocket and large dorsally directed paired lobes on dorsal
surface immediately anterior to midlength (Fig. 46)
New World Cleonini
457
S. parshus Anderson, p. 461
5' Pronotum with punctures small, shallow and sparse, distance between
punctures greater than diameter of a puncture (Fig. 29). Female
metatarsal article 3 entirely spinose, lacking ventral pilose pads. Male with
median lobe in ventral view with apex asymmetrical (Fig. 35b); internal
sac with median dorsal pocket and without dorsally directed lobes from
dorsal surface (Fig. 47) S. plumbeus (LeConte), p. 464
Stephanocleonus confusus Anderson, new species
(Figs. 20, 25,31,37,43,204)
Stephanocleonus plumbeus; LeConte 1876a (in part; desc., misident., mixed type series). Henshaw 1881-1882 (in part;
check., misident.). Henshaw 1885 (in part; check., misident.). Wickham 1902 (check., misident.). Wickham 1909
(check., misident.). Leng 1919 (in part; check., poss. misident.). Leng 1920 (in part; catal., misident.). Danks 1981 (in
part; check., poss. misident.).
Cleonus cristatus; Casey 1891 (key, misident.).
Cleonus ( Stephanocleonus ) plumbeus ; Fall and Cockerell 1907 (in part; check., misident.).
Stephanocleonus cristatus ; LeConte 1878 (distn., misident.). Leng 1920 (in part; catal., misident.).
Cleonus plumbeus : Krombein 1979 (in part; prey, poss. misident.).
Cleonis cristaticollis ; O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Cleonis plumbeus; O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Type Material. — Holotype, male, labelled with a red label “HOLOTYPE”, “Spearfish Canyon/ Black Hills,
S.D./ 1 5— 18:VI:1910.”, inverted '‘‘’Cleonus/ cristatus/ Lee.”, “HOLOTYPE/ Stephanocleonus / confusus/ Anderson”
and with abdomen on card and genitaliain microvial attached to pin (USNM). Allotype, female, labelled with red label
“ALLOTYPE”, “Tp.21 Rge.6/ W.4 Mer. Alberta/ 12.V.1980/ Lot 3 BF&JL Carr”, “ALLOTYPE/ Stephanocleonus/
confusus/ Anderson” and with abdomen on card and genitalia in microvial attached to pin (CNCI). Type locality,
Spearfish Canyon, Black Hills, South Dakota.
Paratypes. 37 males, 29 females. CANADA: Alberta; Township 1, Range 4, West 4 Meridian, 23.V.82, B.F. and J.L,
Carr, 1M (RSAN); Township 13, Range 14, West 4 Meridian, 16. IV. 82, B.F. and J.L. Carr, 1M (JLCC); Township 14,
Range 25, West 4 Meridian, 15.V.82, B.F. and J.L. Carr, IF (JLCC); Township 10, Range 2, West 5 Meridian, 11.V.82,
B.F. and J.L. Carr, IF (RSAN); Township 30, Range 21, West 4 Meridian, 17.IV.82, B.F. and J.L. Carr, 1M (RSAN);
Medicine Hat, 22.IV.29, F.S. Carr, IF (UASM), 13. VI. 32, F.S. Carr, 2M, IF (UASM), 24.VI.30, F.S. Carr, IF
(UASM), 27. III. 27, F.S. Carr, IF (UASM), 14.IV.26, F.S. Carr, IF (UASM), 13.VI.33, F.S. Carr, IF (UASM),
2. VII. 23, F.S. Carr, 1M (UASM), 11.V.32, F.S. Carr, 1M (UASM), 13.V.27, F.S. Carr, 1M (RSAN), 13.V.23, F.S.
Carr, 1M (UASM), 6.IX.27, F.S. Carr, IF (UASM), 12.VII.24, F.S. Carr, IF (RSAN), 18.11.24, F.S. Carr, IF
(UASM), 8.V.65, J.L. Carr, IF (JLCC); Township 14, Range 13, West 4 Meridian, 22.VII.79, J.L. Carr, 1M (JLCC);
Ghost Dam, 2.VII.80, B.F. and J.L. Carr, IF (JLCC); Seebe, 7.VI.73, B.F. and J.L. Carr, 1M (JLCC); Suffield,
20. IV. 23, IF (UASM); Township 6, Range 1, West 4 Meridian, 20.V.80, B.F. and J.L. Carr, IF (JLCC); Township 10,
Range 2, West 5 Meridian, 28. IV. 81, B.F. and J.L. Carr, 1M (JLCC); Township 15, Range 14, West 4 Meridian, 9.V.80,
B.F. and J.L. Carr, IF (JLCC); 27mi. sw. Nordegg, 13. VI. 66, R. Frietag/T.L. Erwin, IF (CWOB); Calgary, IV. 11,
Criddle, IF (CNCI); Lethbridge, 6.V.30, J.H. Pepper, 1M (CNCI). Northwest Territories: 1M (CNCI). Saskatchewan:
Township 6, Range 2, West 3 Meridian, 13. VIII. 73, B.F. and J.L. Carr, 1M (RSAN); Cypress Hills Provincial Park,
29.V.63, Cook et al„ 1 F (CNCI).
UNITED STATES OF AMERICA: Arizona: Coconino County, 2.4mi. n. Kaibab Lodge, 28. VII. 71, Lawton/Willis,
2M (CWOB). Colorado: Grant/ Jackson County, Rabbit Ears Pass, 2872 me., 19.VI.81, M. Kaulbars, 1M (RSAN); Top
of Range below Sapello and Pecos Rivers, 11,000 ft., 1. VIII. 00, H.F. Wickham, 1M (USNM); Cumbres Pass, 10,000 ft.,
22.VI.35, E.C. Van Dyke, 1M (CASC); lmi. n. Nederland, 11.VIII.73, C.W. O’Brien, 1M (CWOB); Longs Peak,
10-11,000 ft,, 8.VII.26, E.C. Van Dyke, 1M (CASC); Argentine Road, H.F. Wickham, 1M (USNM); Kenosha Pass,
16.VII.38, J.W. Green, IF (CASC); Gore Pass, 15.IX.48, O. Bryant, IF (CASC); Leadville, 7-14.VII.96, H.F. Wickham,
1M (USNM). Montana: Helena, Hubbard and Schwarz, 2M (USNM); Flathead Lake, 12. VII. 11, IF (USNM); Gallatin
County, 9. IV. 32, 1M (MSU), 8.V.32, IF (MSU), 1VI.31, IF (MSU); Bozeman, 17.V.28, 1M (MSU), 11.V.25, 1M
(MSU); 15mi. ne. Bozeman, 30. VI. 72, Burleson, 1M (MSU); 6mi. e. Dell, 30.VI.54, Anderson, 1M (MSU). Nevada:
White Pine County, 17.7mi. n., 3.5mi. nw. McGill, Monte Neva Hot Springs, 25. VII. 71, Lawton and Willis, 1M, IF
(CWOB). New Mexico: IF (MCZ); Cloudcroft, V.18, 1M (CASC). Utah: Alta, 29. VI. 47, O. Bryant, IF (CASC).
Wyoming: IF (USNM); Beulah, 21. VII., T.D.A. Cockerell, 1M (UASM); Park County, Beartooth Plateau, 6.VII.61,
S.M. Sutton, 1M (UCM), 31. VIII. 59, J.G. Edwards, IF (CWOB); Beartooth Plateau, Hairgrass Canyon, 7. VII. 59, IF
(CWOB); Bighorn Mountains, Medicine Mountain, 18.VIII.62, J.G. Edwards, IF (CWOB); Tie Siding, VIII.25, IF
(USNM).
Quaest. Ent., 1987, 23 (4)
458
Anderson
Notes about synonymy. — An individual of this species from New Mexico was included in
the original type series of S. plumbeus LeConte. Stephanocleonus plumbeus as herein
recognized does not occur in New Mexico.
Derivation of specific epithet. — From the Latin “ confusio ” meaning mixed or confused.
This name is used in reference to the previous confusion of this species with S. plumbeus.
Problems in recognition. — This is the only species of North American Stephanocleonus in
which the elytra have small recurved hair-like scales on the dorsal surface and in which elytral
intervals 3 and 5 are markedly elevated and convex throughout the greater part of their lengths.
The internal sac of the male aedeagus is also distinctive (Fig. 43).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 1. Size. Length, male, 9.2-15.0 mm;
female, 10.5-14.8 mm. Width, male, 4. 7-6. 5 mm; female, 4. 9-7. 2 mm. Head. Frons with punctures large, sparse, shallow,
not confluent; also with small broad appressed white scales, dense laterally, sparse medially. Some specimens with slightly
raised carina on vertex. Rostrum. Moderately robust, slightly more so in males (width at apex 0.85-0.95 times length in
male; 0.810-0.909 in female) (Fig. 25). With well-developed, sharp, steeply declivous, low, median carina from above point
of antennal insertion to base of rostrum; both basally and apically terminated at small, moderately deep fovea (Fig. 25).
Rostrum flat from immediately laterad of median carina to lateral margins, lateral margins sharp from above point of
antennal insertion to anterior margin of eye, indicated in few specimens by slightly raised rounded carina. Dorsal
punctures large, sparse, shallow, not confluent. With scales small broad appressed white, dense laterally, sparse medially.
Pronotum. Dorsal apical margin not or shallowly emarginate at middle. With low broad indistinct median carina in
anterior one-half. Subapical constriction well defined laterally, not so dorsally; some specimens with shallow impressions
laterad of median carina, but discontinuous with lateral portion of subapical constriction. Median basal area narrowly,
moderately deeply impressed. Dorsal punctures moderately large to large, dense and moderately deep; medially, distance
between punctures markedly less than diameter of single puncture, some punctures confluent and irregularly impressed;
laterally, the distance between punctures subequal to or greater than diameter of an individual puncture. Prosternum.
With shallow impression anterior to each procoxal cavity, and with slightly to moderately developed transverse swelling
anterior to each impression. Elytra. Robust in general form (width at midlength 0.64-0.72 times length in male; 0.62-0.71
in female) (Fig. 20). In dorsal view with lateral margins evenly arcuate from midlength to base, humerus rounded, not
distinct (Fig. 20). Sutural interval and intervals 3 and 5 (in some specimens also basal one-half of interval 7) elevated and
convex throughout the greater part of their lengths. Scales white small moderately dense; smaller and less dense in two
posteromedially directed oblique patches, one at basal one-third from intervals 2 to 6, the other at apical one-third from
intervals 2 to 4. With slightly elevated triangular glabrous shiny area at apical one-quarter at confluence of intervals 4 to 6
(Fig. 20). With scattered very short recurved hair-like scales, most evident on declivity from sutural interval to interval 3.
Wings. Short (more or less equal to one-half length of elytra). Legs. Foretibia of female with inner margin with small
denticles in apical one-half; subapical tooth small to moderately large, indistinct to distinct from apical denticles.
Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal
one-quarter to one-third, divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of
foretarsus of male present as small elongate-narrow pads on apical one-quarter of article 1, as large elongate-oval pads on
apical three-quarters of article 2, and as large elongate-oval pads on more or less entire ventral surface of article 3; of
female, pilose vestiture lacking from article 1, lacking to present as elongate-narrow pads on apical one-quarter to one-half
of article 2, as elongate-oval pads on apical one-half of article 3; of mesotarsus of male, lacking to as apical tufts on outer
lobe of article 1, as elongate-narrow pads on apical three-quarters on outer lobe and as apical tufts on inner lobe of article
2, as large elongate-oval pads on apical three-quarters of article 3; of mesotarsus of female, lacking from article 1, lacking
from to as elongate-narrow pads on apical one-half on outer lobe of article 2, as small to large elongate-oval pads on apical
one-third to two-thirds of article 3; of metatarsus of male, lacking from articles 1 and 2, as elongate-oval pads on apical
one-quarter to three-quarters of article 3; of metatarsus of female, lacking from articles 1 to 3 to as elongate-oval pads
covering apical one-third of article 3. Abdomen. Abdominal sternum VII in female evenly rounded, not tumescent; in male
with apical margin slightly emarginate medially. Genitalia. Female (four examined). Abdominal sternum VIII with basal
arm expanded at apex; lateral arms narrow, slightly inwardly and evenly arcuate throughout length (Fig. 37). Male (12
examined). Abdominal sternum VIII with paired sclerite with inner apices truncate (Fig. 31c). Aedeagus elongate-narrow;
in lateral view slightly and evenly arcuate throughout length, narrowed from approximately apical one-third to apex, apex
slightly spatulate; in ventral view with apex symmetrical, medially produced into acuminate apical projection (Figs.
31a,b). Base of aedeagus with at most small ventral tubercle. Internal sac elongate with large dorsally directed paired lobe
on dorsal surface at apical one-third; median dorsal pocket high, with large laterally directed paramedial lobes at basal
one-third near ventral margin (Fig. 43). In lateral view with basal paired sclerite and apical sclerite complex widely
separated (Fig. 43a).
Geographic distribution. — This species is known from southern Alberta and southern
Saskatchewan, south to Arizona and New Mexico (Fig. 204). Most published records of S.
cristaticollis refer to this species.
New World Cleonini
459
Natural history. — Adults of this species have been collected in prairie habitats in southern
Alberta to alpine habitats at elevations of 3400 m in Colorado and New Mexico.
Chorological relationships. — This species is broadly sympatric with S', cristaticollis and S.
parshus, and narrowly sympatric with S. immaculatus along the western limits of the range of
the latter.
Stephanocleonus cristaticollis Csiki
(Figs. 26, 32, 38, 44, 205)
Stephanocleonus cristatus LeConte 1876a: 147 [not Chevrolat 1873:98]. Holotype (examined), female, labelled “Utah”,
“S./ cristatus/ Lee”, “Horn Coll/ H8514” and with my label indicating it as the holotype “Stephanocleonus/ cristatus
LeConte/ HOLOTYPE labelled/ by Anderson” (MCZC). Type locality, Utah. LeConte 1878 (distn., misident. of S.
confusus). Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Leng 1920 (in part; catal., misident.
Stephanocleonus confusus).
Cleonus ( Stephanocleonus ) cristatus ; Casey 1891 (key, misident. Stephanocleonus confusus). Csiki 1934 (catal.).
Cleonus ( Stephanocleonus ) cristaticollis Csiki 1934:26. New name for Stephanocleonus cristatus LeConte.
Cleonis cristatus ; O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Cleonis cristaticollis', O’Brien and Wibmer 1982 (in part; catal., distn., misident. Stephanocleonus confusus).
Problems in recognition. — This is one of two brachypterous species of North American
Stephanocleonus. Unlike S. confusus individuals, in which elytral intervals 3 and 5 are elevated
and convex throughout the greater part of their lengths, individuals of S. cristaticollis have
only the extreme base of interval 3 slightly elevated and convex. They also do not possess the
short recurved hair-like scales on the dorsal surface of the elytra that are found in S. confusus.
Abdominal sternum VII of females has a pair of low rounded swellings, and the shape of the
male aedeagus and form of the internal sac are distinctive (Figs. 32, 44).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 2. Size. Length, male, 13.8 mm;
female, 13.6-14.1 mm. Width, male, 6.4 mm; female, 6. 4-6. 6 mm. Head. Frons with punctures large, sparse, shallow,
indistinct; also with small elongate appressed white scales, dense laterally, sparse medially. Rostrum. Moderately robust
(width at apex 0.86 times length in male; 0.91 in female) (Fig. 26). With well-developed, sharp, steeply declivous, low,
median carina from above point of antennal insertion to base of rostrum; both basally and apically terminated at small,
moderately deep fovea (Fig. 26). Rostrum with shallow trough immediately laterad of median carina, lateral margins
sharp from above point of antennal insertion to anterior margin of eye, indicated by slightly raised rounded carina. Dorsal
punctures large, sparse, shallow, indistinct. With scales small, elongate, appressed, white, dense laterally, sparse medially.
Pronotum. Dorsal apical margin shallowly emarginate at middle. With low broad indistinct median carina in anterior
one-half. Subapical constriction well defined laterally, not so dorsally; with shallow impressions laterad of median carina,
but discontinuous with lateral portion of subapical constriction. Median basal area broadly, moderately deeply impressed.
Dorsal punctures small, sparse and shallow; medially and laterally with distance between punctures subequal to or greater
than diameter of single puncture; punctures irregularly impressed and dense in median basal area. Prosternum. With only
shallow impression anterior to each procoxal cavity. Elytra. Moderately robust in general form (width at midlength 0.63
times length in male; 0.65 in female). In dorsal view with lateral margins evenly arcuate from midlength to base, humerus
rounded, not distinct. Base of interval 3 swollen and convex, otherwise elytral intervals uniformly flat. Scales white small
moderately dense; smaller and less dense in two posteromedially directed oblique patches, one at basal one-third from
intervals 2 to 6, the other at apical one-third from intervals 2 to 4. With slightly elevated triangular glabrous shiny area at
apical one-quarter at confluence of intervals 4 to 6. Suberect or erect vestiture lacking. Wings. Short (more or less equal to
one-half length of elytra). Legs. Foretibia of female with inner margin with small denticles in apical one-half; subapical
tooth small, distinct from apical denticles. Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus
small. Tarsal claws connate in basal one-quarter to one-third, divergent, each with basal internal flange well-developed.
Ventral tarsal pilose vestiture of foretarsus of male present as small elongate-oval pads on apical one-third of article 1, as
large elongate-oval pads on apical three-quarters of article 2, and as large elongate-oval pads on more or less entire ventral
surface of article 3; of female, pilose vestiture lacking from article 1, present as apical tufts of article 2, as large
elongate-oval pads on apical three-quarters of article 3; of mesotarsus of male, as elongate-narrow pads on apical
one-quarter of outer lobe, lacking from inner lobe of article 1, as elongate-narrow pads on apical three-quarters of article 2,
as large elongate-oval pads on more or less entire ventral surface of article 3; of mesotarsus of female, lacking from articles
1 and 2, as elongate-narrow pads covering apical three-quarters of article 3; of metatarsus of male, lacking from articles 1
Quaest. Ent., 1987,23 (4)
460
Anderson
and 2, as elongate-oval pads on apical three-quarters of article 3; of metatarsus of female, lacking from articles 1 to 3.
Abdomen. Abdominal sternum VII in female with pair of rounded swellings at midlength; in male with apical margin
slightly emarginate medially. Genitalia. Female (one examined). Abdominal sternum VIII with basal arm slightly
expanded at apex; lateral arms broad, slightly inwardly and evenly arcuate throughout length (Fig. 38). Male (two
examined). Abdominal sternum VIII with paired sclerite with inner apices acuminate (Fig. 32c). Aedeagus moderately
robust; in lateral view markedly arcuate, more so in basal one-half, narrowed gradually from midlength to apex, apex
markedly spatulate; in ventral view with apex symmetrical, medially produced into elongate rounded apical projection
(Figs. 32a, b). Base of aedeagus with large ventral tubercle (Fig. 32a). Internal sac globose with dorsally directed median
lobe on dorsal surface; with moderately large dorsolaterally directed paired lobe at midlength near dorsal margin (Fig. 44).
In lateral view with basal paired sclerite and apical sclerite complex approximate (Fig. 44a).
Geographic distribution — This species is known from only four specimens: two males and a
female from southern Alberta (Medicine Hat, Monarch, Gorge Creek), and the female
holotype from Utah (no further locality data given) (Fig. 205). Most published records of this
species refer to S. confusus.
Chorological relationships. — The distribution of the few known specimens indicates that
this species is probably broadly sympatric with S. confusus and narrowly sympatric with
western S. parshus.
Stephanocleonus immaculatus Anderson, new species
(Figs. 21,27, 33, 39,45, 202)
Cleonus ( Stephanocleonus ) plumbeus4, Fall and Cockerell 1907 (in part; check., misident.). Csiki 1934 (in part; catal.,
misident.).
Stephanocleonus plumbeus-, Blatchley and Leng 1916 (in part; key, redesc., misident.). Leng 1919 (in part; check., poss.
misident.). Leng 1920 (in part; catal., misident.). Danks 1981 (in part; check., poss. misident.).
Cleonus plumbeus-, Leonard 1926 (in part; check., poss. misident.). Krombein 1979 (in part; prey, poss. misident.).
Cleonis plumbeus-, O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “McMurray,/ Alta. V. 10.53/ W.J. Brown”,
“HOLOTYPE/ Stephanocleonus/ immaculatus / Anderson” and with abdomen on card and genitalia in microvial
attached to pin (CNCI). Allotype, female, with a red label “ALLOTYPE”, “Gillam Man./ 10. VI. 1949/ J.B. Wallis”,
“ALLOTYPE/ Stephanocleonus / immaculatus/ Anderson” and with abdomen on card and genitalia in microvial
attached to pin (CNCI). Type locality, Fort McMurray, Alberta.
Paratypes. 10 males, 16 females. CANADA: Alberta: Township 93, Range 10, West 4 Meridian, 1.VI.85, B.F. and
J.L. Carr, IF (JLCC); McMurray, 21.VI.53, W.J. Brown, IF (CNCI), 29.VII.53, G.E. Ball, 1M (RSAN), 22.VI.53, G.E.
Ball, IF (CNCI); Lac La Biche, NE shore, 30.VI.63, L.M. Kenakin, IF (CWOB); Township 37, Range 18, West 5
Meridian, 18. VII. 73, B.F. and J.L. Carr, IF (RSAN); Township 35, Range 18, West 5 Meridian, 18. VIII. 67, B.F. and
J.L. Carr, IF (JLCC); Exshaw, 27.VI.54, B.F. and J.L. Carr, 1M, IF (JLCC); High Prairie, 26.VII.61, A.R. Brooks, IF
(CNCI). British Columbia: Canal Flats, 13.V.82, B.F. and J.L. Carr, 1M (RSAN). Manitoba: Gillam, 16.VI.50, F.
McAlpine, IF (CNCI); Aweme, 31.V.10, N. Criddle, IF (USNM); Beaujou [sic], 24.VI.61, H.E. Milliron, 1M (CNCI).
Northwest Territories: Fort Smith, 17.V.50, J.B. Wallis, IF (CNCI), 16.VI.50, W.G. Helps, IF (CNCI). Ontario: Lake
Superior, Whitefish Point, 46°38‘N 84°33‘W, Hubbard and Schwarz, 1M, IF (USNM). Saskatchewan: Junction
Highways 2 and 165 east, 22.VI.85, B.F. and J.L. Carr, 1M, IF (JLCC); Hudson Bay, 9.VII.54, Brooks-Wallis, 1M, IF
(CNCI).
UNITED STATES OF AMERICA: Colorado: Douglas County, 19.IV. 15, 1M (CASC). Minnesota: IF (USNM).
Nebraska: Lincoln, 29. VI., Wolcott, IF (UNSM). South Dakota: Englewood, Haggard, 1M (UNSM).
Derivation of specific epithet.— From the Latin “/m” meaning not, and “ macula ” meaning
spot or mark. This name is used in reference to the lack of distinct elytral markings in most
individuals of this species.
Problems in recognition. — This is the only species of North American Stephanocleonus in
which the elytra are more or less uniformly covered with small white scales and in which most
specimens lack a distinct pattern of elytral maculations (Fig. 21). Most specimens also lack the
small triangular shiny glabrous area at the confluence of intervals 4 to 6 which is found in all
other North American Stephanocleonus (Fig. 21). The shape of the male aedeagus and form of
the internal sac are also distinctive (Figs. 33, 45).
New World Cleonini
461
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 3. Size. Length, male, 13.3-15.6 mm;
female, 14.8-16.8 mm. Width, male, 5. 9-6. 8 mm; female, 6. 8-7. 5 mm. Head. Frons with punctures large, moderately
dense, moderately deep, many punctures longitudinally confluent, appearance longitudinally rugose; also with small broad
appressed white scales, sparse laterally, absent to very sparse medially. Most specimens with slightly raised carina on
vertex. Rostrum. Moderately robust, slightly more so in females (width at apex 0.80-0.95 times length in male; 0.79-0.96
in female) (Fig. 27). With well-developed, sharp, steeply declivous, high, median carina from above point of antennal
insertion to base of rostrum; both basally and apically terminated at small, moderately deep fovea (Fig. 27). Rostrum flat
from immediately laterad of median carina to lateral margins, lateral margins rounded to sharp from above point of
antennal insertion to anterior margin of eye. Dorsal punctures large, moderately dense, moderately deep, many punctures
confluent. With scales small uniformly dense, elongate-narrow appressed, white. Pronotum. Dorsal apical margin not or
shallowly emarginate at middle. With low broad distinct median carina in anterior one-half. Subapical constriction slightly
to well defined laterally, not so dorsally; with shallow impressions laterad of median carina, but discontinuous with lateral
portion of subapical constriction. Median basal area narrowly, shallowly impressed. Dorsal punctures moderately large,
dense and moderately deep; medially, the distance between punctures less than diameter of individual puncture, some
punctures confluent or nearly so and irregularly impressed; laterally, distance between punctures subequal to diameter of
individual puncture. Prosternum. With only very shallow impression anterior to each procoxal cavity. Elytra. Moderately
robust in general form (width at midlength 0.58-0.68 times length in male; 0.56-0.67 in female) (Fig. 21). In dorsal view
with lateral margins straight, slightly convergent from slightly past midlength to base, humerus acute, very distinct (Fig.
21). All elytral intervals more or less flat. Scales white, small, dense, no distinct maculations present (Fig. 21). Some
specimens with a slightly elevated, small, triangular, glabrous, shiny area at apical one-quarter at confluence of intervals 4
to 6. Suberect or erect vestiture lacking. Wings. Long (greater than length of elytra). Legs. Foretibia of female with inner
margin with small denticles in apical one-half; subapical tooth moderately large, distinct from apical denticles. Foretibial
and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-quarter to
one-third, divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus of male
elongate-narrow pads on apical one-quarter to one-half of article 1, as large elongate-oval pads on apical three-quarters to
more or less entire ventral surface of articles 2 and 3; of female, pilose vestiture lacking from article 1, as apical tufts to
elongate-narrow pads on apical one-half of article 2, as elongate-oval pads on apical three-quarters of article 3; of
mesotarsus of male, lacking from, to as an elongate-narrow pad on apical one-third of outer lobe of article 1, as
elongate-narrow pads on apical three-quarters of article 2, as large elongate-oval pads on apical three-quarters to more or
less entire ventral surface of article 3; of mesotarsus of female, lacking from article 1, lacking from, to as elongate-narrow
pad on apical one-half of outer lobe of article 2, as large elongate-oval pads on apical two-thirds of article 3; of metatarsus
of male, lacking from articles 1 and 2, as elongate-narrow pads on apical two-thirds of article 3; of metatarsus of female,
lacking from articles 1 and 2, as apical tufts to small elongate-oval pads on apical one-half of article 3. Abdomen.
Abdominal sternum VII in female evenly rounded, not tumescent; in male with apical margin slightly emarginate
medially. Genitalia. Female (five examined). Abdominal sternum VIII with basal arm expanded at apex; lateral arms
broad, inwardly arcuate at midlength (Fig. 39). Male (six examined). Abdominal sternum VIII with paired sclerite with
inner apices rounded (Fig. 33c). Aedeagus elongate-narrow; in lateral view slightly and evenly arcuate throughout length,
narrowed from approximately apical one-third to apex, apex slightly spatulate; in ventral view with apex symmetrical,
medially produced into truncate apical projection (Figs. 33a,b). Base of aedeagus with at most small ventral tubercle (Fig.
33a). Internal sac elongate with small laterally directed paired lobe near ventral margin at or slightly beyond midlength;
median dorsal pocket high, with large dorsolaterally directed paired lobe at basal one-third near ventral margin, separated
from base of median pocket by distinct trough (Fig. 45). In lateral view with basal paired sclerite and apical sclerite
complex widely separated (Fig. 45a).
Geographic distribution. — This species is known from widely scattered localities in the
southwestern Northwest Territories, northern Manitoba, and central Ontario, south in the west
to Colorado and Nebraska (Fig. 202).
Natural history. — A single adult has been collected on Rosa sp. (Rosaceae).
Chorological relationships. — his species is broadly sympatric throughout the northern part
ofits range with S. plumbeus and S. parshus, and is narrowly sympatric in the western part of
its range with S. confusus.
Stephanocleonus parshus Anderson, new species
(Figs. 28, 34, 40, 46, 203)
Cleonus ( Stephanocleonus ) plumbeus-, Casey 1891 (key). Csiki 1934 (in part; catal., misident.). Kissinger 1964 (biol.,
misident.).
Stephanocleonus plumbeus-, Blatchley and Leng 1916 (in part; key, redesc., misident.). Leng 1919 (in part; check., poss.
Quaest. Ent., 1987, 23 (4)
462
Anderson
misident.). Leng 1920 (in part; catal., misident.). Marcovitch 1923 (biol., redesc., larvae, misident.). Danks 1981 (in
part; check., poss. misident.).
Cleonus plumbeus-, Leonard 1926 (in part; check., poss. misident.). Bleasdell 1937 (check., prob. misident.). Krombein
1979 (in part; prey, poss. misident.).
Cleonis plumbeus ; O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “Chalk River, Ont./ May 16, 1938/ N.R.
Brown”, “HOLOTYPE/ Stephanocleonus / parshus/ Anderson” and with abdomen on card and genitalia in microvial
attached to pin (CNCI). Allotype, female, with a red label “ALLOTYPE”, “Tp. 36 Rge. 15/ W.5 Mer Alberta/
16. VII. 1973/ B&J Carr Lot 3”, “ALLOTYPE/ Stephanocleonus/ parshus/ Anderson” and with abdomen on card and
genitalia in microvial attached to pin (CNCI). Type locality, Chalk River, Ontario.
Paratypes. 39 males, 43 females. CANADA: Alberta: Township 24, Range 8, West 5 Meridian, 8. VII. 82, B.F. and
J.L. Carr, 1M (RSAN); Bragg Creek, 31.V.75, F.A.H. Sperling, 1M, IF (RSAN); Pincher Creek, 20.VI.61, H.E.
Milliron, 1M (CNCI); McMurray, 4.VI.53, W.J. Brown, 1M (CNCI), 6.VI.53, W.J. Brown, IF (CNCI); Township 25,
Range 3, West 5 Meridian, 7. VI. 73, B.F. and J.L. Carr, 2M (JLCC, RSAN); Township 29, Range 5, West 5 Meridian,
28.VI.64, B.F. and J.L. Carr, 1M (JLCC); Calgary, 28.VI.64, B.F. and J.L. Carr, 1M (RSAN); Exshaw, 27.VI.54, B.F.
and J.L. Carr, 1M (JLCC), 19.V.60, J.L. Carr, 1M (JLCC); Willow Creek, 9.VIII.28, H. Richmond, 1M (CASC); Ghost
Dam, B.F. and J.L. Carr, IF (JLCC); Edmonton, 26.V.19, F.S. Carr, 1M, IF (UASM), 14.V.20, F.S. Carr, IF (UASM),
13.V.18, F.S. Carr, 1M (UASM); Medicine Hat, 13. VI. 32, F.S. Carr, 2M, IF (UASM); Crow‘s Nest Pass, 9. VI. 30, J.H.
Pepper, 1M (CNCI); Beaverlodge, IF (UASM); near Josephburg, 53°41‘N 113°55‘W, 23.V.79, K. Shaw and J. Sutcliffe,
IF (UASM). British Columbia: Rolla, 21. VII. 27, Vroom, IF (CASC); Pouce Coupe, 1. VII. 27, Vroom, IF (CASC);
Baynes Lake, 9.V.76, B.F. and J.L. Carr, IF (JLCC); Canal Flats, 13.V.82, B.F. and J.L. Carr, 1M, IF (JLCC);
Skookumchuck, 30.V.84, B.F. and J.L. Carr, 1M (JLCC). Manitoba: Aweme, 12.VIII.29, R.H. Handford, IF (RSAN),
2. VII. 29, N. Criddle, 1M (CNCI), 20.IV.06, N. Criddle, IF (CNCI), 31.V.10, N. Criddle, IF (USNM), 9.VI.14, N.
Criddle, 1M, IF (CNCI), 24.IX.28, N. Criddle, 1M (CNCI), VI, N. Criddle, 1M (USNM), 1.VII.05, N. Criddle, IF
(USNM), V.12, N. Criddle, 1M (USNM), 3.VIII., N. Criddle, IF (USNM); Township 14, Range 105, 12.V.24, J.B.
Wallis, IF (CNCI); Riding Mountain National Park, 12.VI.38, W.J. Brown, IF (CNCI); Melita, 8.VI.20, N. Criddle, IF
(CNCI); Shell River, VII.27, E. Criddle, IF (CNCI); Winnipeg, IF (USNM); Treesbank, 24.VI.48, Criddle, IF (CNCI);
Sandilands, 22.VI.30, W.J. Brodie, IF (CNCI); 24km. w. Haddashville, 27.VI.84, I. Askevold, 1M (CWOB). Northwest
Territories: Fort Wrigley, 27. IX. 29, O. Bryant, 1M (CASC); Fort Simpson, Manners Creek, 11. VI. 72, Smetana, IF
(CNCI). Ontario; Blackburn, 6.VI.32, W.J. Brown, IF (CNCI); Ridgeville, VI.35, S.D. Hicks, IF (CNCI); Toronto, 1M
(USNM); Dunrobin, 18.V.77, J.E. O’Hara, 1M (RSAN). Quebec: Kazubazua, 25.V.33, W.J. Brown, 1M (CNCI); Fort
Coulonge, 25. VI. 19, J. Beaulne, 1M (CNCI). Saskatchewan: 8mi. e. Saskatoon, 17. VI. 73, C.K. Starr, IF (CWOB);
Township 35, Range 6, West 3 Meridian, 20. VII. 85, B.F. and J.L. Carr, 1M (JLCC); Pike Lake, 5.1.42, King & Glen, IF
(CNCI); Prince Albert, 4. VII. 54, Brooks- Wallis, IF (CNCI).
UNITED STATES OF AMERICA: Connecticut: New Hartford, 26. IV. 20, Zappe, IF (CWOB). Iowa: Ames, IF
(USNM). Maine: 1M (USNM). Massachusetts: Woodshole, IF (CASC). Michigan: Emmet County, Mackinaw City,
28. VI. 20, M.H. Hatch, 1M (USNM). Minnesota: IF (CASC); Crow Wing County, Pelican Lake, 25. VIII. 10, Wolcott,
IF (UNSM), 28.VIII.08, Wolcott, IF (UNSM). Montana: Missoula, 16.V.04, IF, 11.VII.04, 1M (USNM); Helena,
Hubbard and Schwarz, IF (USNM); Flathead County, Echo Lake, 31.V.35, Eichmann, IF (MSU); Hamilton,
16.VIII.26, 1M (MSU); Lake County, 6. VII. 36, 1M (MSU). Nebraska: Lincoln, VI., IF (UNSM); Bellevue, 20.V.02,
Bruner, 1M (UNSM). New Mexico: IF (MCZC). South Dakota: Englewood, Haggard, IF (UNSM). Tennessee:
Knoxville, 12.lV. 55, H. and A. Howden, IF (HAHC), 17.V.57, H. and A. Howden, 1M (HAHC). Vermont: Bennington
County, G.H. Horn, 1M (USNM); Lyndon, 22.VIII.00, Melander, 1M, IF (OCUC).
Country unknown: Bridge, 31.V.14, IF (CNCI).
Derivation of specific epithet. — An arbitrary combination of letters.
Problems in recognition. — Among macropterous North American Stephanocleonus species
with elytral maculations, individuals of S. parshus can be recognized by the large deep and
dense pronotal punctures, the distance between punctures distinctly less than the diameter of a
puncture (Fig. 28a). As in individuals of S. plumbeus , the pronotum has the lateral subapical
constriction not defined dorsally and has the apical margin entire and not emarginate. In
addition to other features, females of S. parshus can further be distinguished from those of
other North American macropterous species by the metatarsus with article 3 with elongate
pilose pads; males can be distinguished from males of other species by the distinctive shape of
the aedeagus and form of the internal sac (Figs. 34, 46).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 4. Size. Length, male, 10.4-12.0 mm;
.female, 10.9-13.6 mm. Width, male, 4. 8-5. 6 mm; female, 5.0-5. 9 mm. Head. Frons with punctures large, dense,
moderately deep, many punctures confluent, appearance longitudinally rugose; also with small elongate-narrow appressed
New World Cleonini
463
white, dense scales. Most specimens with slightly raised carina on vertex. Rostrum. Moderately elongate-narrow, slightly
more so in males (width at apex 0.76-0.84 times length in male; 0.77-0.84 in female) (Fig. 28). With well-developed,
sharp, gradually declivous, high, median carina from above point of antennal insertion to base of rostrum; both basally and
apically terminated at small, shallow fovea (Fig. 28). Rostrum rounded to flat from immediately laterad of median carina
to lateral margins, lateral margins rounded to sharp from above point of antennal insertion to anterior margin of eye,
indicated in very few specimens by slightly raised rounded carina. Dorsal punctures large, moderately dense, moderately
deep, many confluent. With scales small, elongate-narrow, appressed, white, dense. Pronotum. Dorsal apical margin not or
shallowly emarginate at middle. With low, broad, distinct median carina in anterior one-half. Subapical constriction
slightly to well defined laterally, not so dorsally; with shallow impressions laterad of median carina, but discontinuous with
lateral portion of subapical constriction. Median basal area narrowly, shallowly impressed. Dorsal punctures large, dense
and deep; medially, most confluent and irregularly impressed; laterally, distance between punctures much less than
diameter of individual puncture (Fig. 28a). Prosternum. With shallow impression anterior to each procoxal cavity, and
with or without slightly developed transverse swelling anterior to each impression. Elytra. Moderately robust in general
form (width at midlength 0.61-0.68 times length in male; 0.59-0.65 in female) (Fig. 22). .In dorsal view with lateral
margins straight and convergent from midlength to base, humerus obtuse, distinct. All elytral intervals more or less
uniformly flat. Scales white small moderately dense; smaller and less dense in two posteromedially directed oblique
patches, one at basal one-third from intervals 2 to 6, other at apical one-third from intervals 2 to 4. With slightly elevated
triangular glabrous shiny area at apical one-quarter at confluence of intervals 4 to 6. Suberect or erect vestiture lacking.
Wings. Long (equal to or greater than length of elytra). Legs. Foretibia of female with inner margin with small denticles in
apical one-half; subapical tooth small to moderately large, indistinct to distinct from apical denticles. Foretibial and
mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-quarter to
one-third, divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus of male
present as elongate-oval pads on apical one-half of article 1, as large elongate-oval pads on apical three-quarters of article
2, and as large elongate-oval pads on more or less entire ventral surface of article 3; of female, pilose vestiture lacking from
article 1, present as elongate-oval pads on apical one-half of article 2, as elongate-oval pads on more or less entire ventral
surface of article 3; of mesotarsus of male, lacking from to present as an apical tuft of outer lobe of article 1, as
elongate-narrow pads on apical three-quarters of article 2, as large elongate-oval pads on more or less entire ventral
surface of article 3; of mesotarsus of female, lacking from article 1, lacking from to present as an elongate-narrow pad on
apical one-half of outer lobe of article 2, as large elongate-oval pads on more or less entire ventral surface of article 3; of
metatarsus of male, lacking from articles 1 and 2, as elongate-oval pads on apical one-third to one-half of article 3; of
metatarsus of female, lacking from articles 1 and 2, as an elongate-oval pad on apical one-third to one-half of outer lobe
and apical one-half to two-thirds of inner lobe of article 3. Abdomen. Abdominal sternum VII in female evenly rounded,
not tumescent; in male with apical margin slightly emarginate medially. Genitalia. Female (three examined). Abdominal
sternum VIII with basal arm expanded at apex; lateral arms narrow, inwardly arcuate at midlength (Fig. 40). Male (12
examined). Abdominal sternum VIII with paired sclerite with inner apices rounded (Fig. 34c). Aedeagus elongate-narrow;
in lateral view slightly and evenly arcuate throughout length, narrowed from approximately apical one-third to apex, apex
slightly spatulate; in ventral view with apex symmetrical, medially produced into rounded apical projection (Figs. 34a,b).
Base of aedeagus with at most small ventral tubercle. Internal sac elongate with large dorsally-directed paired lobe on
dorsal surface immediately beyond midlength; median dorsal pocket lacking, with large dorsolaterally directed paired lobe
at basal one-third near dorsal margin (Fig. 46). In lateral view with basal paired sclerite and apical sclerite complex widely
separated (Fig. 46a).
Geographic distribution. — This species is widely distributed in North America from Maine
and Massachusetts south to Tennessee, west to Nebraska and Iowa in the south, British
Columbia and Alberta in the north (Fig. 203). A single state record for New Mexico is in the
Horn collection at the Museum of Comparative Zoology, Harvard University.
Natural history. — At Knoxville, Tennessee, Marcovitch (1923) records larvae of this
species feeding on roots of strawberry in June, July, December and March. Pupae were found
on July 13 and a single adult emerged on July 25. Although identified as S. plumbeus, the
description provided notes that the thorax is coarsely punctured. This, and the verified
occurence of only S. parshus at Knoxville, indicates that this is the species to which he referred.
Chorological relationships. — This, the most widespread species of North American
Stephanocleonus, is sympatric in the northern part of its range with S. plumbeus ; in the
northeastern part of its range with S. immaculatus; and, in the western part of its range with S.
cristaticollis and S. confusus.
Quaest. Ent., 1987, 23 (4)
464
Anderson
Stephanocleonus plumbeus LeConte
(Figs. 22, 29,35,41,47,205)
Cleonus obliquus\ LeConte 1850 (check., misident.).
Stephanocleonus plumbeus LeConte 1876a:146. Lectotype (here designated), male, one of an undetermined number of
syntypes, labelled with a pale blue circle ( = Lake Superior), “Type/ 5259”, and with card with abdomen attached,
microvial containing genitalia, and my designation label “ Stephanocleonus / plumbeus LeC./ LECTOTYPE/ desig.
Anderson” (MCZC). Type locality, north shore of Lake Superior. LeConte 1876a (in part; desc., misident.
Stephanocleonus confusus, mixed type series). Henshaw 1881-1882 (in part; check., misident. Stephanocleonus
confusus). Henshaw 1885 (in part; check., misident. Stephanocleonus confusus). Wickham 1902 (check., misident.
Stephanocleonusconfusus) . Wickham 1909 (check., misident. Stephanocleonus confusus). Blatchley and Leng 1916
(in part; key, redesc., misident. Stephanocleonus immaculatus, Stephanocleonus parshus). Leng 1919 (in part;
check., poss. misident. Stephanocleonus immaculatus , Stephanocleonus parshus , Stephanocleonus confusus ,
Stephanocleonus' stenothorax). Leng 1920 (in part; catal., misident. Stephanocleonus immaculatus, Stephanocleonus
parshus, Stephanocleonus confusus). Marcovitch 1923 (biol., redesc., larvae, misident. Stephanocleonus parshus).
Danks 1981 (in part; check., poss. misident. Stephanocleonus immaculatus, Stephanocleonus parshus,
Stephanocleonus confusus, Stephanocleonus stenothorax).
Cleonus ( Stephanocleonus ) plumbeus-, Casey 1891 (key). Fall and Cockerell 1907 (check., misident. Stephanocleonus
immaculatus, Stephanocleonus confusus). Csiki 1934 (in part; catal., misident. Stephanocleonus immaculatus,
Stephanocleonus parshus, Stephanocleonus confusus).
Cleonus plumbeus-, Leonard 1926 (in part; check., poss. misident. Stephanocleonus immaculatus, Stephanocleonus
parshus). Bleasdell 1937 (check., poss. misident. Stephanocleonus parshus). Kissinger 1964 (biol., misident.
Stephanocleonus parshus). Krombein 1979 (in part; as prey, poss. misident. Stephanocleonus immaculatus,
Stephanocleonus parshus, Stephanocleonus confusus).
Cleonis plumbeus-, O’Brien and Wibmer 1982 (in part; catal., distn., misident. Stephanocleonus immaculatus,
Stephanocleonus parshus, Stephanocleonus confusus).
Problems in recognition. — Among North American Stephanocleonus species that are
macropterous and possess maculated elytra, this species can be recognized by the following
combination of characters: pronotum with lateral subapical constriction not defined dorsally;
apical margin of pronotum entire, not emarginate at middle; and pronotal punctures small,
shallow and sparse, distance between punctures greater than diameter of individual puncture
(Fig. 29a). Females of this species have the metatarsus with article 3 lacking ventral pilose
pads.
Individuals of this species are most likely to be confused with those of S. parshus, but the
two can be readily separated by characters noted in the key and above.
Notes about synonymy. — I have been unable to ascertain the exact constitution of the type
series of S. plumbeus LeConte. The species was described from an unspecified number of
specimens from the north shore of Lake Superior and from New Mexico. In the LeConte
collection at the Museum of Comparative Zoology, there are three specimens labelled as types;
two specimens with pale blue circles attached (= Lake Superior), and a single specimen
labelled “N.M.” (= New Mexico). I have selected one of the two Lake Superior specimens as
lectotype and believe that these three specimens likely initially constituted the type series of S.
plumbeus. The other specimen from Lake Superior is conspecific with the lectotype, but the
specimen labelled “N.M.” is S. confusus.
Other specimens in the LeConte and Horn collections, but not labelled as types and
therefore questionably part of the type series, include one of S. immaculatus with no locality
data; two of S. plumbeus with no locality data; and one of S. parshus from New Mexico.
Description. — Specimens examined. 13 males, 11 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 5. Size. Length, male, 10.9-13.0 mm; female, 11.2-15.6 mm. Width, male,' 4. 9-6.0 mm; female, 5. 2-7. 2 mm.
Head. Frons lacking distinct large punctures; uniformly covered with small elongate-narrow appressed white scales. Some
specimens with slightly raised carina on vertex. Rostrum. Moderately elongate-narrow to moderately robust, (width at
apex 0.82-0.90 times length in male; 0.82-0.94 in female) (Fig. 29). With well-developed, sharp, gradually declivous,
high, median carina from above point of antennal insertion to base of rostrum; basally terminated at small, shallow fovea
New World Cleonini
465
(Fig. 29a). Rostrum rounded from immediately laterad of median carina to lateral margins, lateral margins rounded from
above point of antennal insertion to anterior margin of eye. Dorsally lacking distinct large punctures. With small
elongate-narrow appressed white, dense scales. Pronotum. Dorsal apical margin not or shallowly emarginate at middle.
With sharp distinct median carina in anterior one-half. Subapical constriction slightly defined laterally, not so dorsally;
with shallow impressions laterad of median carina, but discontinuous with lateral portion of subapical constriction. Median
basal area broadly, shallowly to moderately deeply impressed. Dorsal punctures uniformly small, sparse and shallow,
distance between punctures greater than diameter of one puncture (Fig. 29a). Prosternum. With shallow impression
anterior to each procoxal cavity, and slightly developed transverse swelling anterior to each impression. Elytra. Moderately
robust in general form (width at midlength 0.60-0.64 times length in male; 0.57-0.62 in female) (Fig. 22). In dorsal view
with lateral margins straight and convergent from midlength to base, humerus obtuse, distinct (Fig. 22). All elytral
intervals more or less uniformly flat. Scales white small moderately dense; smaller and less dense in two posteromedially
directed oblique patches, one at basal one-third from intervals 2 to 6, the other at apical one-third from intervals 2 to 4
(Fig. 22). With slightly elevated triangular glabrous shiny area at apical one-quarter at confluence of intervals 4 to 6.
Suberect or erect vestiture lacking. Wings. Long (equal to or greater than length of elytra). Legs. Foretibia of female with
inner margin with small denticles in apical one-half; subapical tooth small to moderately large, indistinct to distinct from
apical denticles. Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws
connate in basal one-quarter to one-third, divergent, each with basal internal flange well-developed. Ventral tarsal pilose
vestiture of foretarsus of male present as elongate-oval pads on apical one-third of article 1, as large elongate-oval pads on
apical three-quarters of article 2, and as large elongate-oval pads on more or less entire ventral surface of article 3; of
female, pilose vestiture lacking from articles 1 and 2, present as elongate-oval pads on apical three-quarters of article 3; of
mesotarsus of male, lacking from to as apical tuft of outer lobe of article 1, as elongate-narrow pads on apical
three-quarters of article 2, as large elongate-oval pads on apical three-quarters of article 3; of mesotarsus of female,
lacking from articles 1 and 2, as small elongate-narrow pads on apical one-third of article 3; of metatarsus of male, lacking
from articles 1 and 2, as elongate-oval pads on apical three-quarters of article 3; of metatarsus of female, lacking from
articles 1 to 3. Abdomen. Abdominal sternum VII in female evenly rounded, not tumescent; in male with apical margin
slightly emarginate medially. Genitalia. Female (three examined). Abdominal sternum VIII with basal arm expanded at
apex; lateral arms narrow, inwardly arcuate at midlength (Fig. 41). Male (seven examined). Abdominal sternum VIII
with paired sclerite with inner apices truncate (Fig. 35c). Aedeagus elongate-narrow; in lateral view markedly arcuate,
more so in basal one-half, narrowed from approximately apical one-third to apex, apex slightly spatulate; in ventral view
with apex asymmetrical, produced into rounded apical projection (Figs. 35a, b). Base of aedeagus with at most small
ventral tubercle. Internal sac elongate with small dorsolaterally directed paired lobe on dorsal surface immediately beyond
midlength; median dorsal pocket high, with large dorsolaterally directed paired lobe at basal one-third near ventral
margin, separated from base of median pocket by trough (Fig. 47). In lateral view with basal paired sclerite and apical
sclerite complex widely separated (Fig. 47a).
Geographic distribution. — This species is known from widely scattered localities from
Newfoundland west to Alberta (Fig. 205).
Chorological relationships. — This species is sympatric throughout the central and western
part of its range with S. parshus and S. immaculatus. Individuals of these three species have
been collected within a few days of each other at Fort McMurray, Alberta.
Stephanocleonus stenothorax Anderson, new species
(Figs. 23, 30, 36, 42, 48, 204)
Stephanocleonus plumbeus\ Leng 1919 (in part; check., poss. misident.). Danks 1981 (in part; check., poss. misident.).
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “YT. Bluefish caves/ 67°08‘N 140°48‘W/
2000‘ 4.vii.l983/R.J. Cannings”, “HOLOTYPE/ Stephanocleonus / stenothorax / Anderson” and with abdomen on card
and genitalia in microvial attached to pin (CNCI). Allotype, female, with a red label “ALLOTYPE”, “YUKON: Dog.
Ck./ 30. VII. 77/ R.E. Roughley”, “ALLOTYPE/ Stephanocleonus / stenothorax/ Anderson” and with abdomen on card
and genitalia in microvial attached to pin (CNCI). Type locality, Bluefish Caves archaeological site, Yukon Territory.
Paratypes. 12 males, 1 1 females. Canada: Yukon Territory, Keele Range, Bluefish Caves archaeological site, 67°08‘N
140°47‘W, 1983, J. Cinq-Mars, 12M, 1 IF. (CNC, CWOB, JVM, RSAN, USNM, ZIL)
Derivation of specific epithet. — From the Greek “ stenos ” meaning narrowed, and “ thorax ”
meaning breastplate. This name is used in reference to the apically narrowed form of the
pronotum.
Problems in recognition. — Among North American species of Stephanocleonus with
maculate elytra, individuals of this species are recognized by the pronotum with the lateral
subapical constriction continued and well-defined dorsally, and with the apical margin
Quaest. Ent., 1987, 23 (4)
466
Anderson
shallowly emarginate at the middle. The glabrous shiny tubercle at the confluence of elytral
intervals 4 to 6 is markedly elevated and lateral arms of abdominal sternum VIII of the female
are very broad and medially approximate to continguous near their apices (Fig. 42). The shape
of the aedeagus of the male is also distinctive (Figs. 36a, b).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 6. Size. Length, male, 12.5-13.1 mm;
female, 12.8-14.9 mm. Width, male, 5. 3-6. 2 mm; female, 5. 6-6. 2 mm. Head. Frons with punctures large, moderately
dense and deep, many punctures confluent in some specimens, surface thus longitudinally rugose; uniformly covered with
small broad appressed white scales, dense laterally, sparse medially. Most specimens with slightly raised carina on vertex.
Rostrum. Robust, (width at apex 0.90-0.95 times length in male; 0.90-1.00 in female) (Fig. 30). With well-developed,
sharp, steeply declivous, low, median carina from above point of antennal insertion to base of rostrum; basally and apically
terminated at small, shallow fovea (Fig. 30a). Rostrum flat from immediately laterad of median carina to lateral margins,
lateral margins rounded to sharp from above point of antennal insertion to anterior margin of eye. Dorsally with punctures
large moderately dense and deep, many confluent in some specimens, surface thus longitudinally rugose. With small
elongate-narrow appressed white, scales; dense laterally, sparse medially. Pronotum. Dorsal apical margin broadly and
moderately deeply emarginate at middle. With sharp distinct median carina in anterior one-half. Subapical constriction
well defined laterally and dorsally; with distinct moderately deep impressions laterad of median carina continuous with
lateral portion of subapical constriction. Median basal area broadly and moderately deeply impressed. Dorsal punctures
small, sparse and shallow laterally and medially (except basally at middle where punctures are dense, deep, approximate
and irregularly impressed), distance between punctures subequal to or greater than diameter of one puncture. Prosternum.
With shallow impression anterior to each procoxal cavity, and with or without slightly to moderately developed transverse
swelling anterior to each impression. Elytra. Moderately elongate-narrow in general form (width at midlength 0.57-0.60
times length in male; 0.55-0.61 in female) (Fig. 23). In dorsal view with lateral margins straight and convergent from
midlength to base, humerus obtuse, distinct (Fig. 23). All elytral intervals more or less uniformly flat. Scales white small
moderately dense; smaller and less dense in two posteromedially directed oblique patches, one at basal one-third from
intervals 2 to 6, other at apical one-third from intervals 2 to 4 (Fig. 23). With markedly elevated triangular glabrous shiny
area at apical one-quarter at confluence of intervals 4 to 6. Suberect or erect vestiture lacking. Wings. Long (equal to or
greater than length of elytra). Legs. Foretibia of female with inner margin with small denticles in apical one-half;
subapical tooth moderately large, distinct from apical denticles. Foretibial and mesotibial unci of both sexes moderately
large, metatibial uncus small. Tarsal claws connate in basal one-quarter to one-third, divergent, each with basal internal
flange well-developed. Ventral tarsal pilose vestiture of foretarsus of male present as elongate-oval pads on apical one-half
of article 1 , as large elongate-oval pads on apical two-thirds of article 2, and as large elongate-oval pads on more or less
entire ventral surface of article 3; of female, pilose vestiture lacking from articles 1 and 2, present as elongate-oval pads on
apical one-half of article 3; of mesotarsus of male, lacking from to as apical tuft of outer lobe of article 1, as
elongate-narrow pads on apical two-thirds of outer lobe and apical one-half on inner lobe of article 2, as large elongate-oval
pads on apical three-quarters of article 3; of mesotarsus of female, lacking from articles 1 and 2, as elongate-oval pad on
apical one-half of outer lobe and as apical tuft of inner lobe of article 3; of metatarsus of male, lacking from articles 1 and
2, as elongate-oval pads on apical two-thirds of article 3; of metatarsus of female, lacking from articles 1 to 3. Abdomen.
Abdominal sternum VII in female evenly rounded, not tumescent; in male with apical margin slightly emarginate
medially. Genitalia. Female (two examined). Abdominal sternum VIII with basal arm expanded at apex; lateral arms very
broad, medially approximate or contiguous near apices, inwardly arcuate at midlength (Fig. 42). Male (four examined).
Abdominal sternum VIII with paired sclerite with inner apices truncate (Fig. 36c). Aedeagus elongate-narrow; in lateral
view slightly and evenly arcuate throughout length, narrowed from approximately apical one-third to apex, apex not
spatulate; in ventral view with apex symmetrical, medially rounded but not produced into distinct apical projection (Figs.
36a, b). Base of aedeagus with at most small ventral tubercle. Internal sac elongate with small dorsolaterally directed
paired lobe on dorsal surface at midlength; median dorsal pocket moderately high, with moderately large dorsolaterally
directed paired lobe at basal one-third near dorsal margin (Fig. 48). In lateral view with basal paired sclerite and apical
sclerite complex widely separated (Fig. 48a).
Geographic distribution. — This species is known only from two localities in the northern
Yukon Territory (Fig. 204).
Natural history. — Most specimens of the type series were collected in unbaited pitfall traps
set in a partly forested limestone upland and fell-field. They were collected in association with
Lepidophorus lineaticollis Kirby and Vitavitus thulius Kissinger and several species of
dry-tundra Carabidae.
Chorological relationships. — This species is allopatric to all other Stephanocleonus species.
New World Cleonini
467
Genus Apleurus Chevrolat
Apleurus Chevrolat 1873:78. Gender, masculine. Type species Apleurus fossus Chevrolat (= Cleonus lutulentus
LeConte) by subsequent designation (Casey 1891:186). Chevrolat 1873 (in part; catal., sp. desc.). Faust 1904 (key).
Notes about synonymy. — See “Notes about synonymy” section under subgenus Apleurus.
Diagnosis. — Adult Cleoninae with moderately elongate-narrow to very robust body form
(Figs. 24, 68-80). Eye elongate teardrop shaped, slightly to moderately prominent and convex
in dorsal view (Figs. 81-88). Rostrum with median carina lacking to variously developed (Figs.
84-86). Pronotum punctate, anterolateral margin of pronotum straight to with variously
developed acute postocular projection; postocular vibrissae long, of unequal length and longest
behind base of eye (Figs. 81-88). Disk with elongate white moderately large moderately dense
scales present in lateral stripe of variable width, small and fine in moderately broad to very
broad apically narrowed median stripe, and variously small and fine to moderately large and
elongate along lateral margins; median area largely black in color, underlying dark cuticle' not
obscured by overlying scales (Figs. 24, 68-80). Antenna with funiculus with article 1 longer
than, to more or less equal in length to, article 2; article 2 more or less as long as wide, to
slightly longer than wide (Figs. 81-88). Ventral tarsal pilosity various but reduced to some
extent (not extended over entire ventral surface of tarsal article) to lacking entirely from at
least more basal tarsal articles (especially of metatarsus). Tibia with corbel ridge sharp (Fig.
6). Prosternum with swellings absent or present, if present, located immediately anterior to
each prosternal impression (Fig. 7). Female with abdominal sternum VIII lacking basal arm
(Figs. 89-96).
Description. — Size. Small to large, moderately elongate-narrow to very robust in body form. Mouthparts.
Prementum flat to slightly swollen, lacking or with as many as five large setae on each side (Figs. 61-63). Maxillary
palpus with palpifer with large seta, stipes with or without large seta (Figs. 64-65). Labial palpi separated by a distance
varied from subequal to, to twice width of basal article of a labial palpus (Figs. 61-63). Rostrum. Moderately to markedly
robust, not medially tumescent (Figs. 81-88). Median carina lacking to low, moderately rounded to sharp (Figs. 81-88).
Epistoma not swollen, produced anteriorly, with apical margin emarginate (Fig. 81b) or rounded medially (Figs.
82b-88b). Antenna with funiculus with article 1 subequal in length to, to slightly longer than article 2 (Figs. 81a-88a);
apical three articles of club with placoid sensillae. Head. Eye elongate teardrop shaped, slightly to moderately prominent
and convex in dorsal view (Figs. 81-88). Area behind eye with moderately deep irregular punctures. Upper margin of eye
rounded to sharp, frons convex to more or less flat. Vestiture. Dorsum with suberect or erect vestiture lacking to very long
and dense; with simple elongate-narrow appressed white scales of variable size and density. Prothorax. Dorsal surface of
pronotum punctate. Pronotum with median basal area shallowly to deeply impressed; anterolateral margin, behind eyes,
straight to with variously developed acute postocular projection; postocular vibrissae long, of unequal length and longest
behind base of eye (maximum length greater than or equal to one-half width of eye) (Figs. 81a-88a); disk with elongate
white moderately large moderately dense scales present in lateral stripe of various width, small and fine in moderately
broad to very broad apically narrowed median stripe, and variously small and fine to moderately large and elongate along
lateral margins; median area largely black in color, underlying dark cuticle not obscured by overlying scales (Figs. 24,
68-80). Prosternum with slight to moderately deep impression anterolaterad of each procoxal cavity; with or without slight
to markedly developed swelling immediately anterior to each prosternal impression (Fig. 7). Legs. Foretarsus moderately
broad, articles 2 and 3 more or less subequal in length, at most only slightly longer than broad; article 1 only slightly longer
than articles 2 or 3; article 3 moderately deeply bilobed. Meso- and especially metatarsus slightly more elongate-narrow,
article 2 slightly to distinctly longer than article 3; article 1 distinctly longer than articles 2 or 3; article 3 moderately
deeply bilobed. Ventral tarsal pilosity various but reduced to some extent (not covering entire ventral surface of a tarsal
article) to lacking entirely from at least more basal tarsal articles (especially of metatarsus). Claws variously connate from
near base to through basal one-third, slightly to markedly divergent. Foretibia with inner margin with at most only small
denticles in apical one-half; near apex with second spur very slightly to moderately well-developed. Metatibia of male with
ventral margin of uncus evenly rounded. Tibia with corbel ridge sharp (Fig. 6). Wings. Absent or present (various in
length). Elytra. Intervals, except humerus and bases of intervals 3 and 5 flat, to with sutural interval and intervals 3, 5, 7,
and 9 variously slightly to markedly swollen and convex. Striae distinctly punctate. Humeri acute to rounded. Scale
pattern various (Figs. 24, 68-80). Abdomen. Ventral surface with small shiny glabrous patches, each with single small
appressed scale to uniformly covered with fine dense hair-like scales. Abdominal sternum VII in males with apical margin
at middle with or without small dorsally directed median tooth. Abdominal terga VII and VIII of female not, or slightly to
markedly longitudinally carinate. Genitalia. Female. Abdominal sternum VIII lacking basal arm (Figs. 89a-96a).
Quaest. Ent., 1987, 23 (4)
468
Anderson
Gonocoxite II elongate triangular in form, apex not prolonged into marked lobe; stylus absent or present, small to
moderately large in size; apical to slightly anteapical in position (Figs. 89b-96b). Spermathecal gland round (Fig. 60).
Male. Abdominal sternum VIII with interior angle of each sclerite lacking basal projection to with basal projection slightly
to markedly developed (Figs. 97f, 1 0 1 e, 103e). Aedeagus moderately robust, in lateral view more or less evenly arcuate
throughout length; apex not spatulate (Figs. 97d, 98c-104c). Internal sac various; with dorsal median pocket low to high;
apical and dorsal median pockets individually distinct; various lobes present, lobe A absent (Figs. 97a,b-104a,b). Apical
sclerite complex with individual sclerites scythe-like (Figs. 97c, 98d-104d).
Comparisons. — Species of Apleurus are most likely to be confused with the Palearctic
Chromoderus and the South African Cnemodontus. Members of these three genera either lack
pronotal postocular lobes entirely or possess variously developed acute postocular projections.
Apleurus species are separated from species of these other two genera by the ventral tarsal
pilosity reduced in extent and also by characters in the male genitalia (see “Phylogenetic
Analysis” section). Species of Pleurocleonus also lack a pronotal postocular lobe but are readily
separated from Apleurus by the possession of a low broad longitudinally sulcate median rostral
carina, flat eyes, and characters of male and female genitalia.
In the New World, some Apleurus species may be confused with large Cleonidius species
which lack a pronotal postocular lobe. Consultation of diagnoses and key characters will readly
separate the two.
Checklist of included species. — Eight species, assigned to two subgenera, are included in
this endemic New World genus. These species are:
Subgenus Gibbostethus Anderson, new subgenus
1. A.(G.) hystrix (Fall)
Subgenus Apleurus Chevrolat
2. A. (A.) aztecus (Champion)
3. A. (A.) lutulentus (LeConte)
4. A. (A.) porosus (LeConte)
5. A. (A.) angularis (LeConte)
6. A. (A.) jacobinus (Casey)
7. A. (A.) albovestitus (Casey)
8. A. (A.) saginatus (Casey)
Phylogenetic relationships. — See “Phylogenetic analysis” section.
Subgenus Gibbostethus Anderson, new subgenus
Dinocleus ; Fall 1913 (in part; sp. desc.). Leng 1920 (in part; catal.).
Cleonus ( Dinocleus ); Csiki 1934 (in part; catal.).
Cleonis\ O’Brien and Wibmer 1982 (in part; catal., distn.).
Derivation of name. — Derived from the Greek “ gibbos ” meaning protuberant or swollen,
and “ stethos ” meaning chest or breast. This name is used in reference to the tumescent
mesosternal process of members of the single species placed in this subgenus.
Type species. — Dinocleus hystrix Fall by monotypy.
Diagnosis. — Apleurus with moderately elongate-narrow body form (Fig. 24). Dorsal erect
vestiture dense and long. Prementum with bases of labial palpi separated by distance more or
less equal to twice width of basal article of labial palpus (Fig. 61). Maxillary palpus with
palpifer with large seta (Fig. 65). Rostrum with epistoma with apical margin emarginate
medially (Fig. 81b). Mesosternum with mesosternal process markedly tumescent and convex
(Fig. 58). Female with abdominal sternum VIII with lateral arms arcuate (Fig. 89a);
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469
gonocoxite II with stylus moderately large, apical in position (Fig. 89b). Male with abdominal
sternum VIII with each sclerite with rounded interior angles, basal projections lacking (Fig.
97f); aedeagus in dorsal view markedly laterally expanded from midlength to apical one-third
(Fig. 97e).
Included species. — Apleurus (Gibbostethus) hystrix Fall from southern California is the
only species in the subgenus.
Phylogenetic relationships. — This subgenus is sister to the remainder of Apleurus.
Recognition as a distinct subgenus is warranted by the primitive phylogenetic position of the
species and the sister-group relationship with the remainder of Apleurus , by a restricted and
likely relictual distribution (Fig. 212), and by various autapotypic character states and
character states exhibited by Gibbostethus which although interpreted as evolved
independently, are homoplasious in derived Apleurus species (see “Phylogeny” section).
Apleurus ( Gibbostethus ) hystrix (Fall), new combination
(Figs. 24, 58, 61, 65, 81, 89, 97, 212)
Dinocleus hystrix Fall 1913:41. Holotype (examined), male, labelled “Pasadena/ Cal.”, “Feb.”, “Not in/ Lee. Coll.”,
“TYPE”, “M.C.Z ./ Type/ 25191”, “H.C.FALL/ COLLECTION”, “ Dinocleus I hystrix/ Fall” (MCZC). Type
locality, Pasadena, California. Leng 1920 (catal.).
Cleonus ( Dinocleus ) hystrix ; Csiki 1934 (catal.).
Cleonis hystrix ; O’Brien and Wibmer 1982 (catal., distn.).
Problems in recognition. — A distinctive and easily recognized species, A. hystrix is
characterized by the markedly tumescent mesosternal process (Fig. 58) and the rostrum with
the apical margin of the epistoma emarginate medially (Fig. 81b). The markedly laterally
expanded aedeagus of males is also characteristic of this species (Fig. 97e).
Although it is unlikely that A. hystrix may be confused with other species of Apleurus ,
individuals may be confused with species of Cleonidius because of their small size and more
elongate general body form (Fig. 24). Among other characters, A. hystrix individuals can be
separated from Cleonidius individuals by the presence in the former of low rounded swellings
on the prosternum immediately anterior to the prosternal impressions (Fig. 7). Swellings may
or may not be present in Cleonidius species, but if present are situated immediately anterior to
the procoxal cavities, adjacent to the prosternal impression (Fig. 8).
Description. — Specimens examined. 18 males, 11 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 7.
Size. Length, male, 7. 0-9. 5 mm; female, 8.0-10.6 mm. Width, male, 2. 8-3. 5 mm; female, 3. 0-4. 2 mm. Head. Eye very
prominent and convex in dorsal view. Frons and vertex with dense, large, irregularly impressed, deep punctures. Frons also
with dense, very long erect white hair-like scales. Area immediately behind posterior margin of eye with very large, deep,
irregularly impressed punctures. Area above eyes slightly to moderately elevated above rest of frons (eyes apparently
browed in anterior view). Width of frons greater than width at apex of rostrum. Rostrum. Moderately robust (width at
apex 0.69-0.85 times length in male; 0.70-0.82 in female) (Fig. 81). Median carina absent. Dorsal and lateral punctation
dense, large, deep and irregularly impressed, some punctures longitudinally confluent medially. Dorsally, excluding
epistoma, with dense, very long, erect white hair-like scales and dense very elongate-narrow recumbent white scales. In
lateral view only slightly declivous from point of antennal insertion to apex (Fig. 81a). Epistoma with apical margin
shallowly emarginate at middle (Fig. 81b). Mouthparts. Maxillary palpus with stipes lacking large seta on outer margin
(Fig. 65). Labial palpi separated by more or less twice width of base of a labial palpus (Fig. 61). Prementum with one pair
of large setae (Fig. 61). Pronotum. In dorsal view with lateral margins subparallel to slightly divergent from base to apical
one-quarter; constricted at oblique angle anterior to apical one-quarter; apical one-quarter and base subequal in width to
slightly wider at apical one-quarter; distinct lateral tubercles not evident (Fig. 24). Dorsal punctation large to very large,
dense and deep; punctures sparser smaller and shallower laterally and on flanks. Scales white, elongate-narrow,
recumbent, sparse and small or lacking medially from disk (except for midline and along apical margin), uniformly dense
laterally, sparser at lateral margins and on flanks (Fig. 24). Median carina lacking. Dorsally covered throughout with
dense very long erect white hair-like scales each situated within large puncture. Anterolateral margin with postocular
projection absent or at most very slightly developed (Fig. 81a). Prosternum. With very shallow impression anterior to each
Quaest. Ent., 1987, 23 (4)
470
Anderson
procoxal cavity and with moderately developed rounded swelling anterior to each prosternal impression (Fig. 81a). Elytra.
Moderately elongate-narrow in general form (width at midlength 0.53-0.66 times length in males; 0.54-0.64 in females)
(Fig. 24). In dorsal view with lateral margins straight to very slightly arcuate from apical one-third to humerus;
moderately and evenly arcuate from apical one-third to apex (Fig. 24). Humerus rounded, indistinct. Dorsally with dense,
very long erect hair-like scales. Sutural interval, and intervals 3 and 5 very slightly elevated and convex, especially basally.
Scales elongate-narrow, recumbent, white, uniformly moderately dense except absent to sparse small and fine on interval 4
at basal one-third and apical one-third (Fig. 24) Wings. Absent. Legs. Foretibia of female with inner margin with small
denticles in apical one-third; subapical tooth small, indistinct from tibial denticles. Foretibial, mesotibial and metatibial
unci of both sexes large. Tarsal claws not to very slightly divergent, basal internal flange distinct. Ventral tarsal pilose
vestiture of foretarsus of male and female present as small rounded apical pads of article 1, as small rounded pads on
apical one-third of article 2 and apical one-half of article 3; of mesotarsus of male present as apical tufts to small rounded
apical pads of article 1, as small rounded apical pads to small rounded pads on apical one-third of article 2 and apical
one-third to one-half of article 3; of mesotarsus of female present as apical tufts of article 1, as small rounded apical pads
of article 2, and as rounded pads on apical one-third of article 3; of metatarsus of male and female present as apical tufts of
article 1, as small rounded apical pads of article 2, and as rounded pads covering apical one-third of article 3.
Mesosternum. Mesosternal process markedly swollen and tumescent (Fig. 58). Abdomen. Ventral surface with moderately
dense, elongate-fine recumbent white scales and short suberect white hair-like scales. Abdominal sterna lacking distinct
glabrous patches. Apex of abdominal sternum VII of male lacking dorsally directed tooth. Abdominal tergum VIII
uniformly convex, apical margin slightly elevated and reflexed. Genitalia. Female (two examined). Abdominal sternum
VIII with lateral arms narrow, straight and divergent in basal one-half, moderately and evenly inwardly arcuate from
midlength to apex (Fig. 89a). Gonocoxite II with stylusmoderately large, slightly anteapical in position (Fig. 89b). Male
(two examined). Abdominal sternum VIII with paired sclerite with inner apices lacking ventral projections (Fig. 97f).
Aedeagus elongate-narrow, in lateral view thickest at midlength; in ventral view markedly laterally expanded at
approximately midlength (Figs. 97d,e). Internal sac short and high, markedly apically deflexed; median dorsal pocket
high, with moderately large dorsolaterally directed paramedial lobe at midheight and midlength and small laterally
directed lobes near basal margin at basal one-third (Figs. 97a, b). Eversible apical sclerite complex with paired scythe-like
sclerites only slightly sclerotized, each with small median projection (Fig. 97c); adjacent ventral surface and basal portion
of sides of apex of internal sac unsclerotized (Fig. 97a).
Geographic distribution. — This species has the most restricted distribution of any species of
Apleurus. It has been found only in southern California, primarily in Los Angeles County at
the El Segundo sand dunes but also at Pasadena; a single specimen is from Palm Springs,
Riverside County (Fig. 212).
Natural history. — This species appears to be restricted to sand dune habitats in the Pacific
semi-desert region of southern California. Adults have been collected on Chaenactis
glabriuscula DC., Aplopappus ericoides (Less.) H. & A. (Compositae); Croton calif ornicus
Muell.-Arg. (Euphorbiaceae); and, Lupinus albifrons Benth. (Leguminosae). Definite host
plants have yet to be determined. Adults have been collected from February to July at
elevations of from 33-313 m (N = 3) (Fig. 232).
Chorological relationships. — This species is sympatric with extreme western A. angularis
and A. albovestitus, and southern A. jacobinus.
Phylogenetic relationships. — This species is the sister-group to the rest of Apleurus (Figs.
234 and 235).
Subgenus Apleurus Chevrolat
Cleonus ; LeConte 1858 (sp. descs.). LeConte 1859 (sp. desc.). Gemminger and von Harold 1871 (in part; catal.). Leng
1920 (in part; catal.). Bradley 1930 (in part; key). Blackwelder 1947 (in part; check.). Arnett 1960-1962 (in part; key,
catal.). Tanner 1966 (in part; sp. redesc., biol.). Krombein 1979 (in part).
Apleurus Chevrolat 1873:78. Gender, masculine. Type species Apleurus fossus Chevrolat (= Cleonus lutulentus
LeConte) by subsequent designation (Casey 1891:186). Chevrolat 1873 (in part; catal., sp. desc.). Faust 1904 (key).
Centrocleonus LeConte 1876a: 145 [not Chevrolat 1873:62]. NEW SYNONYMY Type species Cleonus angularis
LeConte, according to article 67 (h) of the International Code of Zoological Nomenclature. Gender, masculine.
Henshaw 1881-1882 (check.). LeConte and Horn 1883 (key). Henshaw 1885 (check.). Wickham 1889. Horn 1894
(check.).
Cleonopsis LeConte 1876a: 147. NEW SYNONYMY Gender, feminine. Type species Cleonus pulvereus LeConte ( =
Cleonus lutulentus LeConte) by monotypy. LeConte and Horn 1883 (key). Henshaw 1885 (check.). Wickham 1889.
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471
Wickham 1896 (check.).
Cleonaspis LeConte 1876a: 153. NEW SYNONYMY Gender, feminine. Type species Cleonus lutulentus LeConte by
monotypy. Henshaw 1881-1882 (check.). LeConte and Horn 1883 (key). Henshaw 1885 (check.). Wickham 1902
(check.).
Stephanocleonus ; Henshaw 1881-1882 (in part; check.).
Dinocleus Casey 1891:176. NEW SYNONYMY. Gender, masculine. New name for Centrocleonus LeConte. Type
species Cleonus angularis LeConte by subsequent designation (Faust 1904:190, error as “Typus angulatus Lac”,
subsequently listed, p. 274, as ‘‘‘‘angularis Lee”). Wickham 1896 (check.). Fall 1901 (check.). Wickham 1902 (check.).
Champion 1902-1906 (sp. desc., notes). Casey 1904 (sp. desc.). Faust 1904 (key). Fall and Cockerell 1907 (check.).
Leng 1920 (catal.). Bradley 1930 (key). Tanner 1934 (check.). Ting 1936 (morphol.). Bruhn 1947 (morphol.). Van
Dyke 1953 (sp. desc.). Essig 1958 (biol.). Sanders 1960 (morphol.). Arnett 1960-1962 (key, catal.). Hatch 1971
(key).
Cleonus ( Apleurus ); Casey 1891 (key). Champion 1902-1906 (sp. desc., redescs.). Fall and Cockerell 1907 (check.). Csiki
1934 (catal.).
Cleonurus Faust 1904:274. Nomem nudum , incorrectly attributed to LeConte 1876a: 152.
Cleonus ( Dinocleus ); Csiki 1934 (catal.). Blackwelder 1939 (check.). Kissinger 1964 (check.).
Cleonus ( Cleonopsis)-, Kissinger 1964 (check.).
Cleonis ; O’Brien and Wibmer 1982 (in part; catal., distn.). O’Brien and Wibmer 1984 (in part).
Notes about synonymy. — The genus Apleurus was initially established by Chevrolat
(1873) to include three new Mexican and southwestern United States species, and one
previously described southwestern United States species. Three of these species, A. boucardi,
A. trivittatus and A. quadrilineatus, were regarded as “especes lyxiformes” by Chevrolat
(1873:109) thereby indicating their similarity to members of the genus Lixus and focusing
attention on their lack of similarity to A.fossus, the fourth member of the genus.
No type species was designated by Chevrolat although he (1873:80) stated in his treatment
of A. boucardi that “le corps de cet insecte est plus ovalaire que chez le type”. This suggests
that he considered as type one of the other two species ( Cleonidius trivittatus was not treated in
detail but merely placed in Apleurus by Chevrolat). Apleurus fossus was subsequently clearly
designated as the type species by Casey (1891:76).
LeConte (1876a) undertook the first comprehensive revision of North American Cleoninae,
wherein he proposed three new genera; Centrocleonus (a junior homonym of Centrocleonus
Chevrolat), Cleonopsis and Cleonaspis. Species of Centrocleonus LeConte were characterized
by the supposed unique presence of prosternal spines in front of the procoxae. Both Cleonopsis
and Cleonaspis were monobasic genera distinguished from other Cleoninae, most notably
Cleonus , primarily on the basis of features of the tarsi and antennae. I find LeConte‘s proposal
of Cleonopsis and Cleonaspis puzzling for they are based on forms which I consider to be
conspecific, differing only slightly in the extent of ventral tarsal pilosity, but otherwise not to
the extent discussed by LeConte (1876a: 144-1 45, in key) as characteristic (see also “Notes on
synonymy” of A. lutulentus). Apparently LeConte (1876a) was not concerned that these two
genera might be confused, for in his descriptions of each, only characters permitting separation
from Cleonus are discussed.
LeConte (1876a) assigned the “especes lyxiformes” of Chevrolat (1873) to Cleonus along
with a number of newly described North American species which Casey (1891) subsequently
placed in his new subgenus Cleonidius. No mention is made of A.fossus by LeConte (1876a)
and it is likely that he did not see this species for it did not occur within the geographic area of
immediate concern to him.
In the next major work on North American species, Casey (1891) recognized that the
distribution of the character states considered diagnostic of Apleurus also necessitated
inclusion of the species of Cleonopsis and Cleonaspis , and that these latter two taxa did not
warrant separate generic group status. He did not, however, consider Apleurus as warranting
any more than subgeneric status under Cleonus. Dinocleus , proposed as a new name for
Quaest. Ent., 1987, 23 (4)
472
Anderson
Centrocleonus LeConte, and considered as of generic rank, included all of LeConte‘s
Centrocleonus as well as many new species.
My use of Apleurus is decidedly broader than that of any of these authors, including not
only those species regarded as Apleurus by Casey (1891) but also all those placed in Dinocleus
Casey (1891), all subsequently described Dinocleus species and lastly, Cleonus aztecus
Champion. Support for this broader definition is seen in the reconstructed phylogeny of these
species wherein species of Dinocleus are placed as no more than derived species of Apleurus.
Recognition of Dinocleus as a distinct genus would make Apleurus paraphyletic, an
undesirable result because Apleurus are insufficiently structurally or biologically distinct from
Dinocleus to warrant recognition as a formal paraphyletic taxon. The broader definition of
Apleurus employed herein was also indirectly suggested by Van Dyke (1953) in his description
of Dinocleus bryanti. Because D. bryanti is clearly conspecific with forms then regarded by
Casey (1891), and I suspect also by Van Dyke, as Apleurus , Van Dyke’s placement of his new
species suggests a broader definition of Dinocleus and implies recognition of the similarities of
members of that genus to those previously assigned to Apleurus even though it is doubtful he
examined representatives of the latter.
Diagnosis. — Adult Apleurus with moderately robust to very robust body form (Figs.
68-80). Dorsal suberect or erect vestiture absent to very long and dense. Prementum with labial
palpi separated by distance subequal to width of basal article of labial palpus (Figs. 62-63).
Maxillary palpus with palpifer and stipes each with large seta, or with only palpifer with large
seta (Fig. 64). Rostrum with epistoma with apical margin rounded medially (Figs. 82b-88b).
Mesosternum with mesosternal process flat to only slightly convex (Fig. 59). Female with
abdominal sternum VIII with lateral arms straight or arcuate (Figs. 90a-96); gonocoxite II
with stylus absent or present, small to moderately large, apical to anteapical in position (Figs.
90b-96b). Male with abdominal sternum VIII with each sclerite with basal projection at
interior angle slightly to markedly developed (Figs. lOle, 103e). Aedeagus in dorsal view more
or less uniform in width throughout median portion of length.
Included species. — Seven species are placed in the subgenus Apleurus. They are found in
the southwestern United States of America south to southern Mexico.
Phylogenetic relationships. — This subgenus is the monophyletic sister-group of the
subgenus Gibbostethus (see “Phylogeny” section).
Key to species of adult Apleurus (Apleurus)
1 Elytra with sutural interval, intervals 3, 5, 7, and 9 markedly elevated and
convex throughout greater part of their lengths. Metathoracic wings absent 2
V Elytra with all intervals flat or with sutural interval, intervals 3, 5, and 7
only very slightly elevated and convex. Metathoracic wings present,
variable in length from short (approximately one-half length of elytra) to
long (equal to or greater than elytra in length) 6
2 (1) Prosternum lacking swelling or with at most low rounded swelling anterior
to each prosternal impression (Fig. 82a). Rostrum lacking median carina.
Elytra very robust (width at midlength greater than 0.70 times length)
(Fig. 68) A. aztecus (Champion), p. 474
2' Prosternum with prominent swelling anterior to each prosternal impression
(Figs. 84a-88a). Rostrum with median carina (obscured in some specimens
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473
by large deep punctures) (Figs. 84-86). Elytra less robust (width at
midlength less than 0.75 times length) (Figs. 70-73) 3
3 (2') Tarsal claws slender, widely divergent, each lacking basal internal flange,
tapered evenly from base to apex (Fig. 66). Dorsal erect vestiture of
pronotum and elytra short 4
3' Tarsal claws robust, not widely divergent, each with basal internal flange,
tapered abruptly from midlength to apex (Fig. 67). Dorsal erect vestiture
of pronotum and elytra short or long 5
4 (3) Abdomen ventrally with scales broad, moderately dense; at least sterna III
and IV with distinct glabr us shiny patches, each with single long and erect
hair A. angularis (LeConte) (in part), p. 481
4' Abdomen ventrally with scales fine and elongate, extremely dense; no
glabrous patches present A. porosus (LeConte) (in part), p. 478
5 (30 Elytra with sutural interval, intervals 3, 5, 7, and 9 (especially 3 and 5 near
base) markedly elevated and convex. Most specimens with dorsal vestiture
of head, pronotum and elytra very long and erect and with rostrum with
carina moderately well-developed (Fig. 86). Abdomen ventrally with scales
fine and elongate, moderately dense; sterna III and IV with distinct
glabrous shiny patches, each with single long and erect hair
A. jacobinus (Casey), p. 484
5' Elytra with sutural interval, intervals 3, 5, 7, and 9 less markedly elevated
and convex. Dorsal vestiture of head, pronotum and elytra short and
suberect. Rostrum with carina only slightly developed, almost entirely
obscured in some specimens by large deep punctures (Fig. 84). Abdomen
ventrally with scales very fine and elongate, extremely dense; no glabrous
patches present A. porosus (LeConte) (in part), p. 478
6 (ly) Pronotum laterally expanded immediately posterior to subapical
constriction (giving distinctly tuberculate appearance); with base and
apical one-quarter approximately equal in width to distinctly widest at
apical one-quarter (Figs. 72-73). Tarsal claws slender, widely divergent,
each lacking basal internal flange and tapered evenly from base to apex
(Fig. 66)
A. angularis (LeConte) (in part), p. 481
6' Pronotum not laterally expanded immediately posterior to subapical
constriction (not appearing distinctly tuberculate); with base and apical
one-quarter approximately equal in width to distinctly widest at base (Figs.
69, 74-80). Tarsal claws robust, not widely divergent; each with basal
internal flange and tapered abruptly from midlength to apex (as in Fig. 67) 7
7 (60 Tarsus of hind leg with venter of article 3 with large pubescent pad.
Prosternum lacking swelling or with swelling anterior to each prosternal
impression only slightly developed (Fig. 83a). Female with tergum VII not
longitudinally carinate; gonocoxite II with stylus large (Fig. 91b)
A. lutulentus (LeConte), p. 475
T Tarsus of hind leg with venter of article 3 lacking large pubescent pad, with
at most small apical tuft of pubescence. Prosternum with prominent
swelling anterior to each prosternal impression (Figs. 87-88). Female with
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474
Anderson
tergum VII longitudinally carinate; gonocoxite II lacking stylus (Figs.
95-96) 8
8 (7') Tarsus of fore- and middle leg with venter of article 3 with large pubescent
pad A. saginatus (Casey) (in part; males), p. 491
8' Tarsus of fore- and middle leg with venter of article 3 lacking large
pubescent pad, with at most small apical tuft of pubescence 9
9 (8') Pronotum with lateral margins with outer margins of large punctures
swollen, shiny and glabrous (appearing as small distinct tubercles).
Abdomen ventrally (especially sterna III and IV) with distinct large
glabrous patches, each with single short and suberect scale situated in the
center A. saginatus (Casey) (in part; females), p. 491
9 ' Pronotum with lateral margins with outer margins of large punctures not
swollen, glabrous or shiny. Abdomen ventrally either lacking glabrous
patches and with scales extremely dense; with small glabrous patches; or,
with most scales abraded A. albovestitus (Casey), p. 486
Apleurus (Apleurus) aztecus (Champion), new combination
(Figs. 68,82, 90, 98,210)
Cleonus aztecus Champion 1902-1906:99. Lectotype (here designated), male, one of two syntypes, labelled “Type”,
inverted “Sp. figured”, “Refugio/ Durango/ Hoge.”, “B.C.A. Col. IV. 4./ Cleonus/ aztecus,/ Champ.” and with my
designation label “LECTOTYPE/ Cleonus aztecus/ Champ, desig./ Anderson” (BMNH). Type locality, Refugio,
Durango, Mexico.
Cleonis aztecus\ O’Brien and Wibmer 1982 (catal., distn.).
Problems in recognition. — Members of this species are most likely to be confused with
brachypterous specimens of A. lutulentus. In these latter specimens, the alternate elytral
intervals are slightly elevated and convex and the elytra are moderately robust (more so than in
macropterous A. lutulentus). These individuals are sympatric with A. aztecus and although
superficially similar, A. aztecus individuals are distinguished by the tarsal claws each lacking a
basal internal flange (as in Fig. 66), lack of metathoracic wings, and lack of a dorsally directed
tooth on the apical margin of abdominal sternum VII in males.
A. aztecus individuals may also be confused with those other species of Apleurus in which
alternate elytral intervals are elevated and convex. Key characters should readily serve to
separate these species.
Description. — Specimens examined. 16 males, 7 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 8.
Size. Length, male, 5. 6-9. 4 mm; female, 5. 2-9. 4 mm. Width, male, 2. 9-4. 6 mm; female, 2. 5-4. 8 mm. Head. Eye
prominent and convex in dorsal view. Frons and vertex with moderately dense, small to moderately large, deep punctures.
Frons also with sparse, short suberect white hair-like scales immediately above eyes. Area immediately behind posterior
margin of eye with large, deep, irregularly impressed punctures. Area above eyes slightly elevated above rest of frons (eyes
apparently browed in anterior view). Width of frons greater than width at apex of rostrum. Rostrum. Robust (width at
apex 0.79-1.00 times length in male; 0.78-0.91 in female) (Fig. 82). Median carina absent. Dorsal and lateral punctation
moderately dense, small to large, deep. Dorsally, excluding epistoma, with moderately dense, short, suberect white
hair-like scales and moderately dense elongate-narrow appressed white scales. In lateral view with apical portion flat to
only very slightly declivous from point of antennal insertion to apex (Fig. 82a). Epistoma with apical margin rounded at
middle (Fig. 82b). Mouthparts. Maxillary palpus with stipes lacking large seta on outer margin (as in Fig. 65). Labial
palpi separated by more or less width of basal article of labial palpus (as in Fig. 62-63). Prementum with one pair of large
setae (as in Fig. 62). Pronotum. In dorsal view with lateral margins subparallel to slightly divergent from base to apical
one-quarter; constricted at oblique angle anterior to apical one-quarter; apical one-quarter and base subequal in width to
slightly wider at apical one-quarter; without distinct lateral tubercles (Fig. 68). Dorsal and lateral punctation small, sparse
to moderately dense and shallow; punctures sparser and smaller on flanks. Scales white, elongate-narrow, appressed, sparse
and small or lacking medially from disk and dorsally from flanks (except for midline and area anterior to low transverse
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475
carina if present), dense laterally to lateral margins in anterior two-thirds and ventrally on flanks. Median carina lacking;
some specimens with low median transverse carina or impunctate area at midlength. Dorsally with dense very short fine
suberect white hair-like scales each situated within large puncture. Anterolateral margin with postocular projection absent
or at most very slightly developed (Fig. 82a). Prosternum. With very shallow impression anterior to each procoxal cavity
and with at most a low slightly developed rounded swelling anterior to each prosternal impression (Fig. 82a). Elytra. Very
robust in general form (width at midlength 0.70-0.90 times length in males; 0.72-0.80 in females) (Fig. 68). In dorsal view
with lateral margins moderately arcuate throughout length (Fig. 68). Humerus rounded, indistinct. Dorsally with sutural
interval and intervals 3, 5, 7, and 9 with multiple rows of short fine suberect hair-like scales; intervals 2, 4, 6, and 8 with
only single row of similar hair-like scales. Sutural interval, and intervals 3, 5, 7, and 9 markedly elevated and convex
throughout their lengths. Scales various in density and size; elongate-narrow, pattern various but with at least small
glabrous (or nearly so) area immediately apicad of midlength on interval 4 (Fig. 68). Wings. Absent. Legs. Foretibia of
female with inner margin with small denticles in apical one-third; subapical tooth large, distinct from tibial denticles.
Foretibial and mesotibial unci of both sexes large, metatibial uncus of both sexes small. Tarsal claws widely divergent, no
basal internal flange present (as in Fig. 66). Ventral tarsal pilose vestiture of foretarsus of male present as small rounded
apical pads of article 1, as elongate-oval pads on apical one-half of article 2, and as large elongate-oval pads covering
apical two-thirds of article 3; of foretarsus of female present as small rounded apical pads on articles 1 and 2, as rounded
pads on apical one-third to one-half of article 3; of mesotarsus of male present as minute apical tufts on articles 1 and 2,
and as small rounded apical pads to small rounded pads on apical one-half of article 3; of mesotarsus of female absent to
present as minute apical tufts on articles 1 and 2, and as small rounded apical pads of article 3; of metatarsus of male
present as minute apical tufts on articles 1 and 2, and as minute to small apical tufts of article 3; of metatarsus of female
absent on articles 1 and 2, and present as minute apical tuft of article 3. Mesosternum. Mesosternal process moderately
convex but not tumescent (as in Fig. 59). Abdomen. Ventral surface with moderately dense, elongate-fine appressed white
scales. Abdominal sterna III to VI (especially III and IV) with moderately dense, small, individually indistinct to distinct
rounded glabrous shiny patches, each with large puncture and single long erect hair-like scale situated in or near center.
Apex of abdominal sternum VII of male lacking dorsally directed tooth. Abdominal tergum VIII uniformly convex, apical
margin slightly elevated and reflexed. Genitalia. Female (two examined). Abdominal sternum VIII with lateral arms
narrow, very slightly sinuate and markedly inwardly arcuate near apex (Fig. 90a). Gonocoxite II with stylus moderately
large, slightly anteapical in position (Fig. 90b). Male (three examined). Abdominal sternum VIII with paired sclerite with
inner apices lacking or with only slightly developed ventral projections. Aedeagus elongate-narrow, in lateral view thickest
at midlength; in ventral view more or less parallel sided throughout length. Internal sac elongate and low, only slightly
apically deflexed; median dorsal pocket low, with moderately large dorsolaterally directed paramedial lobe at midheight at
basal one-third, small median lobe on dorsal surface immediately anterior to crest of median dorsal pocket, and moderately
large dorsolaterally directed paramedial lobes at midheight at apical one-third (Figs. 98a, b). Eversible apical sclerite
complex with paired narrow scythe-like sclerites well sclerotized, each lacking median projection (Fig. 98d); adjacent
ventral surface and basal portion of sides of apex of internal sac with pair of well-developed transverse sclerites (Fig. 98a).
Geographic distribution. — This species is found throughout the highlands of central
Mexico (Fig. 210).
Natural history. — This species appears to be restricted to the xeric high-elevation
mesquite-grasslands of central Mexico (pastizal and perhaps pastizal-matorral xerofilo
transition of Rzedowski [1978]). Adults have been collected on Solidago sp. (Compositae).
Other adults have been collected from under prickly-pear pads, stones and dry cow dung.
Definite host plants are unknown. Adults have been collected from June to August at elevations
from 2250-3385 m (N = 8) (Fig. 232).
Chorological relationships. — This species is sympatric throughout its range with A.
lutulentus. Adult specimens of the two species have been collected together at 1.5 mi. S.
Fresnillo (Zacatecas) and 20 mi. NE. Dolores Flidalgo (Guanajuato).
Phylogenetic relationships. — This species is the sister-species of the rest of the species in
the subgenus Apleurus (Figs. 234-235).
Apleurus ( Apleurus ) lutulentus (LeConte), new combination
(Figs. 6, 59, 62, 69, 83, 91, 99, 207)
Cleonus lutulentus LeConte 1859a: 18. Lectotype (here designated), female, one of two syntypes (only one examined),
labelled with a dark green circle ( = New Mexico), “Type/ 5186”, “ Cleonaspis / lutulentus/ (Lee)” and with my
designation label “ Cleonus j lutulentus/ LeC. LECTOTYPE/ desig. Anderson” (MCZC). Type locality, Santa Fe,
New Mexico. Gemminger and von Harold 1871 (catal.). Leng 1920 (catal.).
Quaest. Ent., 1987, 23 (4)
476
Anderson
Cleonus pulvereus LeConte 1 859a: 1 8. NEW SYNONYMY Holotype (examined), female, labelled “Dallas/ Tex/ Boll”,
“Type/ 5185”, “ Cleonopsis / pulvereus/ (Lee)” (MCZC). Type locality, Dallas, Texas. Gemminger and von Harold
1871 (catal.). Leng 1920 (catal., error as Cleonus pluvereus). Blackwelder 1947 (check.). Krombein 1979 (as prey).
Apleurus fossus Chevrolat 1873:78. NEW SYNONYMY Lectotype (here designated), female, one of two syntypes,
labelled “12....”, “Mexico/ Au. Salle”, “TYPUS”, “40”, “473/ 85”, “Riksmuseum/ Stockholm” and with my
designation label “ Apleurus fossus/ Chevrolat/ LECTOTYPE/ desig. Anderson” (Riksmuseum, Stockholm; other
syntype in BMNH). Type locality, Mexico. Faust 1904.
Cleonaspis lutulentus; LeConte 1876a (redesc.). Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1902
(check.).
Cleonopsis pulvereus-, LeConte 1876a (redesc.). Henshaw 1885 (check.). Wickham 1889. Wickham 1896 (check.).
Stephanocleonus pulvereus-, Henshaw 1881-1882 (check.).
Cleonus ( Apleurus ) fossus ; Casey 1891 (designation as type species of Apleurus Chevrolat). Champion 1902-1906 (syn.,
distn.). Csiki 1934 (catal.).
Cleonus ( Apleurus ) lutulentus\ Casey 1891 (key). Fall and Cockerell 1907 (check.). Csiki 1934 (catal.).
Cleonus ( Apleurus ) pulvereus-, Casey 1891 (key). Champion 1902-1906 (syn., distn.). Fall and Cockerell 1907 (check.).
Csiki 1934 (catal.).
Cleonus fossus-, Blackwelder 1947 (check.).
Dinocleus bryanti Van Dyke 1953:101. NEW SYNONYMY Holotype (examined), male, labelled “Seligman,/ Ariz. VIII
3 36/ Bryant. 1 12.”, “From the/ O. Bryant/ Collection”, “Holotype” (CASC). Type locality, Seligman, Arizona.
Cleonis bryanti-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis fossus; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis lutulentus; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis pulvereus; O’Brien and Wibmer 1982 (catal., distn.).
lutatentus, incertae sedis; Chevrolat 1873 (error, misspelling).
pulverosus, incertae sedis; Chevrolat 1873 (error, misspelling).
Notes about synonymy. — I can find no consistent differences that warrant separation of
Cleonus pulvereus LeConte, Cleonus lutulentus LeConte and Dinocleus bryanti Van Dyke as
distinct species. LeConte (1859a) emphasized characters (extent of ventral tarsal vestiture and
degree of development of rostral carina) which are intraspecifically variable in describing C.
pulvereus and C. lutulentus and subsequently placing them in separate genera (LeConte,
1876a) (see “Notes about synonymy” section for Apleurus). I have not seen any specimens of
A. lutulentus from which the ventral tarsal pilose vestiture is entirely lacking and in which the
tarsal articles are not bilobed, as was given by LeConte (1876a) as in part characteristic of C.
lutulentus , and I do not find that the characters subsequently given by Casey (1891) to
separate the two species represent anything more than intraspecific variation.
Dinocleus bryanti Van Dyke is clearly A. lutulentus. Van Dyke (1953) considered D.
bryanti a member of the genus Dinocleus, however, since he did not mention either Cleonus
pulvereus or Cleonus lutulentus, and since neither was then considered to be Dinocleus, I
suspect that Van Dyke did not look at representatives of these forms in describing D. bryanti,
but only at those forms already placed in Dinocleus. As a result, he found his new species to be
very distinct from those then included in Dinocleus. As discussed elsewhere (“Notes about
synonymy” section for Apleurus) this placement of D. bryanti in Dinocleus concurs with my
classification of species of Dinocleus and Apleurus (including Cleonaspis and Cleonopsis)
herein as congeneric.
I also consider the Mexican brachypterous A. fossus as conspecific with the other forms
discussed previously in this section. No specific type locality was given for this species but
because brachypterous individuals are otherwise only known from central and southern Mexico
I suspect this is the area of collection of the type series. These brachypterous forms are
otherwise distinguished from typical C. lutulentus by a more robust and globose elytral form,
possession of elytral intervals 3, 5 and 7 slightly elevated and convex, and possession of deep
and large punctures on the head, pronotum and elytra. Although only this brachypterous form
occurs in the southern Mexican highlands of Oaxaca and Puebla, some of these similarly
brachypterous individuals are also known to be sympatric with typically macropterous
New World Cleonini
All
individuals in central Mexico. Because wing length polymorphism and associated variation in
elytral form is found in sympatric individuals of other species of Apleurus (A. angularis and A.
albovestitus), I concur with Champion (1902-1906:98) that A. fossus is “probably nothing
more than a southern form of C. pulvereus LeC.”.
Problems in recognition. — Brachypterous specimens of this species may be confused with
individuals of A. aztecus as noted in the “Problems in recognition” section for that species.
They may also be confused with small A. saginatus or A. albovestitus, or macropterous A.
angularis, but the presence of distinct prosternal swellings anterior to the prosternal
impressions in members of the latter three species (Figs. 85, 87-88), among other key
characters, will readily separate them from A. lutulentus.
Description. — Specimens examined. 334 males, 369 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 9.
Size. Length, male, 5.3-10.3 mm; female, 6.4-11.4 mm. Width, male, 2.4-4. 8 mm; female, 2. 9-5. 5 mm. Head. Eye
prominent and convex in dorsal view. Frons and vertex with moderately dense, small to large, deep punctures. Frons
lacking suberect or erect vestiture; with only sparse elongate-narrow appressed white scales. Area immediately behind
posterior margin of eye with large, deep, irregularly impressed punctures. Area above eyes flat, not distinct from rest of
frons (eyes not appearing distinctly browed in anterior view). Width of frons greater than to subequal to width at apex of
rostrum. Rostrum. Moderately robust (width at apex 0.65-0.91 times length in male; 0.67-0.84 in female) (Fig. 83).
Median carina absent or at most only slightly developed and indistinct. Dorsal and lateral punctation moderately dense,
small to large, deep; sparser apically and irregularly impressed medially in some specimens). Dorsally, excluding epistoma,
with scattered very short, fine, suberect hair-like scales towards apex, and sparse elongate-narrow appressed white scales.
In lateral view with apical portion flat to only very slightly declivous from point of antennal in ertion to apex (Fig. 83a).
Epistoma with apical margin rounded at middle (Fig. 83b). Mouthparts. Maxillary palpus with stipes with large seta on
outer margin (as in Fig. 64). Labial palpi separated by more or less width of basal article of a labial palpus (Fig. 62).
Prementum with one pair of large setae (Fig. 62). Pronotum. In dorsal view with lateral margins slightly arcuate from
base to apical one-quarter; constricted at oblique angle anterior to apical one-quarter; pronotum widest at base to subequal
in width at base and at apical one-quarter; distinct lateral tubercles not evident (Fig. 69). Dorsal and lateral punctation
small to large, moderately dense to dense, and deep; punctures sparser and smaller on flanks. Scales white,
elongate-narrow, appressed, sparse and small or lacking medially from disk in a broad, apically narrowed patch (except for
midline in some specimens), and at lateral margins near base; uniformly dense laterally to lateral margins in anterior
one-half and ventrally on flanks. Median carina lacking. Dorsally lacking suberect or erect vestiture. Anterolateral margin
with postocular projection absent or at most very slightly developed (Fig. 83a). Prosternum. With very shallow impression
anterior to each procoxal cavity and with at most low slightly developed rounded swelling anterior to each prosternal
impression (Fig. 83a). Elytra. Moderately robust in general form (width at midlength 0.57-0.72 times length in males;
0. 52-0.76 in females) (Fig. 69). In dorsal view with lateral margins slightly arcuate and convergent to subparallel from
apical one-third to humerus; markedly arcuate from apical one-third to apex; humerus obtuse to acute, moderately distinct
to distinct (Fig. 69). Dorsal suberect or erect vestiture absent or at most with sutural interval and intervals 2 and 3 with
very short suberect hair-like scales. Dorsally with all elytral intervals flat to slightly convex, to with sutural interval and
intervals 3, 5, and 7 slightly swollen and convex in brachypterous individuals. Scales various in density and size;
elongate-narrow, pattern various with numerous irregularly distributed small glabrous or nearly so areas (Fig. 69). Wings.
Short (very slightly shorter than elytra in length [2.1%, N= 15]) to long (greater than elytra in length [97.9%, N = 688]).
Branches of 2A not joined at base. Legs. Foretibia of female with inner margin with small to moderately large denticles
throughout most of length; subapical tooth moderately large, distinct from tibial denticles. Foretibial and mesotibial unci
of both sexes large, metatibial uncus of both sexes moderately large. Tarsal claws not divergent, with well-developed basal
internal flange present. Ventral tarsal pilose vestiture of foretarsus of male present as small elongate apical pads of article
1, as elongate-oval pads on apical one-half of article 2, and as large elongate-oval pads on apical two-thirds of article 3; of
foretarsus of female present as small elongate apical pads of article 1, as moderately large elongate-oval pads on apical
one-half of article 2, and as large elongate-oval pads on apical one-half to two-thirds of article 3; of mesotarsus of male
present as elongate-narrow pads on apical one-third of article 1 and apical one-half of article 2, and as large elongate-oval
pads on apical one-half to two-thirds of article 3; of mesotarsus of female present as elongate-narrow pads on apical
one-third of article 1, as elongate-oval pads on apical one-half of article 2, and as large rounded pads on apical one-half of
article 3; of metatarsus of male absent of article 1, present as elongate very narrow pads of article 2, and as elongate-oval
pads on apical one-half of article 3; of metatarsus of female absent to as small elongate-narrow pads on apical one-quarter
of article 1 , as elongate very narrow pads on apical one-third of article 2, and as large rounded pads on apical one-half to
two-thirds of article 3. Mesosternum. Mesosternal process flat to very slightly convex but not tumescent (Fig 59).
Abdomen. Ventral surface with dense, elongate-fine appressed white scales. Abdominal sterna III to VI (especially III and
IV) with sparse to moderately dense, small, individually indistinct to distinct rounded glabrous shiny patches, each with
large puncture and single short erect hair-like scale situated in or near center. Apex of abdominal sternum VII of male
Quaest. Ent., 1987, 23 (4)
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Anderson
with large dorsally directed median truncate tooth. Abdominal tergum VIII uniformly convex, apical margin slightly
elevated and reflexed. Genitalia. Female (three examined). Abdominal sternum VIII with lateral arms narrow, very
slightly inwardly arcuate in basal three-quarters, then markedly inwardly arcuate to apex (Fig. 91a). Gonocoxite II with
stylus moderately large, anteapical in position by more or less length of stylus (Fig. 91b). Male (12 examined). Abdominal
sternum VIII with paired sclerite with inner apices with slightly to moderately developed ventral projections. Aedeagus
elongate-narrow, in lateral view subequal in thickness from base to midlength (Fig. 99c); in ventral view more or less
parallel sided throughout length. Internal sac elongate and low, only slightly apically deflexed; median dorsal pocket low,
with moderately large dorsolaterally directed paramedial lobe at midheight at basal one-third and small median lobe on
dorsal surface immediately anterior to crest of median dorsal pocket (Figs. 99a, b). Eversible apical sclerite complex with
paired narrow scythe-like sclerites well sclerotized, each with short median projection (Fig. 99d); adjacent ventral surface
and basal portion of sides of apex of internal sac with pair of well-developed transverse sclerites (Fig. 99a).
Geographic variation. — Some specimens from central Mexico have larger and deeper
punctures of the head, pronotum and elytra; more robust and globose elytra; elytral intervals 3,
5, and 7 slightly elevated and convex; and short wings. Only specimens with extreme states in
all of these characters are found in the area of the southern border of the species range in the
Mexican states of Oaxaca and Puebla.
Geographic distribution. — This species is found from Kansas, Oklahoma and eastern
Texas, west to Arizona, disjunct to central and southern Mexico (Fig. 207). The species does
not appear to be found in the Chihuahuan desert region in Mexico. There is a single
questionable record from northeastern California.
Natural history.— This is the most widely distributed species of Apleurus. It is found in
mesquite-grassland, grassland, and desert-grassland transitional habitats in Mexico and the
United States, and in the grassland-deciduous forest transitional zone. Adults have been
collected in dry upland desert washes and grasslands on a wide variety of plants, but mostly
Compositae, as follows: Gossypium hirsutum L. (cotton, Malvaceae); Chenopodium sp.
(Chenopodiaceae); Asclepias sp., A. subverticillata (Gray) Vail (Asclepiadaceae); Koeberlinia
spinosa Zucc. (Capparaceae); Acacia sp. (Leguminosae); Baccharis sp., B. glutinosa Pers.,
Baileya pleniradiata Harv. and Gray, Chrysothamnus nauseosus (Pall.) Britton,, Flourensia
cernua DC., Gutierrezia sp., Solidago sp., Verbesina enceloides (Cav.) Benth. and Hook., V.
oreophila Woot. and Standi., Xanthium sp. (all Compositae). Definite host plants are not
known. Adults have been collected from March to October at elevations from 9-3058 m
(N = 77) (Fig. 232).
Chorological relationships. — This species is sympatric with A. saginatus in southern
Arizona and New Mexico; with A. aztecus in central Mexico; with A. angularis in New
Mexico, western Texas, Colorado, western Oklahoma, and Arizona; and with A. albovestitus in
Arizona, New Mexico, and western Texas.
Adults of A. lutulentus and A. albovestitus have been caught together near Portal, Arizona,
on Chrysothamnus nauseosus. Adult specimens of A. lutulentus and A. aztecus have been
collected together 1.5 mi. S. Fresnillo (Zacatecas), on Solidago sp., and 20 mi. NE. Dolores
Hidalgo (Guanajuato).
Phylogenetic relationships. — This species is the sister-species of the A. porosus-A.
angularis-A. jacobinus-A. saginatus-A. albovestitus lineage of Apleurus (Figs. 234-235).
Apleurus ( Apleurus ) porosus (LeConte), new combination
(Figs. 70, 84, 92, 102,210)
Centrocleonus porosus LeConte 1876a: 1 46. Holotype (examined), female, labelled with a silver circle with portion cut
away ( = Baja California) “Type/ 5239”, “C. porosus/ Lee” (MCZC). Type locality, Cape San Lucas, Baja
California Sur, Mexico. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Horn 1894 (check.).
Dinocleus farctus Casey 1891:181. NEW SYNONYMY Holotype (examined), male, labelled “Cal”, “CASEY/ bequest/
New World Cleonini
479
1925”, “TYPE USNM/ 37271”, “D./ farctus/ Cas.” (USNM). Type locality, California, near the southern border.
Fall 1901 (check.). Leng 1920 (catal.).
Dinocleus porosus-, Casey 1891 (key, redesc.). Leng 1920 (catal.).
Cleonus ( Dinocleus ) farctus ; Csiki 1934 (catal.).
Cleonus ( Dinocleus ) porosus-, Csiki 1934 (catal.).
Cleonus porosus-, Blackwelder 1947 (check.).
Cleonus farctus ; Hardy and Andrews 1976 (distn., prob. misident. A. angularis).
Cleonis farctus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis porosus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — Both LeConte (1876a) and Casey (1891) based their species
descriptions of Centrocleonus porosus and Dinocleus farctus on unique specimens. Having
examined larger numbers of individuals from various localities, I conclude that the two are
conspecific and the differences noted by Casey (1891) as species^specific are due to
intraspecific variation or secondary sexual characteristics (the holotype of C. porosus is a
female; that of D. farctus, a male).
The type locality of D. farctus is given by Casey (1891) as California, near the southern
border. I have not seen any other A. porosus from the state of California.
Problems in recognition. — Members of this species are likely to be confused with A.
angularis but can be distinguished from individuals of that species by the robust form of the
tarsal claws, each possessing a basal internal flange in most specimens (as in Fig. 67); by the
lack of distinct glabrous patches on the abdominal sterna; and by presence of only very short
suberect dorsal vestiture in the former. Some specimens of A. porosus from various localities
however, possess tarsal claws, which although moderately robust, lack a distinct internal flange.
Apleurus porosus specimens are also likely to be confused with A. jacobinus individuals
particularly in Baja California Norte (and possibly also in southern California) where the
species appear to be narrowly sympatric. Apleurus jacobinus specimens are distinguished from
those of A. porosus by the long dorsal erect vestiture; alternate elytral intervals markedly
elevated and convex; and abdominal sterna with distinct glabrous patches in the former.
However, in the southern part of the range of A. jacobinus in Baja California Norte,
individuals I examined have shorter dorsal erect vestiture and less markedly elevated elytral
intervals than elsewhere throughout the species range, thus, approaching the states in A.
porosus in these characters. Although the differences in degree of elevation of the elytral
intervals and length of dorsal vestiture in these individuals should still allow for separation of A.
jacobinus and A. porosus , the presence of distinct glabrous patches on the abdominal sterna of
the former will allow for unequivocal separation of the two species. Apleurus porosus
specimens also have the pronotum with scales uniformly very dense laterally and on the flanks,
more or less obscuring the individual large punctures.
Many A. porosus can also be separated from A. jacobinus and A. angularis by their larger
size, more elongate-narrow rostrum, and by the pronotum being widest at the base (see Figs.
105-106, 109).
Description. — Specimens examined. 23 males, 32 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 10.
Size. Length, male, 5.8-13.6 mm; female, 6.9-15.2 mm. Width, male, 2.6-6. 1 mm; female, 3. 0-7. 2 mm. Head. Eye very
prominent and convex in dorsal view. Frons and vertex with dense, large to very large, deep punctures. Frons and area
immediately above eyes with sparse to moderately dense short suberect hair-like scales. Area immediately behind posterior
margin of eye with large, deep, irregularly impressed punctures. Area above eyes slightly elevated above rest of frons (eyes
appearing distinctly browed in anterior view). Width of frons greater than width at apex of rostrum. Rostrum. Moderately
elongate-narrow (width at apex 0.61-0.68 times length in male; 0.62-0.69 in female) (Fig. 84). Median carina absent to
slightly developed but distinct, almost entirely obscured in some specimens by large deep punctures. Dorsal and lateral
punctation moderately dense, large, deep. Dorsally, excluding epistoma, with scattered short, fine, suberect hair-like scales
towards apex, and dense elongate-narrow appressed white scales. In lateral view with apical portion steeply declivous from
Quaest. Ent., 1987, 23 (4)
480
Anderson
point of antennal insertion to apex (Fig. 84a). Epistoma with apical margin rounded at middle (Fig. 84b). Mouthparts.
Maxillary palpus with stipes lacking large seta on outer margin (as in Fig. 65). Labial palpi separated by more or less
width of basal article of a labial palpus (as in Figs. 62-63). Prementum with two or three pairs of large setae (as in Fig.
63). Pronotum. In dorsal view with lateral margins subparallel to slightly convergent from base to apical one-quarter;
constricted at acute angle anterior to apical one-quarter; widest at base to subequal in width at base and apical
one-quarter; distinct lateral tubercles evident (Fig. 70). Dorsal and lateral punctation large, moderately deep to deep,
dense; punctures sparser and smaller on flanks. Scales white, elongate-narrow, appressed, sparse and small or lacking
medially from disk in broad, apically narrowed patch (except for midline in some specimens); uniformly dense laterally, at
lateral margins and ventrally on flanks, more or less obscuring large punctures. Median carina lacking. Dorsally covered
throughout with sparse, fine, short, erect hair-like scales. Anterolateral margin with postocular projection absent or at
most very slightly developed (Fig. 84a). Prosternum. With shallow impression anterior to each procoxal cavity and with
well-developed rounded swelling anterior to each prosternal impression (Fig. 84a). Elytra. Moderately robust in general
form (width at midlength 0.65-0.70 times length in males; 0.65-0.73 in females) (Fig. 70). In dorsal view with lateral
margins moderately arcuate throughout length; humerus rounded, not distinct (Fig. 70). Sutural interval and intervals 3,
5, 7, and 9 with sparse, very short, suberect hair-like scales. Dorsally with sutural interval and intervals 3, 5, 7, and 9
slightly swollen and convex (more so basally in some specimens). Scales elongate-narrow, various in density and size;
pattern various but in most specimens with two extensive glabrous (or nearly so) areas, one from area behind humerus,
posteromedially directed to sutural interval at midlength, other continous with the former at midlength and directed
laterally from sutural interval to interval 9; some specimens also with glabrous (or nearly so) area of variable extent at
confluence of intervals 4 to 7 (Fig. 70). Wings. Absent. Legs. Foretibia of female with inner margin with small denticles in
apical one-third; subapical tooth small, not distinct from tibial denticles. Foretibial and mesotibial unci of both sexes large,
metatibial uncus of both sexes moderately large. Tarsal claws not divergent, basal internal flange absent to present,
well-developed. Ventral tarsal pilose vestiture of foretarsus of male absent of article 1, present as minute apical tufts of
article 2, as moderately large rounded pads on apical one-third to one-half of article 3; of foretarsus of female, mesotarsus
of male and female, and metatarsus of male, lacking from articles 1 and 2, present as small rounded pads at apex of article
3; of metatarsus of female lacking from articles 1 and 2, present as minute pads at apex of article 3. Mesosternum.
Mesosternal process flat to very slightly convex but not tumescent (as in Fig. 59). Abdomen. Ventral surface with very
dense, elongate-fine appressed white scales and scattered suberect hair-like scales. Abdominal sterna lacking distinct
glabrous patches, with at most only lateral margins of abdominal sterna III and IV with very small indistinct glabrous
patches. Apex of abdominal sternum VII of male with small dorsally directed median truncate tooth. Abdominal tergum
VIII uniformly convex, apical margin slightly elevated and reflexed. Genitalia. Female (four examined). Abdominal
sternum VIII with lateral arms narrow, slightly outwardly arcuate in basal one-half, then parallel to near apex, markedly
inwardly arcuate immediately before apex (Fig. 92a). Gonocoxite II with stylus very small, anteapical in position by more
or less one-half length of stylus (Fig. 92b). Male (five examined). Abdominal sternum VIII with paired sclerite with inner
apices with slightly to moderately developed ventral projections (as in Fig. 10 le). Aedeagus elongate-narrow, in lateral
view thickest at midlength; in ventral view more or less parallel sided throughout length. Internal sac short and high,
markedly apically deflexed; median dorsal pocket high, with moderately large dorsolaterally directed paramedial lobe at
midheight and midlength; small median lobe on dorsal surface immediately anterior to crest of median dorsal pocket (Figs.
102a,b). Eversible apical sclerite complex with paired narrow scythe-like sclerites lightly sclerotized, each lacking median
projection (as in Figs. lOOd-lOld); adjacent ventral surface and basal portion of sides of apex of internal sac with pair of
indistinct lightly sclerotized transverse sclerites (Fig. 102a).
Geographic variation. — None noted, but this may be due to the small number of specimens
examined, especially from Baja California Norte where this species is at most narrowly
sympatric with A. angularis and A. jacobinus.
Geographic distribution. — This species is found throughout Baja California Sud and Baja
California Norte, Mexico (Fig. 210). It may occur in the extreme south of California.
Natural history. — Label data indicate that most adults have been caught in sand dune
areas in the “matorral xerofilo” of Baja California (Sonoran desert) and Pacific semi-desert
region at elevations of from 6-144 m (N = 5) (Fig. 232). Sleeper (pers. comm.) has collected
numerous specimens from a woody Encelia species and a single specimen from Encelia
laciniata Vasey and Rose (Compositae). Definite host plants are not known.
Chorological relationships — This species has one of the more restricted distributions of
Apleurus species. It is sympatric with extreme southern A. albovestitus, and at most narrowly
sympatric to parapatric with its close relatives A. angularis and A. jacobinus.
Phylogenetic relationships. — Apleurus porosus is the sister-species of the A. angularis-A.
jacobinus sister-species pair (Fig. 235). Presence of the apotypic state of tarsal claws each
lacking a basal internal flange in some A. porosus and in A. angularis (Fig. 66) is either a
New World Cleonini
481
result of hybridization or of homoplasy, more likely the former.
Apleurus ( Apleurus ) angularis (LeConte), new combination
(Figs. 66, 72-74, 85, 93, 100, 211)
Cleonus angularis LeConte 1859a: 18. Lectotype (here designated), male, one of two syntypes (only one examined),
labelled with a pale green circle ( = Wyoming Territory) “C. angularis/ Beckwith Lee.”, “Type/ 5182” and with my
designation label "'Cleonus/ angularis/ LeC. LECTOTYPE/ desig. Anderson” (MCZC). Type locality, Kansas.
LeConte 1858 ( nomen nudum). Gemminger and von Harold 1871 (catal.). Hatch 1971.
Centrocleonus angularis ; LeConte 1876a (key, diag.). Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham
1889.
Dinocleus denticollis Casey 1891:180. NEW SYNONYMY. Lectotype (here designated), female, one of five syntypes,
labelled “Ari”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37269”, “D./ denticollis/ Cas.” and with my designation
label “ Dinocleus / denticollis/ Csy. LECTOTYPE/ desig. Anderson” (USNM). Type locality, Peach Springs,
Arizona. Wickham 1896 (check.). Casey 1904. Van Dyke 1953. Hatch 1971 (redesc.).
Dinocleus angularis ; Casey 1891 (key, redescr.). Wickham 1902 (check.). Faust 1904 (desig. as type species of Apleurus
Chevrolat; error as “ angulatus Lac”).
Dinocleus porcatus Casey 1904:321. NEW SYNONYMY Lectotype (here designated), female, one of two syntypes,
labelled “Ogden/ Ut. Solt”, “CASEY/ BEQUEST/ 1925”, “TYPE USNM/ 37270”, “ porcatus ” and with my
designation label “ Dinocleus / porcatus/ Csy. LECTOTYPE/ desig. Anderson” (USNM). Type locality, Ogden,
Utah. Fall and Cockerell 1907 (check.). Leng 1920 (catal.).
Cleonus denticollis ; Leng 1920 (catal.). Tanner 1966 (biol.).
Cleonus ( Dinocleus ) angularis-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) denticollis-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) porcatus-, Csiki 1934 (catal.).
Cleonus farctus-. Hardy and Andrews 1976 (distn., prob. misident.).
Cleonis angularis-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis denticollis-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis porcatus-, O’Brien and Wibmer 1982 (catal., distn.).
angularis, incertae sedis\ Chevrolat 1873.
Notes about synonymy. — Geographic patterns of variation in states of structural features
considered characteristic of Cleonus angularis LeConte, Dinocleus denticollis Casey, and
Dinocleus porcatus Casey, suggest that only a single species warrants recognition and that
these three names be placed in synonymy (see “Geographic variation” section).
I have only been able to locate and examine one of the two syntypes of Cleonus angularis
LeConte.
Problems in recognition. — Apleurus angularis and A. aztecus are the only two species of
Apleurus in which all individuals have the tarsal claws widely divergent and each lacking a
basal internal flange (Fig. 66). This character state should serve to separate nearly all A.
angularis from A. jacobinus and A. porosus. Most A. angularis can further be separated from
most A. jacobinus by the presence of long dorsal erect vestiture and markedly elevated
alternate elytral intervals in the latter. A very few specimens of A. jacobinus from Baja
California Norte have less markedly elevated alternate elytral intervals and have shorter dorsal
erect vestiture than throughout the rest of the species range and are thus difficult to separate
from A. angularis. Although these character states are such that they still allow for recognition
of these individuals as A. jacobinus , they can also be more easily recognized by the robust and
less markedly divergent tarsal claws. (See also “Problems in recognition” section for A.
porosus ).
Description. — Specimens examined. 218 males, 246 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 11.
Size. Length, male, 5.7-10.6 mm; female, 6.9-12.2 mm. Width, male, 2. 6-5. 2 mm; female, 3. 0-5. 6 mm. Head. Eye very
prominent and convex in dorsal view. Frons and vertex with moderately dense, small to moderately large, deep punctures.
Frons and area immediately above eyes with sparse short erect hair-like scales. Area immediately behind posterior margin
of eye with large, shallow to deep, irregularly impressed punctures. Area above eyes markedly elevated above rest of frons
Quaest. Ent., 1987, 23 (4)
482
Anderson
(eyes appearing distinctly browed in anterior view). Width of frons greater than to subequal to width at apex of rostrum.
Rostrum. Moderately robust (width at apex 0.66-0.91 times length in male; 0.62-0.92 in female) (Fig. 85). Median carina
slightly to well-developed, low and rounded but distinct, to high and sharp (Fig. 85). Dorsal and lateral punctation small,
moderately deep, sparse to moderately dense. Dorsally, excluding epistoma, with dense short, suberect hair-like scales, and
dense elongate-narrow to broad appressed white scales. In lateral view with apical portion steeply declivous from point of
antennal insertion to apex (Fig. 85a). Epistoma with apical margin rounded at middle (Fig. 85b). Mouthparts. Maxillary
palpus with stipes lacking large seta on outer margin (as in Fig. 65). Labial palpi separated by more or less width of basal
article of labial palpus (as in Figs. 62-63). Prementum with two or three pairs of large setae (as in Fig. 63). Pronotum. In
dorsal view with lateral margins subparallel from base to almost apical one-quarter, then divergent to apical one-quarter
(giving a distinctly laterally tuberculate appearance); markedly constricted at acute angle anterior to apical one-quarter,
then straight or slightly convergent to apex; widest at tubercles to subequal in width at base and apical one-quarter (Figs.
72-74). Dorsal and lateral punctation moderately large to large, deep, sparse to moderately dense, in some specimens areas
between punctures irregularly elevated thus apparently sculptured; punctures sparser and smaller on flanks. Scales white,
elongate-narrow to broad, appressed, sparse and small or lacking medially from disk in broad, apically narrowed patch
(except for midline in some specimens); dense laterally in pair of posterolaterally directed arcuate stripes; scales sparser at
lateral margins and ventrally on flanks, to uniformly dense laterally, at lateral margins and on flanks, individual large
punctures not obscured. Median carina lacking. Dorsally covered throughout with moderately dense, fine, short, erect
hair-like scales. Anterolateral margin with postocular projection absent or at most very slightly developed (Fig. 85a).
Prosternum. With shallow impression anterior to each procoxal cavity and with moderately to well-developed rounded
swelling anterior to each prosternal impression (Fig. 85a). Elytra. Moderately robust in general form (width at midlength
0.57-0.76 times length in males; 0.59-0.75 in females) (Figs. 72-74). In dorsal view with lateral margins slightly arcuate
throughout length or with margins slightly convergent to subparallel from apical one-third to humerus; humerus rounded
to obtuse, not distinct to distinct (Figs. 72-74). Sutural interval and intervals 3, 5, 7, and 9 with moderately dense, short,
erect hair-like scales; sparser on other intervals. Dorsally with sutural interval and intervals 3, 5, 7, and 9 slightly to
moderately swollen and convex (more so basally in some specimens). Scales elongate-narrow, various in density and size.
Pattern extremely various; many specimens with two variously sized posteromedially directed glabrous or nearly so areas,
one at basal one-third from intervals 4 to 6, other at apical one-third from intervals 2 to 7, and with various number of
scattered small glabrous (or nearly so) patches (Fig. 72); other specimens with glabrous areas of various extent on interval
2 at basal one-quarter and from midlength to apical one-quarter, interval 4 at basal one-third, interval 6 at basal
one-quarter and apical one-quarter, and interval 8 at midlength; scales confined to median portion of intervals, striae and
immediately adjacent portion of intervals lacking scales (Fig. 73). Wings. Absent (95.5%, N = 443) or present, long
(greater than elytra in length [4.5%, N = 21 ] ). Branches of 2A not complete, not joined at base. Legs. Foretibia of female
with inner margin with small denticles in apical one-third; subapical tooth small, not distinct from tibial denticles.
Foretibial and mesotibial unci of both sexes large, metatibial uncus of both sexes small. Tarsal claws widely divergent,
basal internal flange absent (Fig. 66). Ventral tarsal pilose vestiture of foretarsus of male absent from article 1, present as
minute apical tufts of article 2, as small apical pads of article 3; of foretarsus of female, mesotarsus of male and female,
and metatarsus of female, lacking from articles 1 and 2, present as minute to small pads at apex of article 3; of metatarsus
of male lacking from articles 1 and 2, lacking from to present as minute pads at apex of article 3. Mesosternum.
Mesosternal process flat to very slightly convex but not tumescent (as in Fig. 59). Abdomen. Ventral surface with
moderately dense, elongate-fine appressed white scales. Abdominal sterna III to VI (especially III and IV) with
moderately dense, individually distinct circular glabrous patches, each with large puncture and single long erect hair-like
scale situated in or near center. Apex of abdominal sternum VII of male with small dorsally directed median truncate
tooth. Abdominal tergum VIII uniformly convex, apical margin slightly elevated and reflexed. Genitalia. Female (nine
examined). Abdominal sternum VIII with lateral arms narrow, slightly outwardly arcuate in basal one-half, then parallel
to near apex, markedly inwardly arcuate immediately before apex (Fig. 93a). Gonocoxite II with stylus very small,
anteapical in position by more or less one-half length of stylus (Fig. 93b). Male (10 examined). Abdominal sternum VIII
with paired sclerite with inner apices with slightly to moderately developed ventral projections (as in Fig. lOle). Aedeagus
elongate-narrow, in lateral view thickest at midlength; in ventral view more or less parallel-sided throughout length.
Internal sac short and high, markedly apically deflexed; median dorsal pocket high, with moderately large dorsolaterally
directed paramedial lobe at midheight and midlength; small median lobe on dorsal surface immediately anterior to crest of
median dorsal pocket (Figs. 100a,b). Eversible apical sclerite complex with paired narrow scythe-like sclerites lightly
sclerotized, each lacking median projection (Fig. lOOd); adjacent ventral surface and basal portion of sides of apex of
internal sac with pair of indistinct lightly sclerotized transverse sclerites (Fig. 100a).
Geographic variation. — Geographic variation is extensive in a number of characters in
members of this species. As a result there has been a lot of confusion regarding the specific
limits of the previously recognized species (the names of which are here placed in synonymy)
and identity of various specimens. Specimens I examined bear label notes either indicating a
questionable identification or indicating that the specimen represents a probable new species.
One manuscript name is also represented on these labels.
New World Cleonini
483
Both macropterous and apterous individuals are known. Macropterous individuals have only
slightly elevat d and convex alternate elytral intervals and have more or less parallel-sided
elytra withdistinct humeri. Apterous individuals on the other hand have alternate elytral
intervals that are more markedly elevated and convex, and have elytra with the lateral margins
more arcuate with indistinct humeri. There appears to be no geographic component to this
variation because macropterous individuals, although few in number, are found throughout
most of the species range, with the notable exception of Colorado. This has no doubt
contributed to confusion, but the main cause of the confusion is due to the patterns of
geographic variation in other structural features. Specimens from eastern and central Colorado,
western Oklahoma, and western Texas (called hereafter the “eastern morph” and including
typical Cleonus angularis LeConte), are different from those to the west in California, Nevada,
Utah, and northern Arizona (called hereafter the “western morph” and including typical
Dinocleus denticollis Casey and Dinocleus porcatus Casey).
Eastern morphs (Fig. 73) differ from western morphs (Fig. 72) in states of the following
characters: shorter and more robust rostrum; rostral carina higher and sharper; eyes slightly
less prominent and convex in dorsal view; pronotal punctures denser, larger, deeper and more
irregularly impressed; slightly more markedly elevated alternate elytral intervals; generally a
more globose and robust elytral form; elytra with glabrous patches primarily transversely
oriented (Fig. 72) not longitudinally oriented (Fig. 73). Specimens from southern Arizona and
southern New Mexico however, are not reliably assignable to either form. They are
intermediate in many structural features which characterize the eastern and western morphs
and it is in this area that apparent intergradation takes place. Elytral scale patterns are not
clearly of one or the other form (Fig. 74); rostra are variable in form, and carinae both in form
and in degree of elevation; pronotal punctation is variable in depth, density and regularity of
impression; and elytral characters (other than scale pattern) are also variable. Based upon this
intergradation, I conclude that the eastern and western morphs and the individuals from
southern New Mexico and southern Arizona are all conspecific.
It appears that members of certain species of Apleurus (and also Cleonidius ) are largely
restricted to sandy habitats such as dune fields, sand hills, dry washes, stream beds, and
riparian habitats. These are either localized and discontinuous dune fields, or although
widespread, washes and riparian communities restricted in their degree of continuity. Dispersal
possibilities are therefore, I believe, markedly influenced by the continuity of drainage patterns
or other suitable habitats, and also as is true for all insects, by the presence or absence of wings.
Population distinctiveness and apparent lack of intergradation in the north in Colorado and
adjacent Utah and northern Arizona, is perhaps due to a combination of a higher continental
divide in this area with a more marked discontinuity between eastern and western drainage
patterns, consequently more localized and disjunct habitats, and general overall low number of
macropterous individuals including presence of only apterous individuals in Colorado. These
features may all combine to reduce the probability of dispersal between eastern and western
drainages resulting in local selection for differences between the two morphs. On the other
hand, this is not so in southern Arizona and New Mexico where the continental divide is much
lower, drainage patterns are not as markedly discontinuous and localized and where
macropterous individuals although still few in number, are present in both drainages. Dispersal
and interbreeding of individuals is therefore possible in this area with the result that there is no
local selection for differences between the two drainages.
Quaest. Ent., 1987, 23 (4)
484
Anderson
This pattern is similar to that found in C. canescens which also differs between Colorado
and adjacent Utah in the north but not so in the south in New Mexico and Arizona, and all
individuals of which are brachypterous and incapable of flight. Species which are found in these
same areas but which are macropterous do not show distinctive populations on the eastern and
western sides of the continental divide.
Geographic distribution. — This species is widespread in the southwestern United States of
America from Colorado, western Oklahoma and western Texas, west to Nevada in the north,
extreme southern coastal California and Baja California Norte, Mexico in the south (Fig. 211).
Natural history.— This species is associated with grassland, desert-grassland transitional
habitats, Pacific semi-desert, and Great Basin, desert. Specimens have been collected in sand
dune habitats in California, Arizona, Utah, and New Mexico, and otherwise in dry washes and
streambeds throughout the species range. Adults have been collected largely on Compositae as
follows: Chaenactis stevioides Hook, and Arn., Chrysothamnus nauseosus (Pall.) Britton, C.
viscidiflorus (Hook.) Nutt. var. pumilis, Gutierrezia lucida Greene, G. microcephala (DC.)
Gray, Aplopappus arcadenius (Greene) Blake, A. linearifolius DC (all Compositae); Atriplex
lentiformis (Torr.) Wats. (Chenopodiaceae). Adults have also been collected from roots of
Aplopappus tenuisectus (Greene) Blake and Psilostrope cooperi (Gray) Greene (Compositae).
It is likely that many shrubby Compositae, especially those on which adults are commonly
found, act as hosts for this species. Adults have been collected throughout the year at elevations
of from 15-2876 m (N = 63) (Fig. 232).
Adults have been collected in the stomach contents of the roadrunner, Geococcyx
californicus (Aves) at Carlsbad, New Mexico.
Chorological relationships. — This species is sympatric throughout nearly its entire range
with A. albovestitus', throughout Arizona, New Mexico, and western Texas with A. lutulentus ;
and throughout southeastern Arizona with A. saginatus. It is at most narrowly sympatric with
its close relatives A. jacobinus in southeastern California, and A. porosus in Baja California
Norte, Mexico.
Phylogenetic relationships. — Apleurus angularis is the sister-species of A. jacobinus (Fig.
235). Presence of the apotypic state of tarsal claws each lacking a basal internal flange in some
A. porosus and in A. angularis is a result of either hybridization or of homoplasy, more likely
the former.
Apleurus ( Apleurus ) jacobinus (Casey), new combination
(Figs. 67,71,86, 94, 101,208)
Centrocleonus pilosus LeConte 1 876a: 1 45 [not Chevrolat 1873:42]. NEW SYNONYMY Holotype (examined), male,
labelled “Calif.”, “type/ 5240”, “ Centrocleonus j pilosus Lee.” (MCZC). Type locality, California. Henshaw
1881-1882 (check.). Henshaw 1885 (check.).
Dinocleus jacobinus Casey 1891:179. Lectotype (here designated), female, one of eight syntypes, labelled “Cal.”, “Casey/
bequest/ 1925”, “TYPE USNM/ 37268”, “Z)./ jacobinus/ Cas.” and with my designation label “ Dinocleus /
jacobinus/ Casey LECTOTYPE/ desig. Anderson” (USNM). Type locality, San Diego, California. Fall 1901
(check.). Fall 1913. Leng 1920 (catal.).
Dinocleus pilosus-, Casey 1891 (key, redesc.). Fall 1901 (check.). Fall 1913. Leng 1920 (catal.). Essig 1958 (biol.,
misident. of Apleurus albovestitus).
Cleonus ( Dinocleus ) capillosus Csiki 1934:66. NEW SYNONYMY New name for Centrocleonus pilosus LeConte.
Blackwelder 1939 (check.).
Cleonus ( Dinocleus ) jacobinus-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) pilosus-, Csiki 1934 (catal., as jr. homonym).
Cleonis capillosus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis jacobinus-, O’Brien and Wibmer 1982 (catal., distn.).
New World Cleonini
485
Cleonis pilosus\ O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Notes about synonymy. — Based upon examination of very few specimens, Casey (1891)
distinguished Dinocleus jacobinus Casey from Dinocleus pilosus (LeConte) on the basis of the
smaller size, coarser darker and sparser elytral erect vestiture, more parallel-sided elytra with
exposed humeri, and less markedly elevated alternate elytral intervals of the former. Having
now examined large numbers of individuals I find that unequivocal assignment to either of the
forms is not possible because of variation in all but one of the above characters, and conclude
that the two forms are conspecific. I can see no differences in degree of coarseness or color of
the erect elytral vestiture as noted by Casey (1891).
Problems in recognition. — Many specimens of A. jacobinus are confused with individuals
of A. angularis and A. porosus. Characters and their states allowing for the separation of these
species from A. jacobinus are discussed in the respective “Problems in recognition” sections for
those species.
Description. — Specimens examined. 123 males, 255 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 12.
Size. Length, male, 7.8-1 1.8 mm; female, 5.8-14.3 mm. Width, male, 3. 7-5. 5 mm; female, 2. 8-6. 5 mm. Head. Eye very
prominent and convex in dorsal view. Frons and vertex with sparse to dense, large to very large, deep punctures. Frons and
area immediately above eyes with dense very long erect hair-like scales (short or lacking in few specimens). Area
immediately behind posterior margin of eye with large, deep, irregularly impressed punctures. Area above eyes markedly
elevated above rest of frons (eyes apparently distinctly browed in anterior view). Width of frons greater than width at apex
of rostrum. Rostrum. Moderately robust (width at apex 0.67-0.79 times length in male; 0.65-0.80 in female) (Fig. 86).
Median carina slightly to well-developed, low to high, rounded, in part obscured in many specimens by large deep
punctures (Fig. 86). Dorsal and lateral punctation large to very large, deep, moderately dense to dense. Dorsally, excluding
epistoma, with dense very long, erect hair-like scales (short or lacking from few specimens), and dense elongate-narrow
appressed white scales. In lateral view with apical portion steeply declivous from point of antennal insertion to apex (Fig.
86a). Epistoma with apical margin rounded at middle (Fig. 86b). Mouthparts. Maxillary palpus with stipes lacking large
seta on outer margin (as in Fig. 65). Labial palpi separated by more or less width of basal article of labial palpus (as in
Figs. 62-63). Prementum with two or three pairs of large setae (as in Fig. 63). Pronotum. In dorsal view with lateral
margins subparallel from base to almost apical one-quarter, then divergent to apical one-quarter (thus distinctly laterally
tuberculate in appearance); markedly constricted at acute angle anterior to apical one-quarter, then straight or slightly
convergent to apex; widest at tubercles to subequal in width at base and apical one-quarter (Fig. 71). Dorsal and lateral
punctation moderately large to large, deep, sparse to moderately dense, in some specimens areas between punctures
irregularly elevated thus appearing sculptured; punctures sparser and smaller on flanks. Scales white, elongate-fine to
elongate-narrow, appressed, sparse and small or lacking medially from disk in broad, apically narrowed patch (except for
midline in most specimens); dense laterally in pair of posterolaterally directed arcuate stripes; scales sparser at lateral
margins and ventrally on flanks, to uniformly dense laterally, at lateral margins and on flanks, not obscuring individual
large punctures. Median carina absent to present and distinct in some specimens. Dorsally covered throughout with sparse
to moderately dense, fine, very long, erect hair-like scales, each situated in large puncture. Anterolateral margin with
postocular projection absent or at most very slightly developed (Fig. 86a). Prosternum. With shallow impression anterior
to each procoxal cavity and with well-developed rounded swelling anterior to each prosternal impression (Fig. 86a). Elytra.
Moderately robust in general form (width at midlength 0.65-0.75 times length in males; 0.61-0.76 in females) (Fig. 71).
In dorsal view with lateral margins slightly arcuate throughout length; humerus rounded, not distinct (Fig. 71). Sutural
interval and intervals 3, 5, 7, and 9 with dense, very long, erect hair-like scales; sparser and slightly shorter on other
intervals (short or lacking in some specimens). Dorsally with sutural interval and intervals 3, 5, 7, and 9 moderately to
markedly swollen and convex (interval 5 at basal one-third and intervals 7 and 9 near base only slightly swollen in some
specimens). Scales elongate-narrow, various in density and size; pattern slightly various, but generally with two variously
sized posteromedially directed glabrous or nearly so areas, one at basal one-third from intervals 4 to 6, other at apical
one-third from intervals 2 to 7, and with various number of scattered small glabrous or nearly so patches (Fig. 71). Wings.
Absent. Legs. Foretibia of female with inner margin with small denticles in apical one-third; subapical tooth small, not
distinct from tibial denticles. Foretibial and mesotibial unci of both sexes moderately large to large, metatibial uncus of
both sexes small. Tarsal claws slightly divergent, basal internal flange present, slightly to well-developed (Fig. 67). Ventral
tarsal pilose vestiture of foretarsus of male absent of article 1, present as minute apical tufts of article 2, as small apical
pads to moderately large pads on apical one-third of article 3; of foretarsus of female, lacking from article 1, present as
minute to small pads at apex of articles 2 and 3; of mesotarsus and metatarsus of male, lacking from all three articles to as
minute apical pads of article 2, and as small apical pads to moderately large pads on apical one-third of article 3; of
mesotarsus and metatarsus of female, lacking from articles 1 and 2, present as minute apical tufts of article 3.
Mesosternum. Mesosternal process flat to very slightly convex but not at all tumescent (as in Fig. 59). Abdomen. Ventral
surface with moderately dense, elongate-fine appressed white scales. Abdominal sterna III to VI (especially III and IV)
Quaest. Ent., 1987, 23 (4)
486
Anderson
with moderately dense, individually distinct circular glabrous patches, each with large puncture and single long erect
hair-like scale situated in or near center. Apex of abdominal sternum VII of male with small dorsally directed median
truncate tooth. Abdominal tergum VIII uniformly convex, apical margin slightly elevated and reflexed. Genitalia. Female
(six examined). Abdominal sternum VIII with lateral arms narrow, slightly outwardly arcuate in basal one-half, then
parallel to near apex, markedly inwardly arcuate immediately before apex (Fig. 94a). Gonocoxite II with stylus very small,
anteapical in position by more or less one-half length of stylus (Fig. 94b). Male (seven examined). Abdominal sternum
VIII with paired sclerite with inner apices with slightly to moderately developed ventral projections (Fig. lOle). Aedeagus
elongate-narrow, in lateral view thickest at midlength; in ventral view more or less parallel-sided throughout length.
Internal sac short and high, markedly apically deflexed; median dorsal pocket high, with moderately large dorsolaterally
directed paramedial lobe at midheight and midlength; small median lobe on dorsal surface immediately anterior to crest of
median dorsal pocket (Figs. 101a, b). Eversible apical sclerite complex with paired narrow scythe-like sclerites lightly
sclerotized, each lacking median projection (Fig. 1 01 d); adjacent ventral surface and basal portion of sides of apex of
internal sac with pair of indistinct lightly sclerotized transverse sclerites (Fig. 101a).
Geographic variation. — Specimens from the southern part of the species range in extreme
southern California and Baja California Norte, Mexico have shorter and sparser dorsal erect
vestiture than do individuals from the northern inland portions of the range. These same
specimens tend to have the elytral intervals less markedly elevated throughout (Baja California
Norte) or have only intervals 5 and 7 variously swollen basally (extreme southern California).
Tarsal claws of the Baja California Norte individuals, although robust and not widely
divergent, each have only a slightly developed basal internal flange. All of these southern
individuals possess distinct glabrous patches on the abdominal sterna.
Two specimens from Point Reyes National Seashore and two specimens from Redwoods
Regional Park near Oakland lack or else have extremely short dorsal erect vestiture
(apparently not due to abrasion), but otherwise are typical A. jacobinus.
Geographic distribution. — This species is found in California from the San Francisco Bay
area south through coastal and central California to northern coastal Baja California Norte,
Mexico (Fig. 208).
Natural history. — This species is associated exclusively with Pacific semi-desert habitat.
Adults of have been collected on Aster sp., Hemizonia sp., H. pungens (H. and A.) T. and G.
(Compositae); beans (Leguminosae); Gossypium hirsutum L. (cotton; Malvaceae); and carrots
(Umbelliferae). They have also been found in various shipments of beans and raisins. Definite
hosts are not known. Adults have been collected throughout the year at elevations of from
5-424 m(N= 15) (Fig. 232).
Chorological relationships. — This species is sympatric throughout its range with western
A. albovestitus, and is at most narrowly sympatric with southern Californian A. angularis and
A. porosus from southern California or northern Baja California Norte, Mexico.
Phylogenetic relationships.- — Apleurus jacobinus is the sister-species of A. angularis (Fig.
235).
Apleurus ( Apleurus ) albovestitus (Casey), new combination
(Figs. 75-79, 88, 95, 103,209)
Cleonus molitor LeConte 1858:78 [not Gyllenhal 1834:174], NEW SYNONYMY Holotype (examined), female, labelled
with a gold circle ( = California), ''‘‘Cleonus/ molitor/ Lee.”, “Type/ 5183”, “Anobium!/ parasitic!” (MCZC). Type
locality, California. Gemminger and von Harold 1871 (catal.).
Centrocleonus molitor, LeConte 1876a (key, diag.). Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham
1889.
Dinocleus albovestitus Casey 1891:183. Lectotype (here designated), male, one of an undetermined number of syntypes,
labelled “Los Angeles/ Co. CAL”, “TYPE USNM/ 37273”, “CASEY/ bequest/ 1925”, “ZX/ albovestitus / Cas.” and
with my designation label “ Dinocleus albovestitus / Csy. LECTOTYPE/ desig. Anderson” and genitalia in microvial
under labels (USNM). Type locality, Los Angeles, California. Fall 1901 (check.). Casey 1904. Leng 1920 (catal.).
Ting 1936 (morphol.). Bruhn 1947 (morphol.). Sanders 1960 (morphol.).
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487
Dinocleus densus Casey 1891:185. NEW SYNONYMY Lectotype (here designated), male, one of four syntypes, labelled
“Winslow”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37277”, “D./ densus/ Cas.” and with my designation label
“ Dinocleus / densus Csy./ LECTOTYPE/ desig. Anderson” (USNM). Type locality, Winslow, Arizona. Wickham
1896 (check.). Leng 1920 (catal.).
Dinocleus molitor, Casey 1891 (key, redesc.). Fall 1901 (check.). Champion 1902-1906 (syn., distn.). Casey 1904. Leng
1920 (catal.).
Dinocleus wickhami Casey 1891:184. NEW SYNONYMY Holotype (examined), male, labelled “Indio/ California/
Wickham”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37276”, aD.f wickhami/ Cas.” (USNM). Type locality,
Indio, California. Fall 1901 (check.). Leng 1920 (catal.). Tanner 1934 (check.).
Dinocleus interruptus Casey 1904:322 [not Zoubkoff 1829:162]. NEW SYNONYMY Lectotype (here designated), male,
one of three syntypes, labelled “Ut”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37274”, “ interruptus ” and with my
designation label "Dinocleus/ interruptus Csy./ LECTOTYPE/ desig. Anderson” (USNM). Type locality, Utah.
Leng 1920 (catal.).
Dinocleus mexicanus Casey 1904:322. NEW SYNONYMY Lectotype (here designated), male, one of two syntypes,
labelled “Guer.”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37275”, “ mexicanus ” and with my designation label
” Dinocleus / mexicanus Csy./ LECTOTYPE/ desig. Anderson” (USNM). Type locality, Guerrero, Mexico.
Champion 1902-1906 (distn.).
Cleonus ( Dinocleus ) albovestitus ; Csiki 1934 (catal.).
Cleonus ( Dinocleus ) densus-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) interruptus-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) mexicanus-, Csiki 1934 (catal.).
Cleonus ( Dinocleus ) structor Csiki 1934:67. NEW SYNONYMY New name for Cleonus molitor LeConte. Blackwelder
1939 (check.).
Cleonus ( Dinocleus ) wickhami-, Csiki 1934 (catal.).
Cleonus mexicanus-, Blackwelder 1947 (check.).
Dinocleus pilosus; Essig 1958 (biol., misident.).
Cleonus albovestitus-. Hardy and Andrews 1976 (distn., biol.). Andrews et al. 1979 (distn., biol.).
Cleonis albovestitus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis densus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis interruptus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis mexicanus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis molitor, O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Cleonis structor, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis wickhami-, O’Brien and Wibmer 1982 (catal., distn.).
molitor, incertae sedis\ Chevrolat 1873.
Notes about synonymy. — I consider A. albovestitus to be composed of a number of largely
allopatric morphotypes most of which are distinguished solely on the basis of patterns of surface
vestiture (Figs. 75-79), but which exhibit continuous intergradation in zones of parapatry or
narrow sympatry (see “Geographic variation” section).
One of these morphotypes is largely black and glabrous, most of the surface vestiture having
been abraded (Figs. 78-79). This is Centrocleonus molitor LeConte. Dinocleus mexicanus
Casey is a form of this same morphotype, but in which the scales have not been extensively
abraded. Dinocleus densus Casey, is a form in which there has been no abrasion of scales. This
morphotype is confined to the Colorado and Gila River drainages.
The second morphotype, that with dense elytral and ventral abdominal scales and lacking
abdominal glabrous patches but with small elytral glabrous patches, includes Dinocleus
interruptus Casey, D. wickhami Casey, and D. albovestitus Casey (Figs. 76-77).
The third morphotype is recognized, among other characters, by the larger and more
numerous elytral and abdominal glabrous patches (Fig. 75).
Since I have been unable to find other structural features which correlate with the variation
in patterns of surface vestiture to allow for reliable separation of the various morphotypes, I
conclude that a single species is present, but with allopatric forms having different locally
adaptive patterns of surface vestiture.
Champion (1902-1906) incorrectly stated that D. mexicanus Casey is a new name for
Centrocleonus molitor LeConte.
Quaest. Ent., 1987, 23 (4)
488
Anderson
Problems in recognition. — Individuals of A. albovestitus might only be confused with those
of A. saginatus although key characters should serve to reliably separate all members of the
two species. Special note should be made of the marked secondary sexual dimorphism in extent
of ventral tarsal vestiture in A. saginatus that is not as marked in A. albovestitus. This allows
for easy recognition of males of A. saginatus.
All A. albovestitus that possess glabrous ventral abdominal patches are found only in
western California; those A. albovestitus found sympatrically with A. saginatus have extremely
dense scales on .the abdominal sterna and lack the glabrous patches present in all A. saginatus.
Description. — Specimens examined. 1251 males, 1470 females. Data about variation in LR, WF, WRA, LP,
WPB, WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in
Table 13. Size. Length, male, 11.8-16.8 mm; female, 12.8-21.4 mm. Width, male, 5. 1-7.2 mm; female, 5. 4-8. 8 mm.
Head. Eye slightly prominent and convex in dorsal view. Frons and vertex with sparse to moderately dense, small, shallow
punctures. Frons largely lacking suberect or erect vestiture, with at most only sparse, small suberect scales immediately
above eyes. Area immediately behind posterior margin of eye with small, shallow to moderately deep, irregularly
impressed punctures. Area above eyes continuous with and not elevated above rest of frons (eyes not browed in anterior
view). Width of frons greater than to subequal in width to apex of rostrum. Rostrum. Moderately robust (width at apex
0.62-0.78 times length in male; 0.63-0.79 in female) (Fig. 88). Median carina lacking. Dorsal and lateral punctation small
to moderately large, moderately deep, sparse to moderately dense. Dorsally, excluding epistoma, either lacking suberect or
erect vestiture or with at most scattered short suberect scales; and lacking (abraded) or with dense elongate-narrow
appressed white scales. In lateral view with apical portion steeply declivous from point of antennal insertion to apex (Fig
88a). Epistoma with apical margin rounded at middle (Fig. 88b). Mouthparts. Maxillary palpus with stipes with large seta
on outer margin (as in Fig. 64). Labial palpi separated by more or less width of basal article of labial palpus (as in Fig.
62-63). Prementum with two or three pairs of large setae (as in Fig. 63). Pronotum. In dorsal view with lateral margins
straight and subparallel to slightly convergent from base to almost apical one-quarter, not to slightly expanded laterally at
apical one-quarter (thus indistinctly laterally tuberculate in appearance); moderately constricted at obtuse to subacute
angle anterior to apical one-quarter, then straight and slightly convergent to apex; widest at base to subequal in width at
base and apical one-quarter (Figs. 75-79). Dorsal and lateral punctation small, shallow, sparse to moderately dense;
punctures sparser on flanks; outer margins of larger punctures not swollen or glabrous. Scales white, elongate-narrow to
broad, appressed, entirely lacking (abraded) to sparse and small or lacking medially from disk in broad, apically narrowed
patch (except for midline in most specimens); very dense laterally and ventrally on flanks onto prosternum; scales sparser
at lateral margins. Median carina lacking to present but irregularly developed but with at least a broad low rounded
median swelling in most specimens. Dorsally lacking suberect or erect vestiture or with sparse, short, erect hair-like scales,
each situated in large puncture. Anterolateral margin with postocular projection slightly to moderately developed (Fig.
88a). Prosternum. With shallow impression anterior to each procoxal cavity and with well-developed rounded swelling
anterior to each prosternal impression (Fig. 88a). Elytra. Moderately robust to moderately elongate-narrow in general
form (width at midlength 0.59-0.72 times length in males; 0.56-0.69 in females) (Figs. 75-79). In dorsal view with lateral
margins straight, slightly convergent to divergent from apical one-third to humerus; humerus rounded to obtuse, indistinct
to distinct, with at most only very few small glabrous shiny tubercles. Dorsal suberect or erect vestiture lacking to sparse,
short and fine. Dorsally with all intervals flat. Scales, if present, elongate-narrow to elongate-fine, white to golden. Scale
pattern extremely various; with scales entirely or largely lacking (abraded) except near apex on declivity (Figs. 78-79), to
present to various extent (not abraded), uniformly dense and large on sutural interval and intervals 3, 5, 7, and 9,
uniformly sparse and small on intervals 2, 4, 6, and 8, distinct glabrous patches absent (Fig. 77); or, to scales not abraded,
uniformly dense and large on sutural interval and intervals 3, 5, 7, and 9, and sparse and small to various degree on
intervals 2, 4, 6, and 8, with at most scattered but few glabrous patches present (Fig. 76); or, to sutural interval with scales
sparse and small or large and dense, otherwise with scales of elytral intervals extremely various in size and density, with
numerous irregularly distributed small and moderately large glabrous or nearly so patches present (Fig. 75). Wings. Short
(slightly shorter than elytra in length [0.2%, N = 4]), to long (slightly greater than elytra in length [99.8%, N = 1717]).
Branches of 2A complete and joined at base. Legs. Foretibia of female with inner margin with small to minute denticles in
apical one-third; subapical tooth minute, not distinct from tibial denticles. Foretibial and mesotibial unci of both sexes
moderately large to large, metatibial uncus of both sexes small. Tarsal claws slightly divergent, basal internal flange
present, well-developed (as in Fig. 67). Ventral tarsal pilose vestiture of foretarsus of male absent to present as minute
apical tufts on articles 1 to 3; of mesotarsus of male, lacking from articles 1 and 2, present as minute apical tufts of article
3; lacking from articles 1 to 3 of all tarsi of female and from articles 1 to 3 of metatarsus of male. Mesosternum.
Mesosternal process flat to very slightly convex but not at all tumescent (as in Fig. 59). Abdomen. Ventral surface either
with scales present, white, appressed, along posterior margins of abdominal sterna, also with scattered, suberect short
hair-like scales, each situated in large puncture; or, with white appressed scales uniformly very dense, abdominal sterna III
to VI without glabrous patches to with scales moderately dense, individually distinct circular glabrous patches present
(especially on abdominal sterna III and IV), each with large puncture and single short suberect hair-like scale situated in
or near center. Apex of abdominal sternum VII of male with large dorsally directed median truncate tooth. Abdominal
New World Cleonini
489
tergum VII of female slightly medially longitudinally carinate, apical margin moderately elevated and reflexed; abdominal
tergum VIII of female markedly medially longitudinally carinate, apical margin moderately elevated and reflexed.
Genitalia. Female (12 examined). Abdominal sternum VIII with lateral arms broad, straight and slightly divergent from
base to apical one-third, slightly inwardly arcuate from apical one-third to apex (Fig. 95a). Gonocoxite II with stylus
absent, with darkly sclerotized slightly elevated dorsal subapical ridge (Fig. 95b). Male (eight examined). Abdominal
sternum VIII with paired sclerite with inner apices with well-developed ventral projections (Fig. 103e). Aedeagus
elongate-narrow, in lateral view thickest at midlength; in ventral view more or less parallel-sided throughout length.
Internal sac short and high, slightly apically deflexed; median dorsal pocket high, with moderately large dorsolaterally
directed paramedial lobe near dorsal margin at midlength; small median lobe on dorsal surface immediately anterior to
crest of median dorsal pocket (Figs. 103a,b). Eversible apical sclerite complex with paired narrow scythe-like sclerites
darkly sclerotized, each with long dorsoapically directed median projection (Fig. 103d); adjacent ventral surface and basal
portion of sides of apex of internal sac with pair of indistinct lightly sclerotized transverse sclerites (Fig. 103a).
Geographic variation. — Variation in the pattern of surface vestiture in members of this
species is very extensive (Figs. 75-79). There is also slight variation in other structural features
but these could not be reliably correlated with the variation in vestiture to allow recognition of
more than one species (see also “Notes about synonymy” section). However, I recognize three
largely allopatric or parapatric morphotypes. In the first of these, vestiture is abraded or at
least subject to abrasion. Such specimens are primarily black and glabrous (except for the
tibiae and tarsi) and possess moderately dense to dense scales only along the lateral margins of
the elytra and the posterior margins of the abdominal sterna (especially V and VI), and to a
various extent in some specimens, also in the apical one-third of the elytra, onto the elytral
declivity (Figs. 78-79). Scattered single scales are also found variably on the thoracic and
abdominal sterna, elytra and femora. In individuals in which scales are variably present on the
elytra, the scales are uniformly dense and large on all intervals to slightly less dense and smaller
on intervals 2, 4, and 6. Scales are very easily abraded on these specimens. Individuals of this
morphotype have been collected at elevations of from -67-870 m (N = 23).
This form is found throughout the southern portion of the Colorado River drainage, the
western portion of the Gila River drainage, and throughout Imperial County, California.
Individuals from the latter area tend to have the elytral scales denser and not abraded and
although lacking distinct glabrous elytral patches, the general scale pattern clearly grades into
that pattern to be discussed next in which small elytral glabrous patches are present (Fig. 76).
Individuals of this morphotype seem restricted to Pluchea sericea (Nutt.) Coville (Compositae)
in sand dune habitats along the margins of waterways in these areas.
A second form of A. albovestitus is also found in these and other areas, and although the
ranges of the two morphotypes narrowly overlap, they have not been collected together at the
same time and place. Individuals of this form possess large and dense elytral surface vestiture
that is not prone to abrasion. All abdominal sterna have uniformly very dense scales; no
glabrous patches are present. Elytral scales are uniformly large and dense to very dense, but are
lacking or very sparse in scattered small to moderately large patches on elytral intervals 2 to 8
(Fig. 76). Scales of the sutural interval of many of these specimens are uniformly small and
sparse. Punctures of elytral striae are small and indistinct. This form is widespread from
Nevada and Utah south to extreme western Texas, New Mexico, Arizona and southeastern
California, south into Baja California Sur, Mexico. Individuals of this form in Utah and
northern Arizona, along the northern Colorado and Virgin River drainages, differ from those
elsewhere by largely lacking glabrous or nearly so patches on the elytra (Fig. 77). Scales of
some individuals in these areas are less dense and smaller on intervals 2, 4, and 6, much the
same pattern as in specimens of the largely glabrous morphotype in which elytral scales have
not been extensively abraded. This similarity in scale pattern suggests the notion that the
largely glabrous individuals of morphotype 1 in the southern portion of the Colorado River
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490
Anderson
drainage may have reached this area as a result of downriver dispersal from parent populations
along the Virgin and northern Colorado Rivers. Individuals of this morphotype have been
collected at elevations of from 2-1446 m (N = 49).
A third pattern of vestiture is found in individuals from southern coastal to northern coastal
and central California. In this form, scales of the elytra are less dense and smaller than in the
previous forms, and the elytral glabrous or nearly so patches are generally larger and more
numerous (Fig. 75). Punctures of the elytral striae are generally slightly larger and more
distinct than in the other morphotypes. Abdominal sterna (especially III and IV) of many
individuals with this elytral scale pattern possess small to moderately large indistinct to distinct
glabrous areas, each with a single short suberect hair-like scale situated in the center. Suberect
vestiture, lacking from the elytra on the other morphotypes, is short, sparse and fine on the
elytra of many of these specimens. Elytral scale patterns of this and of the second morphotype
completely intergrade in southern California. Individuals of this third morphotype have been
collected at elevations of from -41-3185 m (N = 61).
The potential adaptive significance of variation in surface vestiture is not known but is
suspected to be influenced by thermoregulatory effects, or, more likely, by cryptic effects
resulting from similarity in color and pattern to the ground substrate (see “Evolutionary
Trends” section).
Brachypterous specimens of A. albovestitus are only known from two localities in Baja
California Norte, Mexico.
Geographic distribution. — This species is distributed from New Mexico and western Texas
west through Utah and Nevada to northern California in the north, through Arizona and
extreme northwestern Mexico to southern California and Baja California Sur, Mexico in the
south (Fig. 209).
Natural history. — This species is associated with a variety of habitats as follows;
desert-grassland transitional. Pacific semi-desert, and Great Basin, Chihuahuan, Mojave, and
Sonoran deserts. Adults of this species have been collected in sand dune habitats in California,
Arizona, Utah, Nevada (label data; Andrews et al. 1979; Hardy and Andrews 1976) and
otherwise primarily in dry washes and stream beds throughout the species range. Andrews et
al. 1979 tentatively suggest that the species is an obligate sand associate.
Adults have been collected on a wide variety of plants as follows: Ephedra sp., E. californica
Wats. (Ephedraceae); Ambrosia sp., A. psilostachya DC., Artemesia tridentata Nutt., Baileya
multiradiata Harv. and Gray, B. pleniradiata Harv. and Gray, Baccharis glutinosa Pers.,
Chrysothamnus nauseosus (Pall.) Britton, Dicoria sp., Franseria sp., F. confertiflora (DC.)
Rydb., F. dumosa Gray, Flourensia cernua DC., Gutierrezia microcephala (DC.) Gray,
Aplopappus acradenius (Greene) Blake, Hymenoclea sp., H. salsola T. and G., Hymenothrix
mslizeni Gray, “ Palafoxia arida'\ P. linearis (Cav.) Lag., Pluchea sericea (Nutt.) Coville,
Verbesina enceloides (Cav.) Benth. and Hook., Xanthium sp., (all Compositae); Larrea
tridentata (DC.) Coville (Zygophyllaceae); Hordeum sp., Sitanion sp., (Graminae); Datura
sp., (Solanaceae); Gossypium sp., (Malvaceae); Eriogonum fasciculatum Benth. var.
polifolium (Polygonaceae); Astragalus sp., Cercidium floridum Benth., Medicago sativa L.,
Olneya tesota Gray, Prosopis juliflora (Swartz) DC. (all Leguminosae); Atriplex sp., A.
lentiformis (Torr.) Wats., Salsola kali L., Sarcobatus vermiculatus (Hook.) Torr. (all
Chenopodiaceae); Vitus sp. (Vitaceae); Prunus persica Batsch. (peach; Rosaceae); Raphanus
sativus L. (radish; Cruciferae). No definite hosts are known, but the vast majority of plant
association are with Compositae; records of occurrence on members of other families are
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491
generally single instances of but one or two individuals and very likely represent chance
occurrences. I suspect that a wide variety of Compositae serve as host plants for this species.
Adults have been collected throughout the year at elevations from -67-3185 m (N=133)
(Fig. 232).
As noted in the “Geographic variation” section, individuals of each of the three morphotypes
occur over slightly different elevational ranges.
Chorological relationships. — This species is sympatric with A. lutulentus in Arizona, New
Mexico and western Texas; with A. angularis throughout most of its range, with the exception
of northern California; with A. jacobinus throughout entire range of that species in California;
narrowly sympatric with A. porosus in Baja California Norte, Mexico and southern California;
and sympatric with its sister-species A. saginatus throughout Arizona, southern New Mexico,
and extreme nothwestern Mexico.
I have caught large numbers of specimens of A. albovestitus together with A. lutulentus on
Chrysothamnus nauseosus (Compositae) in the vicinity of Portal, Arizona and Animas, New
Mexico.
Phylogenetic relationships. — Apleurus albovestitus and A. saginatus are sister-species
(Fig. 235).
Apleurus ( Apleurus ) saginatus (Casey), new combination
(Figs. 63-64, 80, 87, 96, 104, 206)
Dinocleus saginatus Casey 1891:182. Holotype (examined), male, labelled “Ariz.”, “CASEY/ bequest/ 1925”, “TYPE
USNM/ 37272”, “/)./ saginatus/ Cas.” (USNM). Type locality, Arizona. Leng 1920 (catal.).
Dinocleus dentatus Champion 1902-1906:100. NEW SYNONYMY Holotype (examined), female, labelled “Type”,
inverted “Sp. figured”, “Pinos Altos/ Chihuahua,/ Mexico/ Buchan-Hepburn”, “B.C.A. Col. IV. 4./ Dinocleus/
dentatus,/ Champ.”, inverted “ dentatus , Ch.” (BMNH).Type locality, Pinos Altos, Chihuahua, Mexico.
Cleonus ( Dinocleus ) dentatus ; Csiki 1934 (catal.).
Cleonus dentatus-, Blackwelder 1947 (check.).
Cleonus ( Dinocleus ) saginatus-, Csiki 1934 (catal.).
Cleonis dentatus-, O'Brien and Wibmer 1982 (catal., distn.).
Cleonis saginatus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — Following examination of types and numerous specimens I regard
Dinocleus saginatus Casey and D. dentatus Casey as conspecific.
Problems in recognition. — Individuals of this species might only be confused with those of
A. albovestitus (see “Problems in recognition” section for A. albovestitus ).
Description. — Specimens examined. 597 males, 657 females. Data about variation in LR, WF, WRA, LP, WPB,
WPT, WE1M, LEI, WPB/LP, WPT/WPB, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 14.
Size. Length, male, 14.9-19.5 mm; female, 15.9-21.8 mm. Width, male, 5. 8-8.0 mm; female, 6. 2-9.0 mm. Head. Eye
slightly prominent and convex in dorsal view. Frons and vertex with sparse, small, shallow punctures. Frons largely lacking
suberect or erect vestiture, with at most only sparse, small suberect scales immediately above eyes; white elongate-narrow
appressed scales dense above eyes, sparser medially. Area immediately behind posterior margin of eye with small, shallow
to moderately deep, irregularly impressed punctures. Area above eyes continuous with and not elevated above rest of frons
(eyes not browed in anterior view). Width of frons greater than to subequal in width to apex of rostrum. Rostrum.
Moderately robust (width at apex 0.63-0.73 times length in male; 0.63-0.74 in female) (Fig. 87). Median carina lacking
to variously developed as low rounded to sharp fine glabrous line. Dorsal and lateral punctation small, shallow, sparse.
Dorsally, excluding epistoma, with short scattered suberect scales; and with dense elongate-narrow appressed white scales.
In lateral view with apical portion steeply declivous from point of antennal insertion to apex (Fig. 87a). Epistoma with
apical margin rounded at middle (Fig. 87b). Mouthparts. Maxillary palpus with stipes with large seta on outer margin
(Fig. 64). Labial palpi separated by more or less width of basal article of labial palpus (Fig. 63). Prementum with two or
three pairs of large setae (Fig. 63). Pronotum. In dorsal view with lateral margins slightly arcuate to straight and
subparallel to slightly convergent from base to almost apical one-quarter, not to slightly expanded laterally at apical
one-quarter (thus indistinctly laterally tuberculate in appearance); moderately constricted at obtuse to subacute angle
anterior to apical one-quarter, then straight and slightly convergent to apex; widest at base to subequal in width at base
Quaest. Ent., 1987,23 (4)
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Anderson
and apical one-quarter (Fig. 80). Dorsal and lateral punctation small, shallow, sparse to moderately dense; punctures
sparser and shallower on flanks; outer margins of larger punctures swollen, glabrous and shiny (appearing as small
tubercles). Scales white, elongate-narrow to broad, appressed sparse and small or lacking medially on disk in broad,
apically narrowed patch (except for midline in most specimens); very dense laterally and ventrally on flanks onto
prosternum; scales sparser at lateral margins. Median carina lacking to present and irregularly developed in basal one-half
as low, broad glabrous line. Dorsally with sparse, short, erect hair-like scales, each situated in large puncture.
Anterolateral margin with postocular projection slightly developed (Fig. 87a). Prosternum. With shallow impression
anterior to each procoxal cavity and with well-developed rounded swelling anterior to each prosternal impression (Fig.
87a). Elytra. Moderately robust to moderately elongate-narrow in general form (width at midlength 0.59-0.62 times
length in males; 0.56-0.64 in females) (Fig. 80). In dorsal view with lateral margins straight or nearly so, slightly
convergent from apical one-third to humerus; humerus obtuse, distinct, with numerous distinct small glabrous shiny
tubercles (also present at bases of adjacent elytral intervals) (Fig. 80). Dorsal suberect or erect vestiture lacking. Dorsally
with all intervals flat. Scales appressed, white to golden in color, various in size and density. Scale pattern various, with
numerous small irregularly distributed glabrous or nearly so patches (Fig. 80). Wings. Long (slightly greater than elytra in
length). Branches of 2A complete and joined at base. Legs. Foretibia of female with inner margin with small to minute
denticles in apical one-third; subapical tooth minute to small, not distinct from tibial denticles. Foretibial and mesotibial
unci of both sexes moderately large to large, metatibial uncus of both sexes small. Tarsal claws slightly divergent, basal
internal flange present, well-developed (as in Fig. 67). Ventral tarsal pilose vestiture of foretarsus of male present as small
rounded apical pads of article 1, as moderately large rounded pads on apical one-half of article 2, and as large rounded
pads on apical one-half of article 3; of foretarsus of female, lacking from articles 1 and 2, present as minute apical tufts of
article 3; of mesotarsus of male, absent to present as minute apical tufts of article 1, present as small apical pads of article
2, and present as moderately large round pads on apical one-third of article 3; of metatarsus of male, absent from article 1,
present as minute apical tufts of article 2, and present as small apical pads of article 3; lacking from articles 1 to 3 of
mesotarsus and metatarsus of female. Mesosternum. Mesosternal process flat to very slightly convex but not tumescent (as
in Fig. 59). Abdomen. Ventral surface with very dense elongate-narrow to broad, white, appressed scales; abdominal sterna
III to VI (especially III and IV) with moderately dense individually distinct large circular glabrous shiny patches, each
with large puncture and single short suberect hair-like scale situated in or near center. Apex of abdominal sternum VII of
male with large dorsally directed median truncate tooth. Abdominal tergum VII of female slightly medially longitudinally
carinate, apical margin moderately elevated and reflexed; abdominal tergum VIII of female markedly medially
longitudinally carinate, apical margin moderately elevated and reflexed. Genitalia. Female (six examined). Abdominal
sternum VIII with lateral arms broad, straight and slightly divergent from base to near apex, slightly inwardly arcuate
near apex (Fig. 96a). Gonocoxite II with stylus absent, with darkly sclerotized slightly elevated dorsal subapical ridge
(Fig. 96b). Male (six examined). Abdominal sternum VIII with paired sclerite with inner apices with well-developed
ventral projections (as in Fig. 103e). Aedeagus elongate-narrow, in lateral view thickest at midlength; in ventral view more
or less parallel sided throughout length. Internal sac short and high, slightly apically deflexed; median dorsal pocket high,
with moderately large dorsolaterally directed paramedial lobe near dorsal margin at midlength (Figs. 104a,b). Eversible
apical sclerite complex with paired narrow scythe-like sclerites darkly sclerotized, each with long dorsoapically directed
median projection (Fig. 104d); adjacent ventral surface and basal portion of sides of apex of internal sac with pair of
indistinct lightly sclerotized transverse sclerites (Fig. 104a).
Geographic distribution. — This species is found in southeastern Arizona and extreme
southwestern New Mexico, south into northwestern Mexico (Fig. 206).
Natural history. — This species appears restricted to dry washes and riparian habitats in the
desert-grassland transitional habitat bordering the eastern Sonoran desert region. Adults have
been collected on Baccharis glutinosa Pers., Chrysothamnus nauseosus (Pall.) Britton,
Hymenoclea monogyra Torr. and Gray (all Compositae). Definite hosts are not known. Adults
have been collected throughout the year at elevations of from 283-1820 m (N = 36) (Fig. 232).
Chorological relationships. — This species is sympatric throughout its range with A.
angularis, A. lutulentus, and its sister-species A. albovestitus. I know of no records of A.
saginatus being caught with any of these three species.
Phylogenetic relationships. — Apleurus saginatus and A. albovestitus are sister-species
(Fig. 235).
Genus Cleonidius Casey
Curculio ; Pallas 1781 (sp. desc.). Herbst 1795.
Cleonus\ Say 1831 (sp. desc.). Melsheimer 1853 (in part; check.). Gemminger and von Harold 1871 (in part; catal.).
LeConte 1876a (key, sp. descs.). Henshaw 1881-1882 (check.). LeConte and Horn 1883 (key). Henshaw 1885
New World Cleonini
493
(check.). Wickham 1899. Wickham 1896 (check.). Fall 1897 (sp. desc.). Fall 1901 (check.). Wickham 1902 (check.).
Fletcher 1906. Fall and Cockerell 1907 (in part; check.). Pierce 1907 (biol.). Chittenden 1911 (biol.). Ely 1913 (biol.).
Gibson 1914. Anderson 1914. Blatchley and Leng 1916 (key, sp. redescs.). Yothers 1916. Leng 1920 (in part; catal.).
Leonard 1926 (in part; check.). Boving 1927 (in part; larval key). Bradley 1930 (in part; key). Wilcox et al. 1934.
Brimley 1938 (check.). Blackwelder 1947 (in part; check.). Bruhn 1947 (morphol.). Essig 1958 (biol.). Sanders 1960
(morphol.). Tanner 1966 (in part; sp. redesc., biol.). Kingsolver 1972. Kumar et al. 1976 (biol.). Arnett et al. 1980
(key, biol.).
Rhynchophorus\ Say 1831 (in part; sp. desc.).
Cleonis; Kirby 1837 (sp. desc.). O’Brien and Wibmer 1982 (catal., distn.). O’Brien and Wibmer 1984 (in part).
Lixus; Zoubkoff 1833 (sp. desc.). Boheman 1836 (sp. desc.). Randall 1838 (in part; sp. desc.). Mannerheim 1843 (sp.
desc.). Motschulsky 1845 (sp. desc.). Gemminger and von Harold 1871 (in part; catal.). Capiomont and Leprieur 1874
(sp. redesc.). Sprague and Austin 1875. LeConte 1876a (in part; key, sp. descs.). Henshaw 1881-1882 (in part;
check.). LeConte and Horn 1883 (in part; key). Henshaw 1885 (in part; check.). Wickham 1889. Faust 1890. Casey
1891 (in part; key). Horn 1894 (in part; check.). Petri 1905 (key). Petri 1912. Blatchley 4914 (sp. desc.). Blatchley
and Leng 1916 (in part; key, sp. redesc.). Leng 1920 (in part; catal.). Lukjanovitsh 1926. Blatchley 1930. Bradley
1930 (in part; key). Arnett 1960-1962 (in part; key). Kissinger 1964 (in part; key).
Apleurus ; Chevrolat 1873 (in part, especes lyxiformes; check., key, sp. descs.).
Cleonidius Casey 1891:186. As subgenus of Cleonus. Gender, masculine. Type species Cleonis vittatus Kirby ( = Lixus
poricollis Mannerheim) by subsequent designation (Faust 1904:190). Champion 1902-1906 (sp. redesc.). Csiki 1934
(catal.). Blackwelder 1939 (check.). Arnett 1960-1962 (key, catal.). Kissinger 1964 (key). Hatch 1971 (key, sp.
redescs.)
Cleonidius ; Faust 1904 (key, check., type species desig.).
Lixestus Reitter 1916:89. As subgenus of Lixus. NEW SYNONYMY. Gender, masculine. Type species Curculio vibex
Pallas by monotypy. Csiki 1934 (catal.).
Lixus (Lixesthus); Ter-Minasyan 1978 (error, misspelling).
Notes about synonymy. — The subgenus Cleonidius was first proposed by Casey (1891) to
accomodate those Cleonus with a cylindrical rostrum, more or less vittate elytral scale pattern,
and an elongate-narrow body form approaching that of Lixus. Indeed species of Cleonidius
very closely resemble Lixus and species of the two genera are often difficult to separate. This is
evidenced by the placement of some species, now regarded as Cleonidius , originally in Lixus ,
and by the herein proposed new synonymy of Lixestus Reitter, a subgenus of Lixus , with
Cleonidius.
Among Lixus species examined, only Lixus ( Lixestus ) vibex Pallas (type species of
Lixestus ), was found to possess a small dorsal median tubercle at the basal margin of variously
abdominal sterna V to VII, that is characteristic of Cleonidius. However, other Lixus species,
as yet not examined for this character, may also prove to require inclusion in Cleonidius should
they possess the apotypic state as noted above. This is not surprising given the similarity of the
two genera, their apparent close phylogenetic relationship, and previous lack of consideration of
Cleonidius species in past studies and resulting classifications of Palearctic Lixus species. Thus
there is the need for a reevaluation of the validity of at least the subgeneric groupings of Lixus
in this light. Whether Cleonidius will remain as a valid genus when this is done remains to be
seen; undoubtedly Lixus species will require reclassification.
Prior to the present study, Cleonidius had been given generic status by Faust (1904), only.
Diagnosis. — Adult Cleoninae with elongate-narrow to moderately robust body form (Figs.
118-136). Eye more or less oval to elongate-oval, flat (Figs. 137-156). Rostrum
elongate-narrow to moderately robust, with at most only low median carina (indicated in the
majority of species by low glabrous shiny line) (Figs. 137-156). Antennal funiculus with article
1 longer than article 2; article 2 more or less as wide as long (Figs. 137-156). Pronotal
postocular lobes lacking to present and well-developed; postocular vibrissae uniformly short to
long but of unequal length, longest immediately behind base of eye (Figs. 137-156). Pronotal
disk slightly to distinctly vittate, with white scales largest and/or densest immediately laterad
of midline, smaller and/or sparser laterally; largely whitish in color, underlying dark cuticle
largely obscured by white scales (Figs. 118-136). Prosternal swellings absent or present, if
Quaest. Ent., 1987, 23 (4)
494
Anderson
present, situated immediately anterior to each procoxal cavity (Fig. 8). Ventral tarsal pilosity
extensive (covering greater part of ventral surface of each tarsal article) to reduced in extent or
lacking from at least the basal tarsal articles of some species. Tibia with corbel ridge rounded
(Fig. 5). Abdominal sternum VII (at least) of females dorsally (internally) with variously
developed basal median glabrous shiny tubercle, evident externally as shallow impression.
Abdominal sternum VIII of female with basal arm short to long (Figs. 157-176).
Description. — Size. Small to moderately large; elongate-narrow to moderately robust in general body form (Figs.
118-136). Mouthparts. Prementum flat to slightly swollen ventrally; with as many as three large setae on each side.
Maxillary palpus with palpifer and stipes each with at least one large seta. Labial palpi separated by distance subequal to
width of basal article of labial palpus. Rostrum. Elongate-narrow to robust, not to markedly medially tumescent, with at
most only low median carina indicated in majority of species by fine glabrous shiny line (Figs. 137-156). Epistoma not to
moderately swollen, with apical margin emarginate medially (Figs. 137b- 156b). Antenna with funiculus with article 1
longer than article 2; article 2 more or less as long as wide; apical three articles of club lacking placoidal sensillae. Head.
Eye oval to elongate-oval; flat (Figs. 137-156). Upper margin of eye rounded, frons flat to variously convex. Vestiture.
Dorsum with suberect to erect vestiture absent to present and dense, short to very long in length; with simple appressed
white scales of various size and density. Pronotum. Dorsal surface punctate, median basal area shallowly to deeply
impressed; disk with or without various other impressions. Pronotal disk slightly to distinctly vittate, with white scales
largest and/or densest immediately laterad of midline, smaller and/or sparser laterally; largely whitish in color, underlying
dark cuticle medially largely obscured by white scales (Figs. 118-136). Lateral margins with white scales large and/or
dense. Pronotum widest at base to subequal in width from base to near apex, lateral margins arcuate, more or less parallel
or slightly convergent from base to near apex, then variously constricted and more convergent to apex (Figs. 118-136).
Pronotal postocular lobes lacking to present and well-developed; postocular vibrissae uniformly short (length less than
one-half width of an eye) to long but of unequal length, longest behind base of eye (greatest length greater than one-half
width of an eye) (Figs. 137a- 156a). Prosternum. With shallow, rounded impression anterolaterad of each procoxal cavity;
with or without variously developed rounded swelling immediately anterior to each procoxal cavity (Fig. 8). Legs. Tarsus
moderately broad to broad; articles 2 and 3 more or less subequal in length, width and length of each article subequal to
slightly wider than long; article 1 only slightly longer than articles 2 or 3; article 3 moderately deeply bilobed. Ventral
tarsal pilosity various in extent from dense and covering almost entire ventral surface of each tarsal article, to lacking
entirely from at least more basal articles in few species. Claws connate only at very base to from very base through basal
one-third to midlength, not to markedly divergent. Tibia with corbel ridge rounded (Fig. 5). Foretibia with inner margin
with small to large denticles in apical one-half to two-thirds; near apex with small, indistinct to moderately large subapical
tooth. Metatibia of male with uncus with ventral margin evenly rounded. Wings. Absent or present (various in length).
Elytra. All intervals equally flat except humerus and very base of interval 3 variously swollen and convex. Scale pattern
various, more or less vittate (Figs. 131, 133-134, 136) or as marginal band of large white scales (Figs. 118-123). Humeri
acute to rounded. Abdomen. Ventral surface with small shiny glabrous patches, each with single small appressed to
suberect scale-like seta. Female with base of abdominal sternum VII (in some individuals also sterna V and VI) internally
(dorsally) with variously developed rounded glabrous shiny median tubercle, evident externally (ventrally) as rounded
shallow impression. Genitalia. Female. Abdominal sternum VIII with basal arm short to long; lateral arms various in
shape (Figs. 157-176). Gonocoxite II elongate triangular in shape; apex not prolonged into marked lobe; stylus moderately
large, apical in position (Fig. 117). Spermathecal gland round (Fig. 116). Male. Paired sternite of abdominal sternum VIII
lacking distinct basal projections (as in Fig. 97f). Aedeagus moderately robust; in lateral view more or less evenly arcuate
throughout length; apex not spatulate (Figs. 177c-196c). Internal sac various, with median dorsal pocket low to high;
apical and dorsal median pocket individually distinct or not; various lobes present (Figs. 177a,b-196a,b). Apical sclerite
complex present, individual sclerites distinctly scythe-like, simple, lacking median projection.
Comparisons. — Cleonidius species will prove most difficult to distinguish from various
Lixus species. Some Old World species presently regarded as Lixus , but not examined by me,
may prove to be Cleonidius. In North America, north of Mexico, the area where most species
of Cleonidius are found, no Lixus species has a rounded pronotal postocular lobe; at most they
possess a variously developed, but usually small, acute postocular projection or else have the
anterolateral margin of the pronotum straight to slightly sinuate (Fig. 4). Postocular vibrissae
in most New World Lixus are of unequal length, long, and have their greatest length behind
the base of the eye (Fig. 4). All New World Lixus I have examined which possess a variously
developed postocular lobe and that have postocular vibrissae of more or less uniform length
have the inner margin of the femora variously dentate. Femora are not dentate in any
Cleonidius species.
New World Cleonini
495
Unfortunately there are species of Cleonidius which lack or have only slightly developed
postocular lobes and have postocular vibrissae as in most Lixus species (Figs. 145a,
152a- 155a). These can be distinguished from Lixus by the relative lengths of the antennal
articles, by having a more robust rostrum, and by having the ventral tarsal pilosity variously
reduced in extent.
Although there are exceptions (Fig. 141a), world Lixus species in general have a very
elongate-narrow rostrum (Fig. 4) whereas Cleonidius species have a more robust rostrum (Figs.
137a- 140a, 142a- 156a). Eyes are rounded and slightly convex in many Lixus , otherwise are
oval to elongate-oval as in all Cleonidius. Whereas characters used herein to separate Lixus
from Cleonidius work for the New World fauna, the structural diversity of Lixus species in
other geographic areas does not permit separation on the basis of these same characters. In the
Old World, some Lixus species examined are separable from Cleonidius species only by lack of
the internal tubercle at base of abdominal sternum VII in females, the presence of which is, by
definition, universally diagnostic for Cleonidius. Such is the situation with Cylindropterus
luxeri Chevrolat, Lixus (Lixoglyptus) spartii Olivier, and Lixus (Lixoglyptus) circumcinctus
Boheman, the taxa chosen as the out-groups for the phylogenetic analysis of relationships of
species of Cleonidius.
Apleurus (Gibbostethus) hystrix Fall, because of its more elongate-narrow body form (Fig.
24), may be confused with some Cleonidius species, but the tumescent mesosternal process
(Fig. 58), robust and deeply punctate rostrum, and prosternal swellings immediately in front of
the prosternal impressions (Fig. 7) in the former will readily separate the two.
Checklist of included species. — Nineteen species are recognized in the New World from
Nicaragua north to southern Canada. The genus also occurs in the Palearctic Region where it
is represented by at least, species formerly placed in the subgenus Lixestus Reitter of Lixus
Fabricius. Of species placed in Lixestus , I have examined, and therefore herein include only
Lixus ( Lixestus ) vibex (Pallas), the type species of that subgenus by monotypy. Subsequently,
aside from species regarded as synonyms of L. vibex, Csiki (1934) included L. pallasi Faust,
originally described as a variation of L. vibex and therefore likely a Cleonidius, and
Ter-Minasyan (1978) further included L. meles Boheman. Inclusion of L. meles in Lixestus by
Ter-Minasyan (1978) is supported by Petri (1905) wherein L. vibex and L. meles key out
adjacent to one another. A key to separate the three species included in Lixestus is given by
Ter-Minasyan (1978).
The twenty species herein placed in Cleonidius are as follows:
Cleonidius erysimi species group
1. C. erysimi (Fall)
2. C. eustictorrhinus Anderson
3. C. pleuralis (LeConte)
4. C. subcylindricus Casey
5. C. longinasus Anderson
6. C. texanus (LeConte)
Cleonidius americanus species group
7. C. americanus Csiki
8. C. frontalis (LeConte)
9. C. canescens (LeConte)
10. C. infrequens Anderson
Quaest. Ent., 1987,23 (4)
496
Anderson
11. C. puberulus (LeConte)
12. C. collaris (LeConte)
13. C. notolomus Anderson
Cleonidius poricollis species group
14. C. poricollis (Mannerheim)
15. C. calandroides (Randall)
Cleonidius boucardi species group
16. C. boucardi (Chevrolat)
17. C. trivittatus (Say)
18. C. placidus Csiki
19. C. quadrilineatus (Chevrolat)
Incertae sedis
20. C. vibex (Pallas)
Phylogenetic relationships. — See “Phylogenetic analysis” section.
Key to species of adult Cleonidius
1 Tarsal claws widely divergent, lacking basal internal flange (as in Fig.
66) C. vibex (Pallas), p. 544
V Tarsal claws at most only slightly divergent, basal internal flange
well-developed (as in Fig. 67) 2
2 (F) Head above eye with deep straight sulcus extended posteroventrally from
above dorsal margin of eye to area under postocular lobe (Fig. 139). Elytra
with apices produced and acuminate (Fig. 120)
C. pleuralis (LeConte), p. 504
2' Head above eye lacking deep straight sulcus. Elytra with apices rounded to
at most only slightly acuminate and slightly produced 3
3 (2') Pronotum extremely irregularly elevated; markedly constricted
dorsolaterally near apical margin, deeply and broadly impressed medially
at base and longitudinally along lateral margins; with larger punctures on
disk very sparse, shallow and indistinct. Metathoracic wings short
(approximately equal to one-half length of elytra)
C. collaris (LeConte), p. 523
3' Pronotum more regular in elevation and sculpture; only shallowly to
moderately deeply impressed medially at base (also shallowly
longitudinally along lateral margins in very few specimens); with larger
punctures on disk denser, deeper and more distinct. Metathoracic wings
short (approximately equal to or less than one-half length of elytra) to long
(approximately equal to or greater than length of elytra) 4
4 (3") 2 Pronotum with postocular lobes moderately to well-developed and with
postocular vibrissae short to moderately long (maximum length equal to or
less than one-half width of eye in lateral view) (Figs. 137a- 144a,
2An intermediate specimen will key out through both halves of this
couplet.
New World Cleonini
497
1 46a- 151a). Prosternum in lateral view with anterior flat portion one-third
as long as, to approximately equal in length to, more posterior angulate
portion (Figs. 137a-144a, 1 46a- 151a) 5
4 ' Pronotum with postocular lobes lacking to at most moderately developed
and with postocular vibrissae long (maximum length equal to or greater
than one-half width of eye in lateral view) (Figs. 145a, 152a- 155a).
Prosternum in lateral view with anterior flat portion no more than one-half
as long as more posterior angulate portion (Fig. 145a, 152a- 155a) 19
5 (4) Elytra with sutural intervals lacking scales throughout the greater part of
their length, with at most scattered very fine erect hair-like scales (Fig.
118); interval 2 with scales contrastingly dense. . . C. erysimi (Fall), p. 500
5' Elytra with sutural interval with scales uniformly distributed throughout
their length (although sparsely so or abraded in some specimens); interval 2
with scales similar in density to those of sutural interval 6
6 (50 Head behind eye with moderately deep curved sulcus extended
posteroventrally from upper one-half of posterior margin of eye to area
under postocular lobe (largely covered in many specimens by postocular
lobe) (Fig. 150) 7
6' Head behind eye lacking distinct curved sulcus, with at most one or more
wrinkles of cuticle 8
7 (6) Distributed in the northern Atlantic coastal states of the United States and
Atlantic coastal provinces of Canada (Fig. 229). Dorsal surface of
pronotum and elytra with suberect vestiture short, indistinct. Elytral scale
pattern of most specimens not distinctly vittate; scales on interval 2 and
intervals 6 to 8 not markedly smaller than scales on adjacent intervals;
underlying dark cuticle on interval 2 and intervals 6 to 8 largely obscured
by overlying white scales (Fig. 132)
C. calandroides (Randall) (in part), p. 531
7' Distributed in Mexico, and/or Canada and the United States west of 85°
W longitude (Fig. 228). Dorsal surface of pronotum and elytra with erect
vestiture short to moderately long, distinct. Elytral scale pattern distinctly
vittate; scales on interval 2 and intervals 6 to 8 markedly smaller than
scales on adjacent intervals; underlying dark cuticle on interval 2 and
intervals 6 to 8 not largely obscured by overlying white scales (Fig. 131) .
C. poricollis (Mannerheim) (in part), p. 527
8 (6') Elytra distinctly laterally margined with white scales (especially in basal
one-half); intervals 9 to 1 1 with dense moderately elongate broad white
scales; otherwise moderately densely uniformly covered with fine white
scales smaller than those on intervals 9 to 1 1 (Figs. 1 19, 121-123) 9
8' Elytra not laterally margined, either vittate (Figs. 130-131), more or less
uniformly densely scaled (Fig. 129), or mottled with irregularly distributed
patches of larger and denser scales (Figs. 124-125); intervals other than 9
to 1 1 with scales equally large to those on intervals 9 to 1 1 12
9 (8) Head, pronotum and elytra lacking dorsal erect or suberect vestiture (Fig.
140a). Postocular lobes only moderately developed (Fig. 140a)
C. subcylindricus Casey, p. 506
Quaest. Ent., 1987,23 (4)
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Anderson
9' Head, pronotum and elytra with dorsal erect or suberect vestiture short to
long (Figs. 138a, 141a-142a). Postocular lobes well-developed (Figs.
138a, 141 a- 142a) 10
10 (9') Rostrum with large deep dense punctures; medially longitudinally
tumescent and with median carina as fine shiny line at crest of median
tumescence; robust (width at apex greater than 0.58 times length) (Fig.
138) C. eustictorrhinus Anderson, p. 502
10" Rostrum with small shallow moderately dense punctures; not medially
longitudinally tumescent, carina lacking to as variously developed fine
shiny line; robust to elongate-narrow (width at apex less than 0.62 times
length) (Figs. 141-142) 11
11 (10') Rostrum (especially in females) very long and narrow (width at apex less
than 0.50 times length) (Fig. 141a). Head, pronotum and elytral declivity
(and to a lesser extent elytral disk) with dorsal erect vestiture moderately
long to very long (lacking from a very few specimens) (Fig. 141a).
Associated with Leguminosae C. longinasus Anderson, p. 508
1 1 ' Rostrum (of both sexes) shorter and more robust (width at apex greater
than 0.50 times length) (Fig. 142). Head, pronotum and elytra with dorsal
suberect to erect vestiture short to moderately long (Fig. 142a). Associated
primarily with Rosaceae and Rhamnaceae, rarely Leguminosae at lower
altitudes C. texanus (LeConte), p. 510
12 (8') Dorsum (especially of head, pronotum and elytral declivity) with erect
vestiture long to very long (Fig. 143a). Rostrum in lateral view with
prementum swollen ventrally (Fig. 143a); in dorsal view slightly expanded
laterally at midlength (Fig. 143b) C. americanus Csiki, p. 513
12' Dorsum with suberect to erect vestiture either lacking or short variously on
head, pronotum, and elytra (Figs. 144a, 146a-147a, 149a-152a). Rostrum
in lateral view with prementum flat to slightly swollen ventrally (Fig.
144b); in dorsal view not to slightly expanded laterally at midlength (Fig.
144a) 13
13 (120 Distributed in the northern Atlantic coastal states of the United States and
Atlantic coastal provinces of Canada (Fig. 229)
C. calandroides (Randall), p. 531
13' Distributed in Mexico, and/or Canada and the United States west of 85°
W longitude 14
14 (130 All tarsi with similar and extensive ventral pilose vestiture, articles 1 to 3
with large elongate-oval ventral pilose pads on virtually entire ventral
surface of each article (except basal one-third to one-half of article 1) 15
14' Tarsi with ventral pilose pads small, decreased in extent on more posterior
tarsi, either with articles 1 and 2 with small elongate-narrow ventral pilose
pads, apical tufts of pilosity, or, with pads lacking; article 3 with
moderately large pads not on more than apical two-thirds of ventral surface 16
15 (14) Rostrum not dorsally medially tumescent; base in lateral view appearing
continuous with frons (Fig. 152a). Elytra with interval 3 (in some
specimens also intervals 4 and 5), and intervals 9 to 1 1 with dense white
New World Cleonini
499
scales, otherwise with scales very sparse or lacking (Fig. 133)
C. boucardi (Chevrolat) (in part), p. 534
15' Rostrum dorsally medially longitudinally tumescent; base in lateral view
separated from frons by distinct transverse impression (Fig. 150a). Elytra
with scales white, fine and sparse, densest on sutural interval, intervals 3 to
7, and 9 to 1 1 C. poricollis (Mannerheim) (in part), p. 527
16 (14') Rostrum very short and robust (width at apex greater than 0.70 times
length) (Figs. 146-147) 17
16' Rostrum more elongate-narrow (width at apex less than 0.70 times length)
(Figs. 144, 149) 18
17 (16) Rostrum with median carina very distinct, sharp and elevated, also with
faint lateral carinae at lateral margins (Fig. 146b); rostrum flat from
immediately laterad of median carina to lateral margins; rostrum more or
less quadrate in cross-section, lateral margins sharp. Articles of tarsus of
hind leg with ventral pilose pads moderately large. Metathoracic wings
long (greater than length of elytra) to short (approximately one-half or less
than length of elytra) C. infrequens Anderson, p. 520
17 Rostrum with median carina indistinct, low and only slightly elevated,
without trace of lateral carinae (Fig. 147b); rostrum declivous from
immediately laterad of median carina to lateral margins; rostrum less
quadrate in cross-section, the lateral margins rounded. Articles of tarsus of
hind leg with ventral pilose pads small. Metathoracic wings short (less than
one-half length of elytra) C. puberulus (LeConte), p. 522
18(16') Elytra elongate compared to pronotal length (LP/LE1 less than 0.40) (Fig.
125). Metathoracic wings long (greater than elytra in length). Rostrum
more or less circular in cross-section, lateral margins rounded; in dorsal
view, slightly swollen laterally at midlength; in lateral view with
prementum slightly swollen ventrally. Fore-tibia of most females with inner
margin with very large prominent denticles (Fig. 144c)
C. frontalis (LeConte), p. 515
18' Elytra more robust compared to pronotal length (LP/LE1 more than 0.40)
(Fig. 130). Metathoracic wings present, varied in length from short
(approximately equal to one-half length of elytra) to long (greater than
elytra in length). Rostrum more or less quadrate in cross-section, lateral
margins sharp; in dorsal view, not swollen laterally at midlength; in lateral
view with prementum flat. Fore-tibia of female with inner margin with
slightly developed, small denticles C. notolomus Anderson, p. 525
19 (4') Metathoracic wings short, approximately equal to one-half length of elytra.
Eye oval (width greater than 0.60 times length) (Fig. 145a)
C. canescens (LeConte), p. 518
19' Metathoracic wings long (approximately equal to or greater than length of
elytra). Eye elongate-oval (width less than 0.65 times length) (Figs. 152a-
155a) 20
20 (19') Elytra, except sutural interval and portions of interval 2, uniformly covered
with dense white scales (Fig. 135) C. placidus Csiki, p. 539
20' Elytra distinctly vittate, intervals 6 to 8 with scales lacking or very sparse
Quaest. Ent., 1987, 23 (4)
500
Anderson
compared to intervals 3 to 5 and 9 to 1 1 (Figs. 133-134, 136) 21
21 (20') Rostrum not or only slightly dorsally medially tumescent, not distinctly
carinate (very slightly so in few specimens) (Fig. 152a). Metatarsus with
articles 2 and 3 with ventral pilose pads large, onvirtually entire ventral
surface; article 1 with pads elongate-broad, on apical two-thirds of the
ventral surface C. boucardi (Chevrolat) (in part), p. 534
2 1' Rostrum dorsally medially tumescent, with low rounded carina (Figs. 153a,
155a). Metatarsus with article 3 with ventral pilose pads moderately large,
covering the apical one-half to two-thirds of ventral surface; article 2 with
pads elongate-narrow, covering the apical two-thirds of the ventral surface;
article 1 with pads either present as apical tufts, or elongate and very
narrow, covering not more than the apical one-half of the ventral surface 22
22 (2L) Elytra elongate-narrow (width less than 0.55 times length) (Fig. 136).
Pronotal disk with moderately long erect hair-like scales (Fig. 155a).
Associated primarily with Rosaceae and Rhamnaceae, rarely
Leguminosae C. quadrilineatus (Chevrolat), p. 541
22 ' Elytra more robust (width greater than 0.55 times length) (Fig. 134).
Pronotal disk with at most short, suberect, indistinct hair-like scales (Fig.
153a). Associated with Leguminosae C. trivittatus (Say), p. 536
Cleonidius erysimi species group
Diagnosis. — Size small to moderate for Cleonidius (Fig. 197). Dorsal erect or suberect
vestiture absent (Fig. 140a) to present, dense and very long (Fig. 141a). Rostrum very
elongate-narrow (Fig. 141a) to moderately robust (Fig. 138a), not to markedly medially
tumescent, not to variously carinate; lateral margins rounded. Pronotum with postocular lobes
moderately (Fig. 140a) to well-developed (Fig. 141a); postocular vibrissae uniformly short to
more or less uniformly moderately long. Elytra with marginal band of scales, only intervals 9 to
1 1 with white scales large and broad, otherwise with intervals with scales small and fine to
absent (Figs. 1 18-123). Tarsus with ventral pilose vestiture extensive (on almost entire ventral
surface of each article) to slightly reduced in extent (on not less than apical one-half of articles
2 and 3). Wings present, length various. Female with abdominal sternum VIII with basal arm
short; lateral arms more or less straight and divergent throughout length (Figs. 157-162). Male
aedeagus with internal sac with apex of dorsal median pocket rounded (Figs. 177b- 182b); lobes
B, D, and E absent in some species (Figs. 180a,b-182a,b); dorsal median pocket low to high in
lateral view.
Phylogenetic relationships. — The C. erysimi group is hypothesized to be sister group to the
remainder of North American Cleonidius (Figs. 236-237).
Cleonidius erysimi (Fall)
(Figs. 118, 137, 157, 177,219)
Lixus poricollis-, LeConte 1876a,b (misident.). Henshaw 1885 (check., misident.).
Cleonus erysimi Fall 1901:261. Lectotype (here designated), male, one of an undetermined number of syntypes, labelled
‘“male symbol4”, “Redondo/ Cal. 4.7.94.”, “Type/ erysimi ”, “M.C.Z./ Type/ 25193”, “H.C. FALL/
COLLECTION” and with my designation label "'Cleonus/ erysimi/ Fall LECTOTYPE/ desig. Anderson” (MCZC).
Type locality, Redondo, California. Leng 1920 (catal.).
Cleonus ( Cleonidius ) erysimi ; Csiki 1934 (catal.).
New World Cleonini
501
Cleonis erysimi ; O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — This species has infrequently been erroneously referred to as
Lixus poricollis Mannerheim.
Problems in recognition. — Individuals of this species are easily recognized by the sutural
interval of the elytra lacking scales, with at most only scattered very fine erect hair-like scales;
interval 2 has scales that are contrastingly dense (Fig. 118). Although some individuals of C.
pleuralis are superficially similarly patterned, close examination of sutural interval reveals
small scales. Regardless, C. pleuralis individuals are easily recognized by the deep straight
sulcus above each eye (Fig. 139).
Individuals of C. eustictorrhinus and C. texanus, especially the former, may also prove
difficult to separate from those of C. erysimi should the elytra be largely abraded of vestiture.
Cleonidius texanus specimens have a slightly more elongate-narrow rostrum (Fig. 142) and the
length of the elytra as compared to the length of the pronotum is greater (see Figs. 197, 200).
Cleonidius eustictorrhinus and C. erysimi individuals will prove most difficult to separate, if
abraded, although most C. erysimi are brachypterous, whereas all C. eustictorrhinus are
macropterous.
Description. — Specimens examained. 129 males, 179 females. Data about variation in LR, WF, WRA, LP,
WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are
presented in Table 15. Size. Length, male, 7.2-1 1.3 mm; female, 6.1-12.0 mm. Width, male, 2.2-4. 1 mm; female, 2. 1-4.5
mm. Head. Eye elongate-oval. Area behind eye with numerous wrinkles of cuticle. Frons with sparse to moderately dense,
small, shallow to moderately deep punctures, some longitudinally confluent and irregularly impressed immediately above
eyes, punctures sparser medially; interspersed with minute punctures. Frons also with moderately dense to dense,
elongate-narrow erect hair-like scales immediately above eyes; otherwise with scattered erect hair-like scales and
moderately dense to dense, elongate-narrow appressed white scales, sparse to absent medially. Frons separated from base
of rostrum by moderately deep to deep transverse impression (Fig. 137a). Base of median tumescence or carina of rostrum
with small moderately deep fovea. Width of frons greater than or subequal to width at apex of rostrum. Rostrum.
Moderately robust, slightly more so in males (width at apex 0.57-0.80 times length) (Fig. 137). In lateral view straight to
very slightly curved downward. In dorsal view with postgenae not to very slightly expanded laterally, antennal scrobes very
slightly visible. Slightly to markedly medially tumescent from point of antennal insertion to middle of frons (more
markedly so basally), with median carina variously developed as rounded to sharp, moderately elevated, fine to broad,
glabrous shiny line at crest of median tumescence. Dorsal and lateral punctation moderately dense to dense, small to
moderately large, shallow to deep (longitudinally confluent in many specimens), smaller and not longitudinally confluent
apically, dorsally interspersed with minute punctures. Dorsally (especially laterally) with sparse to moderately dense,
moderately long erect hair-like scales, and laterally with sparse to moderately dense, elongate-narrow white appressed to
recumbent scales, medially with vestiture absent to very sparse, appressed, elongate-fine. Epistoma not swollen. Pronotum.
Median carina variously developed in anterior one-half to three-quarters as slightly to moderately elevated glabrous shiny
broad line. Dorsal punctation moderately large to large, moderately dense to dense, deep; smaller and shallower apically;
shallower on flanks; areas between large dorsal punctures with minute regularly impressed punctures. Dorsally with
moderately dense, moderately long erect hair-like scales each situated in large puncture (Fig. 137a). Scales dorsally absent
along midline, moderately dense, elongate-fine, appressed in pair of paramedian apically slightly narrowed stripes;
laterally, absent to very small and sparse. Lateral margins with appressed white scales elongate-fine to elongate-narrow,
moderately dense in moderately broad stripe. Flanks with scales smaller and slightly less dense than along lateral margins.
Median basal area of disk very shallowly to moderately deeply impressed. Anterolateral margin with pronotal postocular
lobes moderately to well-developed; postocular vibrissae short, of uniform length to very slightly longer immediately
behind base of eye (approximately one-half width of eye or less) (Fig. 137a). In dorsal view with lateral margins very
slightly arcuate and convergent from base to apex; subapical constriction very slight and indistinct. Prosternum. With
moderately large, moderately deep to deep impression, and moderately to well-developed rounded swelling anterior to each
procoxal cavity. In lateral view with anterior flat to slightly angulate portion long, approximately one-half to two-thirds
length of posterior more angulate portion, anterior and posterior portions not to slightly differentiated by shallow
transverse impression (Fig. 137a). Elytra. Elongate-narrow in general form (width at midlength 0.45-0.55 times length in
males; 0.48-0.53 in females) (Fig. 1 18). In dorsal view with lateral margins straight to very slightly arcuate or very slightly
sinuate, slightly convergent from apical one-third to humerus; slightly to moderately and evenly arcuate from apical
one-third to apex. Elytral apices slightly produced, rounded to sub-acuminate, not to slightly divergent. Humerus
indistinct, rounded. Dorsally with sparse to moderately dense, suberect to erect, short hair-like scales (abraded in many
specimens). Sutural interval with scales lacking throughout greater part of length, with sparse fine scales only at very base;
intervals 2 to 7 with scales uniformly moderately dense, various in size from elongate-fine to elongate-narrow on alternate
intervals in many specimens; interval 8 with scales absent to very small, fine and sparse; intervals 9 to 1 1 with scales
Quaest. Ent., 1987, 23 (4)
502
Anderson
uniformly moderately dense to dense, moderately large and elongate-narrow, sparser and smaller apically and at base of
interval 11 (Fig. 118). Punctures of elytral striae small, individually distinct, larger and less individually distinct basally,
arranged in regular rows. Wings. Long (greater than length of elytra [9%, N = 29]) to short (less than one-half length of
elytra [91%, N = 279]). Legs. Foretibia of female with inner margin with small denticles in apical three-quarters;
subapical tooth small, indistinct to distinct from tibial denticles. Foretibial and mesotibial unci of both sexes moderately
large, metatibial uncus small. Tarsal claws connate in basal one-third to one-half, not to slightly divergent, each with basal
internal flange well-developed. Ventral tarsal pilose vestiture of all tarsi of male and female present as elongate-narrow
pads on apical one-half to three-quarters of article 1, as rounded pads on apical one-half of article 2, and as large rounded
pads on apical two-thirds to more or less entire ventral surface of article 3. Abdomen. Ventral surface with sparse to dense
appressed elongate-narrow to very elongate-narrow white scales (abraded in part in many specimens). Abdominal sterna
III and IV laterally with individually indistinct glabrous patches, each with large puncture and single elongate recumbent
to suberect hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with
moderately developed median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm
short; lateral arms straight and divergent from base to apical one-third; markedly inwardly arcuate at apical one-third and
convergent to apex, moderately expanded at apex (Fig. 157). Stylus moderately large in length compared to length of
gonocoxite II. Male (four examined). Internal sac (Figs. 177a,b) elongate and low; median dorsal pocket low, with single
elongate moderately broad dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on
dorsal surface (lobe B), with small paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), with
moderately large paired dorsolaterally directed lobe at midlength near dorsal margin (lobe D), with moderately large
paired dorsolaterally directed lobe at apical one-third at dorsal margin (lobe E), and with small ventrally directed paired
lobe near base near ventral margin (lobe F). Apex of median dorsal pocket rounded in dorsal view. Apical pocket
individually distinct from median dorsal pocket. Ventral median pocket moderately large. Apical pocket with paired small
ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple
scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is found from southern British Columbia and
Alberta, east to Manitoba, south to southern California, Arizona and extreme western Texas
(Fig. 219).
Natural history. — This species is associated with grassland, desert-grassland transitional
habitats, Pacific semi-desert, and Great Basin desert. Adults of this species have been collected
on a variety of plants (but especially Cruciferae) as follows; Cleome sp. (Capparaceae);
Aplopappus ericoides (Less.) H. and A., Senecio sp. (both Compositae); Brassica sp., Brassica
oleracea L. (cabbage), Dithyrea californica Harv., Erysimum sp., E. suffrutescens (Abrams)
G. Rossb., Raphanus sativus L. (radish), Stanleya pinnata (Pursh.) Britton var .inyoensis (all
Cruciferae); Adenostoma fasciculatum H. and A., Prunus persica (L.) Batsch. (peach)
(Rosaceae). The type series of Cleonus erysimi Fall was collected from flowers of Erysimum
capitatum (Dougl.) Greene on sandhills at Redondo Beach, California (Fall 1901). Adults have
been reared from larvae found in crowns of Lobularia maritima (L.) Desv. (Cruciferae) at
Harris Ranch, Los Alamos, Santa Barbara Co., California.
Adults of the species have been frequently collected in sand dune habitats in coastal
California and in other sandy habitats throughout the species range. Adults have been collected
throughout the year at altitudes of 15-2359 m (N = 40) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister species of C. eustictorrhinus , the two
species forming a lineage that is a member of an unresolved trichotomy including C. pleuralis
and the C. subcylindricus-C. longinasus-C. texanus lineage (Fig. 237).
Cleonidius eustictorrhinus Anderson, new species
(Figs. 119, 138, 158, 178, 216)
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “SACRAMENTO/ CALIF/
XII-1 3-1949”, “D. GIULIANI/ COLLECTOR”, “Derham Giuliani/ Collection/ Calif. Acad. Sci./ Accession 1967”,
“Collection of the/ CALIFORNIA ACADEMY/ OF SCIENCES, San/ Francisco Calif.”, “HOLOTYPE/ Cleonidius/
eustictorrhinus/ Anderson” and with abdomen on card and genitalia in microvial attached to pin (CASC). Allotype,
New World Cleonini
503
female, with a red label “ALLOTYPE”, “Orange Vale/ Sacto.Co./ July 31, 1938/ Quentin Tornich”, ’’ALLOTYPE/
Cleonidius/ eustictorrhinus / Anderson” and with abdomen on card and genitalia in microvial attached to pin (UCBC, on
indefinite loan to CASC). Type locality, Sacramento, California.
Paratypes. 11 males, 12 females. UNITED STATES OF AMERICA: California: Orange Vale, 31. VII. 38, Q.
Tornish, IF (UCBC); Lancaster, 19.V.37, E.P. Van Duzee, 1M (CASC); 4mi. w. Meadow Valley, 13.V.49, Middlekauff,
IF (UCBC); Madera County, Bates, 25.III.17, IF (AMNH); Antioch, 1.IX.37, M. Cazier, 1M (AMNH), 26.IV.68,
Monroe, IF (CWOB); Irvine, Davis, 1M (CNCI); San Diego County, 26.IV.91, Blaisdell, 1M (CASC); Stanford, III.04,
1M (LACM), 11. III. 06, IF (LACM); Fresno County, Waltham Canyon, 26. III. 37, Blum, IF (RSAN); Baldwin Hills,
11. VII. 53, Menke, IF (LACM); Vine Hill, 23.V.65, Johnson, IF (CWOB); Atascadero, 18.VI.46, Mansfield, 1M
(CASC); Laguna Beach, 30.XII.49, 1M (CWOB); Idyllwild, 12.V.34, Stone, IF (OSUC); Laguna, 20.VI.30, IF
(OSUC), 30.VI.30, 1M (CNCI); Irvine, 1M (RSAN); Yosemite, 17. VI. 31, Essig, IF (UCBC); Stanislaus, XII. 35, 1M
(CFDA); Carmichael, 17.V.58, Wilkey, 1M (CDFA); Napa County, Pope Valley, 9.V.55, Raven, IF (CASC).
Derivation of specific epithet. — From the Greek “ew” meaning very or exceedingly;
“ stiktos ” meaning punctured; and, “ rhinos ” meaning nose. This name is used in reference to
the markedly punctured rostrum of members of this species.
Problems in recognition. — Individuals of this species are likely to be confused only with C.
texanus, C. subcylindricus, and C. erysimi. Individuals of C. eustictorrhinus are easily
separated from sympatric individuals of C. erysimi by the sutural interval lacking scales in the
latter (Fig. 1 18) and by most C. erysimi being brachypterous whereas all C. eustictorrhinus are
macropterous. Individuals of C. eustictorrhinus are easily separated from C. subcylindricus by
geographic distribution, the former being restricted to California (Fig. 216), the latter to the
Atlantic Coastal Plain from Florida north to New York (Fig. 215). C. subcylindricus
individuals also lack suberect or erect dorsal vestiture (Fig. 140a) whereas C. eustictorrhinus
have short but distinct erect dorsal vestiture (Fig. 138a). Specimens of C. eustictorrhinus will
prove most difficult to separate from sympatric individuals of C. texanus. In C. eustictorrhinus
individuals (Fig. 138), the rostrum is medially tumescent, distinctly carinate and has large deep
punctures; in C. texanus specimens (Fig. 142), the rostrum is not medially tumescent or
carinate and has only small shallow punctures. The rostrum is slightly more robust in C.
eustictorrhinus (Fig. 138) than in C. texanus (Fig. 142), C. eustictorrhinus tend to be larger
than C. texanus, and the length of the elytra as compared to the length of the pronotum tends
to be greater in C. texanus than in C. eustictorrhinus (see Figs. 197, 200).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 16. Size. Length, male, 8.8-10.4 mm;
female, 8.8-11.0 mm. Width, male, 3. 2-3. 9 mm; female, 3. 2-4.0 mm. Head. Eye elongate-oval. Area behind eye with
numerous wrinkles of cuticle. Frons with moderately dense, small to moderately large, shallow punctures; longitudinally
confluent and very irregularly impressed immediately above eyes; larger punctures interspersed with sparse minute
punctures. Frons also with scales sparse to moderately dense elongate-narrow, erect immediately above eyes; medially and
laterally with sparse to moderately dense elongate-narrow appressed to suberect white scales; laterally with moderately
dense, short to moderately long fine erect hair-like scales. Frons either continuous with base of rostrum, or slightly
separated by shallow transverse impression (Fig. 138a). Base of median tumescence or carina of rostrum with small
shallow to moderately deep fovea. Width of frons greater than or subequal to width at apex of rostrum. Rostrum.
Moderately robust, slightly more so in males (width at apex 0.56-0.69 times length) (Fig. 138). In lateral view slightly
curved downward. In dorsal view with postgenae not to slightly laterally expanded, antennal scrobes slightly visible.
Moderately to markedly medially tumescent from point of antennal insertion to middle of frons, with median carina
variously developed as a rounded, slightly to moderately elevated, broad, glabrous, shiny line at crest of median
tumescence. Dorsal and lateral punctation moderately dense to dense, moderately large to large, shallow to deep
(longitudinally confluent in many specimens), smaller and shallower apically, dorsally interspersed with few minute
punctures. Dorsally (especially laterally) with sparse to moderately dense, moderately long, suberect to erect hair-like
scales, and laterally and medially with sparse to moderately dense, elongate-fine to elongate-narrow, appressed to
recumbent, white scales. Epistoma not to very slightly transversely swollen at base. Pronotum. Median carina variously
developed in anterior one-half as slightly elevated glabrous narrow to broad line. Dorsal punctation small, dense, shallow;
smaller and sparser apically; sparser on flanks; areas between dorsal punctures with dense minute regularly impressed
punctures. Dorsally with moderately dense, short erect hair-like scales each situated in larger puncture (Fig. 138a). Scales
dorsally absent to sparse, small and fine along midline; moderately dense to dense, elongate-fine, appressed laterad of
median line, but not as distinct stripes; laterally, absent to sparse, small and fine. Lateral margins with scales moderately
Quaest. Ent., 1987,23 (4)
504
Anderson
dense to dense, elongate-narrow to moderately robust, appressed to recumbent in moderately broad stripe. Flanks with
scales moderately dense to dense, elongate-fine. Median basal area of disk very shallowly, broadly impressed.
Anterolateral margin with pronotal postocular lobes well-developed; postocular vibrissae uniformly short (Fig. 138a). In
dorsal view with lateral margins uniformly moderately arcuate and convergent from base to apex; subapical constriction at
apical one-fifth indistinct or slight, then straight and convergent to apex (Fig. 119). Prosternum. With shallow to
moderately deep impression and moderately to well-developed rounded swelling anterior to each procoxal cavity. In lateral
view with anterior flat to slightly angulate portion moderately long, approximately one-half length of posterior more
angulate portion; anterior and posterior portions not or slightly differentiated by shallow transverse impression. Elytra.
Elongate-narrow in general form (width at midlength 0.49-0.58 times length in males; 0.49-0.55 in females) (Fig. 119). In
dorsal view with lateral margins very slightly sinuate, slightly convergent from apical one-third to humerus; slightly to
moderately and evenly arcuate from apical one-third to apex. Elytral apices slightly produced, rounded to sub-acuminate.
Humerus distinct. Dorsally with sparse to moderately dense, suberect to erect, very short to short, hair-like scales. Sutural
interval to interval 7 with uniformly moderately dense, fine to elongate-fine, white scales and with scattered patches of
larger white scates; interval 8 with scales absent to moderately dense, small and fine basally to elongate-fine apically;
intervals 9 to 1 1 with scales moderately dense to dense, moderately large and elongate-narrow to robust, smaller and finer
apically in most specimens (Fig. 119). Punctures of elytral striae small, individually distinct, arranged in regular rows.
Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with small denticles in apical
three-quarters; subapical tooth not distinct from tibial denticles. Foretibial and mesotibial unci of both sexes moderately
large, metatibial uncus small. Tarsal claws connate in basal one-half, not to slightly divergent, each with basal internal
flange well-developed. Ventral tarsal pilose vestiture of all tarsi of male and female present as elongate-narrow pads on
apical one-half to three-quarters of article 1, as rounded pads on apical one-half of article 2, and as large rounded pads on
apical two-thirds to more or less entire ventral surface of article 3. Abdomen. Ventral surface with moderately dense to
dense elongate-narrow appressed white scales. Abdominal sterna III to VI (especially III and IV) with individually
indistinct glabrous patches, each with large puncture and single suberect to erect moderately long hair-like scale situated
in or near center. Base of abdominal sternum VII of female internally with slightly developed median shiny tubercle.
Genitalia. Female (two examined). Abdominal sternum VIII with basal arm short; lateral arms straight and divergent
from base to apical one-third; markedly abruptly inwardly arcuate at apical one-third; apical one-half to one-third
markedly expanded (Fig. 158). Stylus moderately large in length compared to length of gonocoxite II. Male (two
examined). Internal sac (Figs. 178a, b) elongate and low; median dorsal pocket low, with single elongate moderately broad
dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B),
with moderately large paired dorsally directed lobe near midlength on dorsal surface (lobe C), with moderately large
paired dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired anterodorsolaterally
directed lobe at apical one-third at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near
ventral margin (lobe F). Apex of median dorsal pocket rounded in dorsal view. Apical pocket individually distinct from
median dorsal pocket. Ventral median pocket moderately large. Apical pocket with paired moderately large ventrally
directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like
sclerite; adjacent ventral surface of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is known only from California (Fig. 216).
Natural history. — This species is found only in the Pacific semi-desert region. Adults of C.
eustictorrhinus have been collected only on Arctostaphylos sp. (Ericaceae); Eriogonum sp.
(Polygonaceae); and beans (Leguminosae). Definite host records are not known. Adults have
been collected from March to July, and December at altitudes of 9-2002 m (N = 8) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister species of C. erysimi (Figs. 236 and
237).
Cleonidius pleuralis (LeConte)
(Figs. 120, 139, 159, 179,217)
Lixus pleuralis LeConte 1858:78. Holotype (examined), female, labelled with a gold circle ( = California), “1062”,
“Type/ 5177”, “Z../ pleuralis/ Lee.” (MCZC). Type locality, on Colorado River below the Gila River, California.
Gemminger and von Harold 1871 (catal.). LeConte 1876a,b (key, redesc.). Henshaw 1881-1882 (check.). Henshaw
1885 (check.). Wickham 1889. Horn 1894 (check.).
Lixus californicus ; Gemminger and von Harold 1871 (catal.). LeConte 1876a.
Lixus modestus\ Gemminger and von Harold 1871 (catal.). LeConte 1876a,b. Henshaw 1885 (check.).
Cleonus ( Cleonidius ) californicus ; Casey 1891 (as synonym).
Cleonus ( Cleonidius ) modestus ; Casey 1891 (key). Wickham 1896 (check.). Fall 1901 (check.). Leng 1920 (catal.). Csiki
1934 (catal.). Blackwelder 1947 (check.). Hatch 1971 (key, misident. of Cleonidius longinasus).
New World Cleonini
505
Cleonus ( Cleonidius ) pleuralis\ Casey 1891 (as synonym). Fall 1901 (check.).
Cleonus modestus-, Wickham 1896 (check.). Fall 1901 (check.). Leng 1920 (catal.). Blackwelder 1947 (check.).
Cleonis californicus-, O’Brien and Wibmer 1982 (catal., distn., as synonym).
Cleonis modestus ; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis pleuralis-, O’Brien and Wibmer 1982 (catal., distn., as synonym).
Notes about synonymy. — This species has been referred to as Lixus modestus Mannerheim
(1843:291) and L. californicus Motschulsky (1845:378). The holotype of Lixus modestus has
been examined (ZMMU) and as noted elsewhere {“Nomen dubium ” section) is not a
Cleonidius and is doubtfully a New World Lixus. The name Lixus californicus Motschulsky is
herein considered a nomen dubium and not considered in the nomenclature (see “ Nomen
dubium ” section) of any species of Cleonidius.
Problems in recognition. — Adults of this species are very easily recognized by the deep
straight sulcus above each eye (Fig. 139) and the produced and acuminate elytral apices (Fig.
120). Most individuals of C. poricollis and some C. calandroides possess a shallow curved
sulcus behind the eye (covered by the pronotal postocular lobe in some specimens and difficult
to see) (Fig. 150a), but this should not be confused with the deep straight sulcus in members of
C. pleuralis.
Description. — Specimens examined. 89 males, 90 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 17. Size. Length, male, 7.2-10.0 mm; female, 7.3-10.8 mm. Width, male, 2. 1-3.1 mm; female, 2. 2-3. 5 mm.
Head. Eye elongate-oval. Vertex, above eye with deep straight sulcus extended posteriorly from dorsal margin of eye to
area under pronotal postocular lobe. Frons with scattered small shallow punctures and moderately dense minute punctures;
also with moderately dense erect hair-like scales above eyes and moderately dense elongate fine appressed white scales,
slightly sparser medially in most specimens. Frons continuous with base of rostrum, not separated by transverse impression
(Fig. 139a). Most specimens with a shallow median fovea at base of median tumescence or carina of rostrum. Width of
frons greater than or subequal to width at apex of rostrum. Rostrum. Elongate-narrow, slightly more so in females (width
at apex 0.44-0.57 times length) (Fig. 139). In lateral view straight to very slightly curved downward. In dorsal view
slightly narrowed apically, postgenae not laterally expanded, antennal scrobes not distinctly visible. Slightly medially
tumescent with median carina in most specimens very low, glabrous shiny line at crest of median tumescence. Dorsal and
lateral punctation sparse, small and shallow, interspersed with moderately dense minute punctures. Dorsally (especially
laterally) with moderately dense erect fine hair-like scales and with moderately dense fine elongate white appressed scales
smaller, finer and sparser apicad of point of antennal insertion. Epistoma not swollen. Pronotum. Median carina lacking in
most specimens, indicated in very few specimens by low, glabrous shiny line in anterior one-half. Dorsal and lateral
punctation moderately large, moderately dense, deep; smaller, sparser and shallower apically and medially, the areas
between large dorsal punctures with minute, very shallow punctures. Dorsally with moderately dense, short to moderately
long, fine erect hair-like scales each situated in large puncture (Fig. 139a). Scales dorsally very sparse to absent along
median line, moderately dense, moderately large and elongate in pair of broad paramedian apically slightly narrowed
stripes; laterally, small and fine. Lateral margins with appressed white scales large and elongate-narrow in moderately
broad stripe. Flanks with scales moderately dense, small and fine. Median basal area of disk deeply and broadly impressed.
Anterolateral margin with pronotal postocular lobes well-developed; postocular vibrissae very short, of uniform length
(Fig. 139a). In dorsal view with lateral margins slightly arcuate and slightly convergent apically from base to more or less
apical one-fifth; gradually to abruptly constricted at apical one-fifth, convergent to apex (Fig. 120). Prosternum. With
moderately deep impression, and moderately to well-developed rounded swelling anterior to each procoxal cavity. In lateral
view with anterior flat to slightly angulate portion short, not distinctly differentiated from longer posterior more angulate
portion. Elytra. Very elongate-narrow in general form (width at midlength 0.40-0.46 times length in males; 0.39-0.47 in
females) (Fig. 120). In dorsal view with lateral margins straight, subparallel to slightly divergent from apical one-third to
humerus; slightly and evenly arcuate from apical one-third to apex. Elytral apices produced, acuminate and divergent.
Humerus distinct. Dorsally with moderately dense, erect, short to moderately long, fine hair-like scales. Scale pattern
various; sutural interval to interval 8 with uniformly moderately dense, elongate fine appressed white scales to moderately
dense, elongate fine on intervals 2 to 5 and moderately dense but smaller and finer on sutural interval and intervals 6 to 8
(interval 8 with scales very sparse to lacking in some specimens); intervals 9 to 1 1 with scales more or less uniformly
moderately dense, moderately large and elongate-narrow (Fig. 120). Punctures of elytral striae small, individually distinct,
arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with
small denticles in apical three-quarters; subapical tooth small, indistinct from tibial denticles. Foretibial uncus of both
sexes moderately large, mesotibial and metatibial unci small. Tarsal claws connate in basal one-third to one-half, not
divergent, basal internal flange lacking. Ventral tarsal pilose vestiture of all tarsi of male and female present as
elongate-narrow pads on apical one-half of article 1, as rounded pads on apical one-half of article 2, and as large rounded
pads on apical three-quarters of article 3. Abdomen. Ventral surface with moderately dense to dense appressed very
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Anderson
elongate-narrow white scales, sparser along midline. Abdominal sterna III to VI (especially III and IV) with moderately
large individually distinct to indistinct glabrous patches, each with large puncture and single elongate appressed to
suberect hair-like scale in or near center. Base of abdominal sternum VII of female internally (dorsally) with slightly
developed, median, shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm very
short, indistinct; lateral arms straight and divergent from base to apical one-quarter, markedly inwardly arcuate at apical
one-quarter and convergent to apex, slightly expanded at apex (Fig. 159). Stylus moderately large in length compared to
length of gonocoxite II. Male (three examined). Internal sac (Figs. 179a,b) elongate and low; median dorsal pocket low,
with single short broad elongate dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at
midlength on dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal one-third on dorsal
surface (lobe C), with moderately large paired dorsally directed lobe at midlength near dorsal margin (lobe D), with
moderately large paired dorsolaterally directed lobe at apical one-third at dorsal margin (lobe E), and with small ventrally
directed paired lobe near base near ventral margin (lobe F). Apex of median dorsal pocket truncate in dorsal view. Apical
pocket individually distinct from median dorsal pocket. Ventral median pocket moderately large. Apical pocket with
paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex
with paired simple scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large, dark, longitudinal
sclerites.
Geographic distribution. — Members of this species are found in the southwestern United
States of America and adjacent northern Mexico from California, southern Nevada, southern
Utah, central New Mexico and extreme western Texas, south to the Mexican border, except in
the west where they are found south into Baja California Norte, Mexico (Fig. 217).
Natural history. — This species is associated with desert-grassland transitional habitat, and
Chihuahuan, Mojave and Sonoran deserts. Adults of this species have been collected in various
sandy habitats such as dunes and dry stream beds or desert washes on Ephedra californica
Wats. (Ephedraceae); Chrysothamnus nauseosus (Pall.) Britton, C. paniculatus (Gray) H.M.
Hall, Hymenoclea sp., H. salsola Torr. and Gray, (all Compositae); Asclepias sp.
(Asclepiadaceae); Atriplex polycarpa (Torr.) Wats. (Chenopodiaceae); Medicago sativa L.
(Leguminosae); and Brassical sp., Lepidium sp. (Cruciferae). Definite hosts are not known.
Adults have been collected from February to October at altitudes ranging from -96-1747 m
(N = 26) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is a member of an unresolved trichotomy that
also includes the C. erysimi-C. eustictorrhinus lineage and the C. subcylindricus-C.
longinasus-C. texanus lineage (Fig. 237).
Cleonidius subcylindricus Casey
(Figs. 121, 140, 160, 180, 215)
Cleonus ( Cleonidius ) subcylindricus Casey 1891:193. Holotype (examined), female, labelled “Fla”, “CASEY/ bequest/
1925”, “TYPE USNM/ 37283”, “C. Cl./ subcylindricus / Cas” (USNM). Type locality, Florida. Csiki 1934 (catal.).
Cleonus ( Cleonidius ) graniferus Casey 1891:194. NEW SYNONYMY Holotype (examined), female, labelled “Ga.”,
“CASEY/ bequest/ 1925”, “TYPE USNM/ 37284”, “C. Cl./ graniferus/ Cas.” (USNM). Type locality, Georgia.
Csiki 1934 (catal.).
Lixus lupinus Blatchley 1914:248. Lectotype (designated by Blatchley [1930:38, error as female] examined), male,
labelled “TYPE”, “Dunedin, Fla./ W.S.B. Coll./ 1.24.1913”, “Purdue/ Blatchley/ collection” and with lectotype
designation label of Blatchley (Purdue). Type locality, Dunedin, Florida. Blatchley and Leng 1916 (key, redesc.).
Leng 1920 (catal). Blatchley 1930.
Cleonus subcylindricus', Blatchley and Leng 1916 (key, redesc.). Leng 1920 (catal.).
Cleonus graniferus-, Blatchley and Leng 1916 (key, redesc.). Leng 1920 (catal.).
Cleonus lupinus-, Kingsolver 1972 (synonymy with Cleonus subcylindricus).
Cleonis graniferus ; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis lupinus-, O’Brien and Wibmer 1982 (catal., distn., as synonym).
Cleonis subcylindricus ; O’Brien and Wibmer 1982 (catal., distn.).
lupinus, incertae sedis-, Csiki 1934 (catal.).
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507
Notes about synonymy. — Cleonus graniferus Casey is a developmental^ abnormal female
of C. subcylindricus with a short robust rostrum, legs and antennae. Similar developmental
abnormalities are also known in other species of Cleonidius. Lixus lupinus Blatchley was
placed in synonymy with Cleonus subcylindricus Casey by Kingsolver (1972).
Problems in recognition. — This species is one of only two species of Cleonidius found in and
restricted to eastern North America (Figs. 215, 229). These two species differ markedly in
many structural features and there should be no problem in separating individuals of the two.
C. subcylindricus individuals are easily distinguished from other members of the C. texanus
group, with which they may be confused, by the only slightly developed pronotal postocular
lobes and by lack of dorsal suberect or erect vestiture (Fig. 140a).
Problems in separation of C. subcylindricus from eastern species of Lixus are resolved by
Lixus species, among other characters, lacking broadly rounded pronotal postocular lobes and
instead possessing at most a small acute postocular projection and postocular vibrissae that are
not more or less uniform in length, but rather variable in length and distinctly longest at some
point behind the eye (Fig. 4).
Description. — Specimens examined. 11 males, 18 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 18. Size. Length, male, 8.9-10.5 mm; female, 8.5-11.3 mm. Width, male, 2. 9-3. 5 mm; female, 2. 8-3. 9 mm.
Head. Eye elongate-oval. Area behind eye with numerous wrinkles of cuticle. Frons with very dense, small, shallow
punctures, some confluent in some specimens. Frons lacking suberect or erect vestiture, with moderately dense, small to
moderately large, very fine and elongate appressed white scales (scales abraded medially in most specimens). Frons
separated from base of rostrum by shallow transverse impression (Fig. 140a). Base of rostral carina with small shallow
fovea. Width of frons less than width at apex of rostrum. Rostrum. Moderately robust, very slightly more so in males
(width at apex 0.56-0.69 times length) (Fig. 140). In lateral view slightly curved downward. In dorsal view with postgenae
slightly expanded laterally, antennal scrobes partially visible. Slightly medially tumescent from point of antennal insertion
to base of frons, with distinct, low, glabrous median carina from point of antennal insertion to middle of frons. Dorsal and
lateral punctation very dense, small, shallow (many punctures longitudinally confluent in some specimens). Dorsally
lacking suberect or erect vestiture; with moderately dense elongate-fine appressed white scales, scales smaller medially.
Epistoma slightly elevated and swollen. Pronotum. Median carina lacking to variously developed as low, glabrous, shiny
line. Dorsal punctation small, dense and shallow; larger and sparser on flanks; areas between larger dorsal punctures with
very dense shallow minute punctures, areas between shiny and irregularly elevated. Dorsally lacking suberect or erect
vestiture (Fig. 140a). Scales dorsally absent to moderately dense and very small along midline; moderately dense, small
and fine, appressed in pair of broad paramedian apically narrowed stripes; laterally absent to moderately dense and very
small. Lateral margins with appressed white scales large and robust, moderately dense in a moderately broad stripe.
Flanks with scales small and fine. Median basal area of disk very shallowly and very broadly impressed. Anterolateral
margin with rounded pronotal postocular lobes moderately developed; postocular vibrissae of approximately uniform
length (slightly longer behind basal portion of eye in some specimens with less well-developed postocular lobes). In dorsal
view with lateral margins straight, slightly convergent from base to apical one-quarter, then straight and more markedly
convergent to apex; subapical constriction indistinct (Fig. 121). Prosternum. With shallow to moderately deep transverse
impression and with prosternal swelling lacking to slightly developed anterior to each procoxal cavity. In lateral view with
anterior flat to slightly angulate portion short, not distinctly differentiated from posterior longer more angulate portion.
Elytra. Elongate-narrow in general form (width at midlength 0.45-0.50 times length in males; 0.43-0.53 in females) (Fig.
121). In dorsal view with lateral margins subparallel from apical one-third to humerus; moderately evenly arcuate from
apical one-third to apex. Elytral apices not produced, sub-acuminate, not divergent. Humerus distinct. Dorsally lacking
suberect or erect vestiture. Sutural interval to interval 8 with scales uniformly moderately dense, fine and elongate, white,
with scattered patches of slightly larger white scales; intervals 9 to 11 with scales uniformly moderately dense, large,
robust and white (Fig. 121). Punctures of elytral striae small, individually distinct, larger and less individually distinct
towards base, arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner
margin with small denticles in apical three-quarters; subapical tooth small, indistinct from tibial denticles. Foretibial,
mesotibial, and metatibial unci of both sexes moderately large. Tarsal claws connate in basal one-half, not divergent, each
with basal internal flange well-developed. Ventral tarsal pilose vestiture of all tarsi of male and female extensive, present
as rounded pads which cover more or less entire ventral surface of all articles, with exception of basal portion of article 1 .
Abdomen. Ventral surface with moderately dense appressed very elongate white scales. Abdominal sterna III and IV
(especially laterally), with distinct moderately large glabrous patches, each with large puncture and single elongate
appressed scale situated in or near center; lacking suberect or erect vestiture except for small, scattered, sparse hair-like
scales on sternum VII of male. Base of abdominal sternum VII of female internally (dorsally) with slightly developed
median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm very short; lateral
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arms straight to very slightly sinuate and divergent from base to apex, apices not expanded (Fig. 160). Stylus large in
length compared to length of gonocoxite II. Male (two examined). Internal sac (Figs. 180a,b) elongate, moderately high;
median dorsal pocket moderately high, with single narrow moderately elongate dorsally directed basal lobe (lobe A), with
moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), and with small ventrally
directed paired lobe near base near ventral margin (lobe F); lobes B, D, and E lacking. Apex of median dorsal pocket
rounded in dorsal view. Apical pocket individually distinct from median dorsal pocket. Ventral median pocket moderately
large. Apical pocket with paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible
apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large,
dark, longitudinal sclerites.
Geographic distribution. — This species is distributed, perhaps discontinuously, along the
Atlantic Coastal Plain from New York south to Florida (Fig. 215).
Natural history. — This species is found at various localities on the Atlantic Coastal Plain in
the southeastern mesophytic evergreen forest region. This species was described from several
specimens collected on flowers of the hoary lupine, Lupinus diffusus Nutt. (Leguminosae) near
Dunedin, Florida between January 24 and March 18 (Blatchley 1914; Blatchley and Leng
1916). Other adults have been collected in roots of Lupinus sp. (Wilmington, North Carolina)
and have been reared from Lupinus cumulicola Small (Spring Valley, Florida). Adult
specimens have been collected from January to May at elevations of sea level to 182 m (N = 7)
(Fig. 233).
Chorological relationships. — Table V. This species is sympatric with C. calandroides in
the northern part of its range in New York.
Phylogenetic relationships. — This species is the sister species of the C. longinasus-C.
texanus lineage (Fig. 237).
Cleonidius longinasus Anderson, new name
(Figs. 122, 141, 161, 181,214)
Lixus mixtus LeConte 1 876a:4 1 6 [not Fabricius 1792:417], Holotype (examined), female, labelled “Col”, “B.D. Smith”,
“Type/ 5178”, “L. mixtus/ Lee.” (MCZC). Type locality, Colorado. Henshaw 1881-1882 (check.). Henshaw 1885
(check.) Leng 1920 (catal.).
Cleonus ( Cleonidius ) modestus ; Hatch 1971 (key, redesc., misident.).
Cleonus mixtus-, Kingsolver 1972.
Cleonis mixtus-, O’Brien and Wibmer 1982 (catal., distn.).
mixtus, incertae sedis\ Csiki 1934 (catal.).
Notes about synonymy. — The name Lixus mixtus LeConte is a junior homonym and is
here replaced with the name C. longinasus Anderson.
Derivation of specific epithet. — From the Latin “ longus ” meaning long and “ nasus ”
meaning nose. This name is used in reference to the very long rostrum of females of this species
(Fig. 141a).
Problems in recognition. — Most individuals, especially females, of this species are easily
recognized by the long and narrow rostrum (Fig. 141a). Otherwise, individuals, especially
males, are likely only to be confused with C. eustictorrhinus, C. subcylindricus and C. texanus.
Individuals of C. longinasus are separated from C. subcylindricus by the latter entirely lacking
suberect or erect dorsal vestiture and being distributed only along the Atlantic Coastal Plain
from New York south to Florida. C. longinasus and C. eustictorrhinus are sympatric in
California but individuals are easily distinguished by the more robust, medially tumescent
rostrum of the latter (Fig. 138). On the other hand, although most C. texanus and C.
longinasus, where they are sympatric in California and Oregon, can be separated on the basis
of the length and width of the rostrum, length of erect dorsal vestiture, and association with
different plant taxa, some specimens, especially males, are very difficult to reliably separate as
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509
discussed under the “Problems in recognition” section for C. texanus.
A very few individuals of C. longinasus have scales on elytral intervals 3 to 5 larger than
those on adjacent intervals giving the elytra an indistinctly vittate scale pattern and perhaps
resulting in confusion with species with distinctly vittate scale patterns. Specimens from San
Francisco, California entirely lack suberect or erect vestiture but have the long narrow rostrum
typical of members of this species.
Description. — Specimens examined. 44 males, 80 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 19. Size. Length, male, 7. 2-9. 8 mm; female, 6.1-11.2 mm. Width, male, 2. 2-3. 4 mm; female, 2. 1-3-8 mm.
Head. Eye elongate-oval. Area behind eye with numerous wrinkles of cuticle. Frons with sparse to moderately dense,
small, shallow punctures, some longitudinally confluent and irregularly impressed immediately above eyes, interspersed
with minute punctures. Frons also with sparse to moderately dense, elongate-fine to elongate-narrow, white, appressed to
suberect scales medially, elongate-narrow and erect immediately above eyes. Some specimens also with moderately dense
very elongate-fine erect hair-like scales medially and immediately above eyes. Frons continuous with base of rostrum or
only very slightly separated by very shallow indistinct transverse impression (Fig. 141a). Base of median carina of rostrum
(if present) with small, shallow fovea. Width of frons greater than width at apex of rostrum. Rostrum. Very
elongate-narrow, especially so in females (width at apex in male 0.40-0.56 times length; 0.34-0.44 in female) (Fig. 141). In
lateral view straight to moderately curved downward (markedly so near apex in some specimens). In dorsal view with
postgenae not laterally expanded, antennal scrobes not or only very slightly visible. Rostrum not medially tumescent, with
median carina lacking to variously developed as rounded slightly elevated fine glabrous shiny line. Dorsal and lateral
punctation dense, small, shallow (longitudinally confluent in many specimens), smaller and not longitudinally confluent
apically, dorsally interspersed with minute punctures. Dorsally with suberect or erect vestiture lacking (few specimens) to
with sparse to moderately dense, moderately long to very long, erect hair-like scales in basal one-half, erect vestiture
lacking from apical one-half; with sparse to moderately dense appressed to suberect elongate-fine scales in basal one-half,
lacking from (perhaps abraded) or very sparse in apical one-half in most specimens. Epistoma not swollen. Pronotum.
Median carina lacking to variously developed in anterior one-half as slightly elevated narrow shiny glabrous line. Dorsal
punctation moderately large, moderately dense to dense, moderately deep; smaller and shallower apically; sparser on
flanks; areas between large dorsal punctures with minute regularly impressed punctures. Dorsally (especially laterally)
with sparse to moderately dense, moderately long to very long, erect hair-like scales, each situated in large puncture (Fig.
141a). Scales dorsally absent to moderately dense, small and fine along median line; sparse to dense, elongate-fine to
elongate-narrow laterad of median line (forming distinct apically narrowed stripes in many specimens); laterally, absent or
sparse to moderately dense, small and fine. Lateral margins with scales moderately dense to dense, elongate-narrow to
moderately robust, appressed to recumbent in moderately broad stripe. Flanks with scales moderately dense to dense,
elongate-narrow. Median basal area of disk broadly and shallowly impressed. Anterolateral margin with pronotal
postocular lobes well-developed; postocular vibrissae uniformly short. In dorsal view with lateral margins uniformly very
slightly arcuate and convergent from base to apex; subapical constriction indistinct. Prosternum. With very shallow,
indistinct impression and very slightly to moderately developed rounded swelling anterior to each procoxal cavity. In
lateral view with anterior flat to slightly angulate portion long, approximately one-half to two-thirds length of posterior
more angulate portion; anterior and posterior portions not to slightly differentiated by shallow transverse impression.
Elytra. Elongate-narrow in general form (width at midlength 0.48-0.54 times length in males; 0.48-0.58 in females) (Fig.
122). In dorsal view with lateral margins straight to very slightly arcuate or slightly sinuate, slightly convergent from
apical one-third to humerus; slightly to moderately and evenly arcuate from apical one-third to apex. Elytral apices slightly
produced, rounded to sub-acuminate. Humerus distinct. Dorsally with sparse to moderately dense, short to very long erect
hair-like scales (longest and densest on declivity of most specimens; absent on specimens from San Francisco, California).
Sutural interval to interval 7 with uniformly moderately dense, fine to elongate-fine scales, scales on intervals 3 to 5 larger
than those on sutural interval and interval 2, and intervals 6 and 7 in some specimens; interval 8 with scales small, very fine
to fine, sparse to moderately dense; intervals 9 to 1 1 with scales moderately dense to dense, elongate-narrow to moderately
robust, sparser and smaller apically in some specimens (Fig. 122). Punctures of elytral striae small, individually distinct,
arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with
small denticles in apical one-half; subapical tooth small, indistinct from tibial denticles. Foretibial and mesotibial unci of
both sexes large to moderately large, metatibial uncus small. Tarsal claws connate in basal one-third to one-half, not
divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of all tarsi of male and female
present as elongate-narrow pads on apical one-half to two-thirds of article 1, as large oval pads on apical three-quarters to
more or less entire ventral surface of article 2, and as large rounded pads on apical three-quarters to more or less entire
ventral surface of article 3. Abdomen. Ventral surface with moderately dense, elongate-narrow appressed white scales.
Abdominal sterna III to VI (especially III and IV) with individually indistinct to distinct glabrous patches, each with large
puncture and single suberect to erect moderately long hair-like scale situated in or near center. Base of abdominal sternum
VII of female internally (dorsally) with well-developed median shiny tubercle. Genitalia. Female (two examined).
Abdominal sternum VIII with basal arm short; lateral arms straight and divergent from base to apical one-quarter,
inwardly arcuate at apical one-quarter and convergent to apex, apices slightly expanded (Fig. 161). Stylus moderately
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Anderson
large compared to length of gonocoxite II. Male (two examined). Internal sac (Figs. 181 a,b) elongate, moderately high;
median dorsal pocket moderately high, with single moderately broad elongate dorsally directed basal lobe (lobe A), with
large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), and with small ventrally directed paired
lobe near base near ventral margin (lobe F); lobes B, D, and E lacking. Apex of median dorsal pocket rounded in dorsal
view. Apical pocket individually distinct from median dorsal pocket. Ventral median pocket moderately large. Apical
pocket with paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible apical
sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large, dark,
longitudinal sclerites.
Geographic variation. — Specimens from San Francisco, California lack dorsal suberect or
erect vestiture but are otherwise typical C. longinasus. Some specimens from California and
Oregon have less elongate-narrow rostra than typical for this species, perhaps due to
hydridization with C. texanus which in these areas also occurs on Leguminosae. It does not
appear to be host plant induced for all C. longinasus are found on Leguminosae.
Geographic distribution. — This species is distributed from southern British Columbia and
Washington, south to Idaho, western Montana and Wyoming in the east, south to southern
California in the west (Fig. 214).
Natural history. — This species is associated with grassland, Pacific semi-desert and Great
Basin desert habitats. Adults of this species have only been found associated with Leguminosae.
They have been collected on Lupinus spp. and Astragalus spp. (both Leguminosae) throughout
California. No definite host records are known but it is likely that various annual legumes serve
as potential host plants throughout the species range. Adults have been collected from February
to December at elevations of from 31-2293 m (N= 14) (Fig. 233). Unlike adults of C. texanus
which are known to occur, albeit rarely, on low altitude Leguminosae, C. longinasus are not
known from Rosaceae, the presumed primary host of the former species.
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister species of C. texanus (Fig. 237).
Cleonidius texanus (LeConte)
(Figs. 123, 142, 162, 182,218)
Lixus texanus LeConte 1876a: 155. Lectotype (here designated), one of two syntypes, male, labelled with a dark red circle
(= Texas), “67”, “TYPE/ 5250”, uL.texanus/ Lee” and with my designation label “ Lixus / texanus LeC./
LECTOTYPE/ desig. Anderson” (MCZC). Type locality, Texas. LeConte 1876a (in part; misident. of Cleonidius
quadrilineatus , [mixed type series]). Henshaw 1881-1882 (check.). Henshaw 1885 (check.).
Cleonus ( Cleonidius ) circumductus Casey 1891:192. NEW SYNONYMY Holotype (examined), male, labelled “Ari.”,
“CASEY/ bequest/ 1925”, “TYPE USNM/ 37282”, “C.C/./ circumductus / Cas.” (USNM). Type locality, Arizona.
Csiki 1934 (catal.).
Cleonus ( Cleonidius ) texanus ; Casey 1891 (key). Csiki 1934 (catal.).
Cleonus circumductus-, Wickham 1902 (check.). Leng 1920 (catal.).
Cleonus texanus-, Wickham 1902 (check.). Leng 1920 (catal.).
Cleonus canescens-, Yothers 1916 (misident.). Essig 1958 (biol., misident.).
Cleonis circumductus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis modestus\ O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Cleonis texanus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — Casey (1891) distinguished C. circumductus Casey from C.
texanus (LeConte) by the longer, denser and more erect dorsal vestiture. These features of the
dorsal vestiture vary throughout the species range and do not warrant separate species status.
(See also sections on “Problems in recognition” and “Variation”.)
Problems in recognition. — Individuals of this species are likely to be confused only with
individuals of C. longinasus , C. eustictorrhinus, and C. subcylindricus. C. subcylindricus
individuals are easily separated as they lack suberect or erect dorsal vestiture and are found
only along the Atlantic Coastal Plain of the eastern United States of America. Separation of C.
New World Cleonini
511
eustictorrhinus and C. texanus individuals is more difficult. Both species occur in California
but can be separated by the more robust and medially tumescent rostrum of C. eustictorrhinus
(Fig. 138) as outlined in the key.
Individuals, especially males, of C. texanus and C. longinasus are perhaps the most difficult
species of Cleonidius to reliably separate. Both species are sympatric over a wide geographic
range but are especially problematical to distinguish in California and Oregon. Most C.
longinasus are slightly larger in size, have long to very long dorsal erect vestiture (Fig. 141a), a
very elongate-narrow rostrum (especially so in females) (Fig. 141a), and are associated with
both low and high altitude Leguminosae. Most C. texanus on the other hand have generally
shorter dorsal erect vestiture (Fig. 142a), a shorter, slightly more robust rostrum (Fig. 142a),
and are associated with Rosaceae or Rhamnaceae. All individuals from Rosaceae or
Rhamnaceae are structurally typical C. texanus, although not all specimens associated with
Leguminosae are C. longinasus. I have seen both male and female specimens with short rostra
and short dorsal erect vestiture, both character states of C. texanus, from low altitude
Leguminosae. I consider these as C. texanus. Various individuals from the area of sympatry,
including some females, but especially males and those individuals lacking data on plant
associations, are less easy to separate based on these structural differences. The rostrum in
these specimens is generally of a length precluding reliable assignment to either species,
however, length of dorsal erect vestiture of many specimens allows for a tentative identification.
As a consequence of this variation, some may consider these twoforms as questionably
warranting separate species status, but the moderately extensive distribution of C. longinasus,
yet the lack of C. longinasus or C. texanus on Leguminosae from areas (particularly Arizona
and Texas) where C. texanus is found on Rosaceae and Rhamnaceae, and the lack of
individuals with structural character states of C. longinasus from Rosaceae or Rhamnaceae yet
presence of individuals with structural features of C. texanus from Leguminosae, argues
against conspecificity (the structural differences perhaps induced by association with different
host plants). I suspect hybridization to be the likely cause of the complexities in variation but
because of the noted assymmetry in plant associations, do not regard it as warranting
conspecificity of the two forms.
Description. — Specimens examined. 156 males, 174 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 20. Size. Length, male, 6. 2-8. 6 mm; female, 5. 3-8. 8 mm. Width, male, 1.9-3. 2 mm; female, 1. 8-3.0 mm. Head.
Eye elongate-oval. Area behind eye with numerous wrinkles of cuticle. Frons with sparse to moderately dense, small,
shallow punctures, some longitudinally confluent and irregularly impressed immediately above eyes, interspersed with
minute punctures. Frons also with moderately dense, elongate-fine, white appressed to suberect scales medially,
elongate-narrow and erect immediately above eyes. Frons continuous with base of rostrum or only very slightly separated
by very shallow indistinct transverse impression (Fig. 142a). Base of median carina of rostrum (if present) with small,
shallow fovea. Width of frons greater than or subequal to width at apex of rostrum. Rostrum. Elongate-narrow, slightly
more so in females (width at apex 0.51-0.68 times length in male; 0.46-0.66 in female) (Fig. 142). In lateral view slightly
curved downward (markedly so near apex in some specimens). In dorsal view with postgenae not to very slightly laterally
expanded, antennal scrobes only very slightly visible. Rostrum not to very slightly medially tumescent, with median carina
lacking. Dorsal and lateral punctation moderately dense to dense, small, shallow (longitudinally confluent in some
specimens), smaller and not longitudinally confluent apically, dorsally interspersed with minute punctures. Dorsally with
suberect or erect vestiture lacking (few specimens) to with sparse, short, sub-erect to erect hair-like scales laterally and
basally, erect vestiture lacking from apical one-half; with sparse to moderately dense recumbent elongate-very-fine to
elongate-fine scales laterally in basal one-half, lacking from (perhaps abraded) or very sparse in apical one-half and
medially in most specimens. Epistoma not swollen. Pronotum. Median carina lacking to variously developed in anterior
one-half as indistinct slightly elevated narrow shiny glabrous line. Dorsal punctation moderately large, moderately dense to
dense, moderately deep; smaller andshallower apically; sparser on flanks; areas between large dorsal punctures with
minute regularly impressed punctures. Dorsally (especially laterally) with sparse to moderately dense, short to moderately
long, erect hair-like scales, each situated in large puncture (Fig. 142a). Scales dorsally sparse to moderately dense, small
and fine along median line; moderately dense, elongate-fine to elongate-narrow laterad of median line; laterally, sparse to
Quaest. Ent., 1987, 23 (4)
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moderately dense, small and fine. Lateral margins with scales moderately dense to dense, elongate-narrow to moderately
robust, appressed to recumbent in moderately broad stripe. Flanks with scales moderately dense, elongate-narrow. Median
basal area of disk broadly and shallowly impressed. Anterolateral margin with pronotal postocular lobes well-developed;
postocular vibrissae uniformly short. In dorsal view with lateral margins uniformly very slightly arcuate and convergent
from base to apex; subapical constriction indistinct, or straight and convergent from base to apical one-quarter, slightly
constricted at apical one-quarter then straight and convergent to apex. Prosternum. With shallow to moderately deep
impression and very slightly to well-developed rounded swelling anterior to each procoxal cavity. In lateral view with
anterior flat to slightly angulate portion long, approximately one-half to two-thirds length of posterior more angulate
portion; anterior and posterior portions not to slightly differentiated by shallow transverse impression. Elytra.
Elongate-narrow in general form (width at midlength 0.45-0.51 times length in males; 0.44-0.51 in females) (Fig. 123). In
dorsal view with lateral margins straight to very slightly arcuate or slightly sinuate, slightly convergent from apical
one-third to humerus; slightly to moderately and evenly arcuate from apical one-third to apex. Elytral apices slightly
produced, rounded to sub-acuminate. Humerus distinct. Dorsally with sparse to moderately dense, short suberect to erect
hair-like scales (longest and densest on declivity of most specimens). Sutural interval to interval 7 with uniformly
moderately dense, fine to elongate-fine scales, interval 8 with scales small, very fine to fine, sparse to moderately dense;
intervals 9 to 1 1 with scales moderately dense to dense, elongate-narrow to moderately robust, sparser and smaller apically
in some specimens (Fig. 123). Punctures of elytral striae small, individually distinct, arranged in regular rows. Wings.
Long (greater than elytra in length). Legs. Foretibia of female with inner margin with small denticles in apical one-half;
subapical tooth small, indistinct from tibial denticles. Foretibial and mesotibial unci of both sexes large to moderately
large, metatibial uncus small. Tarsal claws connate in basal one-half, not divergent, each with basal internal flange
well-developed. Ventral tarsal pilose vestiture of all tarsi of male and female present as elongate-narrow pads on apical
one-half to two-thirds of article 1, as large oval pads on apical three-quarters to more or less entire ventral surface of
article 2, and as large rounded pads on apical three-quarters to more or less entire ventral surface of article 3. Abdomen.
Ventral surface with moderately dense, elongate-narrow appressed white scales. Abdominal sterna III to VI (especially III
and IV) with individually indistinct to distinct glabrous patches, each with large puncture and single suberect to erect
moderately long hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally)
with well-developed median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm
short; lateral arms straight and divergent from base to apical one-third, inwardly arcuate at apical one-third and
convergent to apex, apices slightly expanded (Fig. 162). Stylus moderately large compared to length of gonocoxite II. Male
(four examined). Internal sac (Figs. 182a,b) elongate, moderately high; median dorsal pocket moderately high, with single
moderately broad elongate dorsally directed basal lobe (lobe A), with moderately large paired dorsally directed lobe at
basal one-third on dorsal surface (lobe C), and with small ventrally directed paired lobe near base near ventral margin
(lobe F); lobes B, D, and E lacking. Apex of median dorsal pocket rounded in dorsal view. Apical pocket individually
distinct from median dorsal pocket. Ventral median pocket moderately large. Apical pocket with paired moderately large
ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple
scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic variation.. — None noted, but see “Geographic variation” section for C.
longinasus and “Problems in recognition” section for C. texanus.
Geographic distribution. — This species is distributed from Washington and Idaho, east to
Colorado, south to southeastern Texas in the east, and southern California in the west (Fig.
218).
Natural history. — This species is associated with the western xerophytic evergreen forest
habitat. Adults have been collected primarily on Rosaceae and Rhamnaceae throughout the
species range, but also on Leguminosae, although only in coastal California and Oregon.
Records from Rosaceae and Rhamnaceae are primarily at higher altitudes, regardless of
geographic location (although especially in oak-pinon-juniper woodlands). Records from
Leguminosae are known only from lower altitudes. Adults have been collected on the following
plants; Lupinus sp. (Leguminosae); Ceanothus sp., C. cuneatus (Hook.) Nutt., C. leucodermis
Greene, C. integerrimus Hook, and Arn. (Rhamnaceae); Cercocarpus sp., C. ledifolius Nutt.,
C. montanus Raf., Cowania mexicana D. Don., Prunus amygdalus Batsch. (almond), P.
persica Batsch. (peach), Pyrus communis L. (pear), Purshia tridentata (Pursh.) DC. (all
Rosaceae). Adults have been reared only from Lupinus affinis J.G. Agardh. (Leguminosae) at
Berkeley, California, but I suspect that a variety of Rosaceae and Rhamnaceae serve as hosts at
higher altitudes throughout most of the species range. Adults have been collected from January
to October at altitudes ranging from 15-2998 m (N = 56) (Fig. 233). Other individuals and I
New World Cleonini
513
have collected specimens of C. texanus and C. quadrilineatus together on various Rosaceae and
Rhamnaceae in Arizona and Texas.
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister species of C. longinasus (Fig. 237).
Cleonidius americanus species group
Diagnosis. — Size small to moderate for Cleonidius (Fig. 197). Dorsal erect or suberect
vestiture absent to present, dense, short to very long. Rostrum moderately elongate-narrow
(Fig. 143) to very robust (Figs. 146-147), slightly to markedly medially tumescent, not to
variously carinate; lateral margins rounded to sharp. Pronotum with postocular lobes absent
(Fig. 145) to present and well-developed (Fig. 143); postocular vibrissae uniformly short to
long, of unequal length, and longest behind base of eye. Elytra with scale pattern vittate,
intervals other than 9 to 1 1 with white scales equal in size to those on intervals 9 to 1 1 (Figs.
146,149), or, with scattered patches of large white scales in mottled appearance (Figs.
143-144). Tarsus with ventral pilose vestiture slightly to markedly reduced in extent (on apical
one-half to two-thirds of articles 2 and 3) to absent or nearly so on all articles. Wings present,
length various. Female with abdominal sternum VIII with basal arm short to long; lateral arms
arcuate to more or less straight throughout length. Male with aedeagus with internal sac with
apex of dorsal median pocket rounded; lobes A to F present; dorsal median pocket low in lateral
view.
Phylogenetic relationships. — Monophyly of this species group is based only on distribution
of the apotypic states of extent of ventral tarsal pilosity (homoplasious within Cleonidius in
species of the C. boucardi group). This monophyly is further weakly supported by the tendency
for individuals of some species placed in the C. americanus group to be brachypterous more so
than individuals of species in other groups and to be the only species group whose members may
exhibit a mottled elytral scale pattern. These states however cannot be considered synapotypic
for they are not shared by all members of the species group and independent evolution of the
apotypic state within species of the group is equally parsimonious with consideration of the
apotypic state as a groundplan state for the group with subsequent reversions to the plesiotypic
state having taken place. The C. americanus group is hypothesized to be the sister group to the
C. poricollis group-C. boucardi group lineage, this lineage then sister to the C. erysimi group
(Figs. 236-237).
Cleonidius americanus Csiki
(Figs. 124, 143, 163, 183,223)
Cleonus ( Cleonidius ) sparsus; Casey 1891 (key, misident.).
Cleonus basalis Fall 1897:242 [not Chevrolat 1860:80]. Holotype (examined), male, labelled “S. Clemente/ Id. CAL./
5.29.97”, “TYPE”, “ basalis / Fall”, “M.C.Z./ Type/ 25192”, “H.C. FALL/ COLLECTION” (MCZC). Type
locality, San Clemente Island, California.
Cleonus {Cleonidius) americanus Csiki 1934:64. New name for Cleonus basalis Fall. Blackwelder 1939 (check.).
Cleonis americanus ; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis basalis; O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Problems in recognition. — Unabraded specimens of this species should be easily recognized
because of the long dorsal erect vestiture (Fig. 143a). Only specimens of C. longinasus have
equally long erect dorsal vestiture, but they have a different elytral scale pattern and a more
elongate-narrow rostrum that is neither laterally expanded at midlength nor with the
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Anderson
prementum swollen ventrally, as in members of C. americanus (Fig. 143). Indeed, C.
americanus and C. frontalis are the only species of Cleonidius in which the rostrum, in lateral
view, has the prementum slightly to markedly swollen ventrally, and in dorsal view is variously
expanded laterally at midlength (Figs. 143-144). Abraded specimens throughout the species
range in California might therefore only be otherwise confused with C. frontalis. From C.
frontalis, individuals of C. americanus can be distinguished by their larger, deeper, and denser
dorsal pronotal punctation, absence of large denticles on the inner margin of the fore-tibia of
females (Fig. 143c), and the rostrum straight, not slightly curved downward.
Description. — Specimens examined. 65 males, 69 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 21. Size. Length, male, 4.9-10.9 mm; female, 5.4-11.3 mm. Width, male, 1. 9-4.1 mm; female, 2. 1-4.1 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with sparse to moderately dense,
small, shallow punctures, some longitudinally confluent and irregularly impressed immediately above eyes, interspersed
with minute punctures. Frons also with sparse, elongate-fine, white appressed to recumbent scales medially, moderately
dense, elongate-narrow and erect immediately above eyes; with dense, very long erect hair-like scales laterally, lacking
medially. Frons continuous with base of rostrum or only very slightly separated by shallow transverse impression (Fig.
143a). Base of median carina of rostrum (if present) with small, shallow fovea. Width of frons greater than width at apex
of rostrum. Rostrum. Moderately elongate-narrow (width at apex 0.51-0.63 times length in males; 0.43-0.59 in females)
(Figs. 143a,b). In lateral view straight. In dorsal view with postgenae not to very slightly laterally expanded, antennal
scrobes not distinctly visible. Rostrum not to very slightly medially tumescent; slightly laterally expanded at midlength
(Fig. 143b); lateral margins more or less rounded, not sharp; with median carina variously developed as moderately
elevated broad shiny glabrous line. Dorsal and lateral punctation sparse to moderately dense, small, shallow to deep
(longitudinally confluent in many specimens), dorsally interspersed with minute punctures. Dorsally with moderately
dense to dense, very long, erect hair-like scales (especially laterally); with moderately dense recumbent elongate-fine scales
laterally in basal one-half, lacking from (perhaps abraded) or very sparse in apical one-half and medially in most
specimens. Epistoma not swollen. Rostrum in lateral view with submentum slightly to markedly swollen ventrally (Fig.
143a). Pronotum. Median carina lacking to variously developed in anterior one-half as low, narrow shiny glabrous line.
Dorsal punctation and that of flanks large, dense, deep; smaller, sparser and shallower apically; areas between large dorsal
punctures with minute regularly impressed punctures. Dorsally with moderately dense, very long, erect hair-like scales,
each situated in large puncture (Fig. 143a). Scales dorsally absent to sparse, small and fine along median line; moderately
dense, elongate-fine, appressed to recumbent in pair of paramedian slightly apically narrowed stripes; laterally, absent to
very sparse, small and fine. Lateral margins with scales moderately dense, elongate-narrow, appressed in narrow stripe.
Flanks with scales uniformly moderately dense, elongate-fine. Median basal area of disk broadly and shallowly to
moderately deeply impressed. Anterolateral margin with pronotal postocular lobes moderately-developed; postocular
vibrissae uniformly short to of unequal length, moderately long and with greatest width approximately one-half width of
eye, immediately behind base of eye (Fig. 143a). In dorsal view with lateral margins slightly arcuate and convergent from
base to apical one-third, slightly constricted at apical one-third then straight and convergent to apex. Prosternum. With
small shallow impression and very slightly to well-developed rounded swelling anterior to each procoxal cavity. In lateral
view with anterior flat to slightly angulate portion long, approximately two-thirds length of posterior more angulate
portion; anterior and posterior portions slightly differentiated by shallow transverse impression. Elytra. Elongate-narrow in
general form (width at midlength 0.50-0.60 times length in males; 0.50-0.56 in females) (Fig. 124). In dorsal view with
lateral margins straight, slightly convergent from apical one-third to humerus; moderately and evenly arcuate from apical
one-third to apex. Elytral apices not produced, rounded, not divergent. Humerus distinct. Dorsally with moderately dense,
moderately long erect hair-like scales. Scales various in color from golden to white; scale pattern various from mottled,
with uniformly moderately dense small, fine scales and irregularly distributed patches of denser and larger white scales
(Fig. 124); to vittate, with scales uniformly moderately dense, sutural interval and interval 2 (except at base) with golden
scales elongate-very-fine, stria 1 with irregularly distributed patches of larger elongate-narrow white scales, base of
interval 2 and intervals 3 to 5 with scales elongate-narrow, white, intervals 6 to 1 1 various from with irregularly distributed
patches of denser and larger, elongate-narrow white scales interspersed among elongate-very-fine white or golden scales; to
uniformly covered (except intervals 6 and 7 at basal one-third) with large elongate-narrow white scales. Punctures of
elytral striae small, individually distinct, larger and less individually distinct basally; arranged in regular rows. Wings.
Long (greater than elytra in length). Legs. Foretibia of female with inner margin with small denticles in apical
three-quarters; subapical tooth small, indistinct from tibial denticles. Foretibial and mesotibial unci of both sexes large to
moderately large, metatibial uncus small. Tarsal claws connate in basal one-third, not divergent, each with basal internal
flange slightly developed. Ventral tarsal pilose vestiture of foretarsus of male present as elongate-narrow pads on apical
one-third to one-half of article 1, as elongate-oval pads on apical one-half to two-thirds of article 2, and as large rounded
pads on apical two-thirds to more or less entire ventral surface of article 3; of foretarsus of female, as apical tufts to small
elongate pads on apical one-quarter of article 1, as small rounded pads on apical one-third of article 2, as moderately large
rounded pads on apical two-thirds of article 3; of mesotarsus of male, as elongate-narrow pads on apical one-half of article
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515
1, as elongate-oval pads on apical one-half to two-thirds of article 2, as large rounded pads on apical two-thirds to
three-quarters of article 3; of mesotarsus of female, as apical tufts to small elongate pads on apical one-quarter of article 1,
as small rounded pads on apical one-third of article 2, as moderately large rounded pads on apical one-half to two-thirds of
article 3; of metatarsus of male, as elongate-narrow pads on apical one-third to one-half of article 1, as elongate-oval pads
on apical one-half to two-thirds of article 2, as large rounded pads on apical two-thirds to three-quarters of article 3; of
metatarsus of female, as apical tufts to small elongate pads on apical one-quarter of article 1, as small rounded pads on
apical one-third of article 2, as moderately large rounded pads on apical one-half of article 3. Abdomen. Ventral surface
with moderately dense, elongate-narrow appressed white scales. Abdominal sterna III to VI (especially III and IV) with
individually indistinct to distinct moderately large glabrous patches, each with large puncture and single erect very long
hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with only very
slightly developed median shiny tubercle. Genitalia. Female (three examined). Abdominal sternum VIII with basal arm
short; lateral arms moderately and evenly inwardly arcuate at point slightly beyond midlength, apices slightly expanded
(Fig. 163). Stylus moderately large compared to length of gonocoxite II. Male (three examined). Internal sac (Figs.
183a,b) elongate and low; median dorsal pocket low, with single elongate narrow dorsally directed basal lobe (lobe A),
with small median dorsally directed lobe at midlength on dorsal surface (lobe B), with moderately large paired dorsally
directed lobe at basal one-third on dorsal surface (lobe C), with moderately large paired dorsolaterally directed lobe near
midlength near dorsal margin (lobe D), with large paired dorsolaterally directed lobe just beyond midlength at dorsal
margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin (lobe F). Apex of median
dorsal pocket truncate in dorsal view. Apical pocket individually distinct from median dorsal pocket. Ventral median
pocket moderately large. Apical pocket with paired moderately large ventrally directed lobe near apical one-third at
ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface and basal
portion of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is found only in California, from the San Francisco
Bay region south along the coast and central valleys to the Mexican border (Fig. 223).
Natural history. — This species is found only in the Pacific semi-desert habitat. Adults of
this species have been collected on Artemesia tridentata Nutt., Chrysothamnus nauseosus
(Pall.) Britton, Franseria sp., Gutierrezia sp., and Aplopappus sp. (all Compositae). No
definite hosts are known but it is probable that a variety of Compositae serve as potential hosts.
Adults have been collected from March to November at elevations of from 15-1446 m (N = 7)
(Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. frontalis (Figs. 236 and
237).
Cleonidius frontalis (LeConte)
(Figs. 125, 144, 164, 184, 226)
Cleonus frontalis LeConte 1876a: 150. Lectotype (here designated), female, one of three syntypes, labelled with a pale
green circle ( = Wyoming Territory), “Black Hills/ Hammond”, “TYPE/ 5245”, “C. frontalis/ Lee.” and with my
designation label “ Cleonus / frontalis LeC./ LECTOTYPE/ desig. Anderson” (MCZC). Type locality, Black Hills,
South Dakota. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1889. Wickham 1902 (check.). Fall
and Cockerell 1907 (check.). Blatchley and Leng 1916 (key, redesc.). Leng 1920 (catal.). Bleasdell 1937 (check.).
Cleonus sparsus LeConte 1876a: 152 [not Zoubkoff 1833:334]. NEW SYNONYMY. Holotype (examined), female,
labelled “Col”, “C./ sparsus/ TYPE Lee.”, “Horn Coll/ H8528” and with a red square label (MCZC). Type locality,
Colorado. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1902 (check.). Leng 1920 (catal.). Essig
1958 (biol., in part misident. of Cleonidius poricollis). Arnett et al. 1980 (probable misident. of Cleonidius
poricollis).
Cleonus ( Cleonidius ) frontalis ; Casey 1891 (key). Csiki 1934 (catal.).
Cleonus ( Cleonidius ) sparsus ; Casey 1891 (key, misident. of Cleonidius americanus). Hatch 1971 (key, redesc., in part
misident. of Cleonidius poricollis).
Cleonus ( Cleonidius ) stratus Csiki 1934:65. NEW SYNONYMY New name for Cleonus sparsus LeConte. Blackwelder
1939 (check.).
Cleonis frontalis-, O’Brien and Wibmer 1982 (catal., distn., in part misident. of Cleonidius poricollis and Cleonidius
puberulus).
Cleonis sparsus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis stratus-, O’Brien and Wibmer 1982 (catal., distn.).
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Anderson
Notes about synonymy. — - To initially distinguish C. sparsus and C. frontalis as separate
species, LeConte (1876a) emphasized differences in the extent of ventral pilosity and form of
hind tarsal article 3. Variation in these characters is continuous and known to occur in a
number of other widespread species, thus I consider these two forms as conspecific. Cleonidius
frontalis was artibrarily chosen over C. sparsus as the valid name for this species.
Casey (1891) incorrectly placed C. puberulus as a junior synonym of C. frontalis and
misidentified individuals of C. americanus as C. sparsus in his key. Likewise, other authors
have frequently confused C. frontalis, C. puberulus, C. sparsus, and less frequently also C.
poricollis.
Problems in recognition. — Individuals of this species are very likely to be confused with
those of C. americanus, C. notolomus and perhaps, C. poricollis and C. trivittatus. They are
easily separated from C. americanus by the long to very long erect dorsal vestiture and dense,
large and deep pronotal punctures of the latter. Furthermore, only a few C. frontalis are known
from California (Fig. 226), whereas C. americanus is restricted to that state (Fig. 223).
Separation from C. poricollis is quite simple as members of that species, where the two are
sympatric, possess a curved sulcus behind each eye (Fig. 150a). Similarly, separation from C.
trivittatus is simple, as individuals of that species lack or at mosthave only very slightly
developed postocular lobes (Fig. 153a).
Separation from C. notolomus is most difficult and discussed under “Problems in
recognition” section for that species.
Most individuals of Cleonidius with an elytral scale pattern that is mottled with irregularly
distributed patches of larger white scales, and not distinctly vittate (Fig. 125), are this species.
Description. — Specimens examined. 121 males, 115 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 22. Size. Length, male, 5.7-10.5 mm; female, 5.7-10.8 mm. Width, male, 2.2-3. 9 mm; female, 2.2-4. 1 mm.
Head. Eye elongate-oval (Fig. 145a). Area behind eye with numerous slight wrinkles of cuticle. Frons with sparse to
moderately dense, small, shallow punctures, some longitudinally confluent and irregularly impressed immediately above
eyes, interspersed throughout with minute punctures. Frons also with moderately dense, elongate-narrow, white appressed
to recumbent scales medially, moderately dense, elongate-narrow and suberect to erect immediately above eyes; otherwise
lacking suberect or erect vestiture. Frons slightly separated from base of rostrum by shallow to moderately deep transverse
impression (Fig. 144a). Base of median carina of rostrum (if present) with small, shallow fovea. Width of frons greater
than, to subequal to, width at apex of rostrum. Rostrum. Moderately robust, slightly less so in females (width at apex
0.56-0.71 times length in male; 0.54-0.65 in female ) (Figs. 144a, b). In lateral view straight to very slightly curved
downward. In dorsal view with postgenae not to very slightly laterally expanded, antenna scrobes very slightly visible.
Rostrum slightly to moderately medially tumescent from point of antennal insertion to middle of frons (more markedly so
basally); not distinctly to slightly laterally expanded at midlength; lateral margins more or less rounded, not sharp; with
median carina absent to variously developed as low glabrous shiny line at crest of median tumescence. Dorsal and lateral
punctation sparse to dense, small, shallow (longitudinally confluent in some specimens); smaller and less dense apically;
dorsally interspersed with minute punctures. Dorsally with sparse to moderately dense, short, suberect hair-like scales
(especially laterally); with moderately dense appressed elongate-narrow scales laterally in basal one-half, sparse in apical
one-half and medially in most specimens. Epistoma not swollen. Rostrum in lateral view with submentum slightly swollen
ventrally (Fig. 144a). Pronotum. Median carina lacking to variously developed in anterior one-half as low, rounded,
moderately broad shiny glabrous line. Dorsal punctation moderately large, moderately dense, moderately deep to deep;
smaller, sparser and shallower apically and on flanks; areas between large dorsal punctures with minute regularly
impressed punctures. Dorsally with moderately dense, short to moderately long, suberect to erect hair-like scales, each
situated in large puncture (Fig. 144a). Scales dorsally absent to very sparse, small and fine along median line; moderately
dense, elongate-fine, appressed in pair of narrow paramedian stripes; laterally, absent to very sparse, small and fine.
Lateral margins with scales moderately dense to dense, elongate-fine to elongate-narrow, appressed to recumbent in
moderately broad stripe. Flanks with scales moderately dense, elongate-fine to elongate-narrow. Median basal area of disk
broadly and shallowly to moderately deeply impressed. Anterolateral margin with pronotal postocular lobes moderately to
well-developed; postocular vibrissae uniformly short (Fig. 144a). In dorsal view with lateral margins slightly to moderately
arcuate and slightly convergent from base to apex, or slightly constricted at apical one-third to one-quarter then straight
and convergent to apex. Prosternum. With small shallow to moderately deep impression and slightly to well-developed
rounded swelling anterior to each procoxal cavity. In lateral view with anterior flat to slightly angulate portion long.
New World Cleonini
517
approximately two-thirds length of, to slightly shorter than, length of posterior more angulate portion; anterior and
posterior portions not to very slightly differentiated by shallow transverse impression. Elytra. Elongate-narrow in general
form (width at midlength 0.50-0.56 times length in males; 0.49-0.58 in females) (Fig. 125). In dorsal view with lateral
margins straight to very slightly sinuate, slightly convergent from apical one-third to humerus; moderately and evenly
arcuate from apical one-third to apex. Elytral apices not produced, rounded, not to very slightly divergent. Humerus
distinct. Dorsally with sparse, indistinct, very short suberect hair-like scales. Scales various in color from golden to white:
scale pattern various with sutural interval and interval 2 with golden to white scales sparse to moderately dense, very small
and fine to elongate-fine, intervals 3 to 5 with scales moderately dense, elongate-fine to elongate-narrow, white, intervals 6
to 8 with scales sparse to moderately dense, small and fine to elongate-narrow, white, intervals 9 to 1 1 with scales
moderately dense, elongate-fine to elongate narrow, white; striae of many specimens variously with irregularly distributed
patches of larger white scales giving mottled appearance (Fig. 125). Punctures of elytral striae small, individually distinct;
arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with
very large denticles in apical three-quarters; subapical tooth large, indistinct to distinct from tibial denticles (Fig. 144c).
Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal
one-third, not divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus of male
present as elongate-oval pads on apical one-quarter to one-third of article 1, as rounded pads on apical one-third to
one-half of article 2, and as large rounded pads on apical one-half to two-thirds of article 3; of foretarsus of female, as
apical tufts of article 1, as small rounded pads on apical one-quarter of article 2, as moderately large rounded pads on
apical one-half of article 3; of mesotarsus of male, as elongate-narrow pads on apical one-third of article 1, as elongate-oval
pads on apical one-third to one-half of article 2, as large rounded pads on apical one-half to two-thirds of article 3; of
mesotarsus of female, as apical tufts of article 1, as small elongate-oval pads on apical one-third of article 2, as moderately
large rounded pads on apical one-half of article 3; of metatarsus of male, as elongate-narrow pads on apical one-quarter to
one-half of article 1 , as elongate-narrow pads on apical one-third to one-half of article 2, as elongate-oval to rounded pads
on apical one-half to two-thirds of article 3; of metatarsus of female, as elongate-narrow pads on apical one-quarter to
one-half of article 1, as elongate-oval pads on apical one-third of article 2, as moderately large rounded pads on apical
one-half of article 3. Abdomen. Ventral surface with dense, elongate-narrow to broad appressed white scales, sparser and
smaller medially on abdominal sterna V to VII. Abdominal sterna III to VI (especially III and IV) with individually
indistinct to distinct large rounded glabrous patches, each with large puncture and single suberect to erect moderately long
hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with moderately to
well-developed median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm short;
lateral arms very slightly and evenly inwardly arcuate from base to apical one-quarter, more markedly inwardly arcuate at
apical one-quarter and convergent to apex, apices slightly expanded (Fig. 164). Stylus moderately large compared to
length of gonocoxite II. Male (three examined). Internal sac (Figs. 184a,b) elongate and low; median dorsal pocket low,
with single broad short dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on
dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C),
with moderately large paired dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired
dorsolaterally directed lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe
near base near ventral margin (lobe F). Apex of median dorsal pocket truncate in dorsal view. Apical pocket individually
distinct from mediandorsal pocket. Ventral median pocket moderately large. Apical pocket with paired moderately large
ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple
scythe-like sclerite; adjacent ventral surface and basal portion of sides of apical pocket with pair of large, dark,
longitudinal sclerites.
Geographic distribution. — This species is distributed from southern Alberta east to
southern Manitoba, south to California in the west, east to Nebraska, Kansas and western
Texas (Fig. 226).
Natural history. — This species is found in grassland, desert-grassland transitional habitats,
and Great Basin desert. Very little is known of the natural history of this species despite the
large number of specimens collected and examined. Plant associations are very few, the only
records known are of adults on Atriplex sp., A. canescens (Pursh) Nutt., (Chenopodiaceae);
and Melilotus sp. (Leguminosae). No definite hosts are known. Adults have been collected
from January to November, primarily in grassland habitats, at elevations of from 339-3130 m
(N = 38) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. americanus (Figs. 236
and 237).
Quaest. Ent., 1987,23 (4)
518
Anderson
Cleonidius canescens (LeConte)
(Figs. 126, 145, 165, 185,227)
Cleonus canescens LeConte 1 876a: 151. Lectotype (here designated), male, one of three syntypes, labelled “Col.”, “Type/
5243”, “C. canescens/ Lee.” and with my designation label “ Cleonus / canescens/ Lee. LECTOTYPE/ desig.
Anderson” (MCZC). Type locality, Colorado. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1902
(check.). Fall and Cockerell 1907 (check.). Yothers 1916 (misident. of Cleonidius texanus). Leng 1920 (catal.). Essig
1958 (biol., misident. of Cleonidius texanus ).
Cleonus ( Cleonidius ) canescens-, Casey 1891 (key).
Cleonus ( Cleonidius ) grandirostris Casey 1891:189. Lectotype (here designated), male, one of five syntypes, labelled
“N.M.”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37278”, “C. Cl./ grandirostris/ Cas.” and with my designation
label '‘‘'Cleonidius/ grandirostris / Cas./ LECTOTYPE/ desig. Anderson” (USNM). Type locality. New Mexico.
Csiki 1934 (catal.).
Cleonus grandirostris-, Wickham 1896 (check.). Fall and Cockerell 1907 (check.). Leng 1920 (catal.).
Cleonus ( Cleonidius ) coloradensis Csiki 1934:64. Unjustified replacement name for Cleonus canescens LeConte.
Blackwelder 1939 (check.).
Cleonis canescens-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis coloradensis-, O’Brien and Wibmer 1982 (catal., distn., as synonym).
Cleonis grandirostris-, O’Brien and Wibmer 1982 (catal., distn., as synonym).
Notes about synonymy. — O’Brien and Wibmer (1982) synonymized C. canescens and C.
grandirostris. Cleonus coloradensis is an unjustified replacement name for C. canescens.
Problems in recognition. — This species is easily recognized by the less elongate-oval form of
the eye (width greater than 0.60 times length) (Fig. 145a), markedly medially tumescent and
robust rostrum (Fig. 145a), anterolateral margin of pronotum lacking postocular lobe (Fig.
145a), and short metathoracic wings.
Description. — Specimens examined. 182 males, 216 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WEIM, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 23. Size. Length, male, 6. 8-8. 8 mm; female, 6.9-10.5 mm. Width, male, 2. 5-3. 7 mm; female, 2. 3-4. 5 mm. Head.
Eye more or less oval (width greater than 0.60 times length). Area behind eye with at most only two or three slight
wrinkles of cuticle. Frons with moderately dense, small, shallow punctures, some longitudinally confluent and irregularly
impressed immediately above eyes, interspersed throughout with minute punctures. Frons also with dense,
elongate-narrow, white suberect scales, slightly smaller medially in some specimens, moderately dense, elongate-narrow
and erect immediately above eyes; with moderately dense short suberect hair-like scales. Frons continuous with base of
rostrum (Fig. 145a). Base of median tumescence or carina of rostrum (if present) with small, shallow fovea. Width of frons
greater than, to subequal to, width at apex of rostrum. Rostrum. Robust, (width at apex 0.64-0.84 times length in males;
0.64-0.87 in females) (Fig. 145). In lateral view markedly curved downward. In dorsal view with postgenae moderately
laterally expanded, antennal scrobes distinctly visible. Rostrum markedly medially tumescent from point of antennal
insertion to middle of frons; not laterally expanded at midlength; lateral margins more or less rounded, not sharp; with
median carina absent to variously developed as low glabrous shiny line at crest of median tumescence. Dorsal and lateral
punctation moderately dense, small, shallow to deep; dorsally interspersed with minute punctures. Dorsally with
moderately dense, short, erect hair-like scales (especially laterally); with dense suberect moderately robust scales, smaller
and finer apicad of point of antennal insertion. Epistoma moderately transversely swollen at base. Pronotum. Median
carina lacking to slightly developed in anterior one-half as low, rounded, shiny narrow glabrous line. Dorsal and lateral
punctation small to moderately large, moderately dense, shallow to deep; smaller apically; areas between large dorsal
punctures with minute regularly impressed punctures. Dorsally with moderately dense, very short, fine erect hair-like
scales, each situated in large puncture (Fig. 145a). Scales dorsally either uniformly moderately dense medially on disk, or
absent along median line; moderately dense, moderately robust, appressed in pair of broad apically slightly narrowed
paramedian stripes; laterally, absent to sparse, small and fine. Lateral margins with scales moderately dense, moderately
robust, appressed. Flanks with scales moderately dense in dorsal one-half, slightly larger in ventral one-half, elongate-fine
to elongate-narrow. Median basal area of disk narrowly and moderately deeply impressed. Anterolateral margin with
pronotal postocular lobes lacking; postocular vibrissae of unequal length, long, greatest length (approximately two-thirds
width of eye) immediately behind base of eye (Fig. 145a). In dorsal view with lateral margins straight to slightly arcuate
and slightly convergent from base to apical one-quarter; slightly constricted at apical one-quarter then straight and
convergent to apex. Prosternum. With small shallow impression and at most very slight rounded swelling anterior to each
procoxal cavity. In lateral view with anterior flat to slightly angulate portion lacking or at most very short and indistinct
from posterior more angulate portion. Elytra. Moderately robust in general form (width at midlength 0.56-0.62 times
length in males; 0.54-0.62 in females) (Fig. 126). In dorsal view with lateral margins very slightly arcuate from apical
one-third to humerus; moderately and evenly arcuate from apical one-third to apex. Elytral apices not produced,
subacuminate, not divergent. Humerus rounded, indistinct. Dorsally with moderately dense, short suberect hair-like scales.
New World Cleonini
519
Scale pattern various; either more or less mottled, with scales moderately dense, small, elongate-narrow to moderately
robust on sutural interval and interval 2, moderately dense, large, moderately robust on intervals 3 to 6, moderately dense,
large, but with scattered patches of very much smaller scales on intervals 7 to 11 (Fig. 126); or vittate, with scales
moderately dense, small and fine on sutural interval and interval 2, moderately dense, uniformly large, moderately robust
on intervals 3 to 5, moderately dense, moderately robust, small on intervals 6 and 7, absent to sparse, small, fine on interval
8, uniformly moderately dense, large, moderately robust on intervals 9 to 1 1. Punctures of elytral striae small, individually
distinct; arranged in regular rows. Wings. Short (approximately equal to one-half length of elytra). Legs. Foretibia of
female with inner margin with small denticles in apical two-thirds; subapical tooth small, indistinct from tibial denticles.
Foretibial and mesotibial unci of both sexes large to moderately large, metatibial uncus small. Tarsal claws connate in
basal one-half, not divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus of
male present as elongate-narrow pads on apical one-quarter of article 1, as elongate-oval pads on apical one-quarter to
one-half of article 2, and as large rounded pads on apical two-thirds of article 3; of foretarsus of female, as apical tufts of
article 1, as small rounded pads at ventral apex of article 2, as moderately large rounded pads on apical one-half to
two-thirds of article 3; of mesotarsus of male, as elongate-narrow pads on apical one-quarter of article 1, as elongate-oval
pads on apical one-third of article 2, as large rounded pads on apical one-half to two-thirds of article 3; of mesotarsus of
female, as apical tufts of article 1, as small rounded pads at ventral apex of article 2, as moderately large rounded pads on
apical one-half to two-thirds of article 3; of metatarsus of male, as apical tufts of article 1, as small elongate pads at ventral
apex of article 2, as large rounded pads on apical one-half to two-thirds of article 3; of metatarsus of female, as apical tufts
of article 1, as small elongate pads at ventral apex of article 2, as moderately large elongate pads on apical one-half to
two-thirds of article 3. Abdomen. Ventral surface with moderately dense to dense, elongate-narrow appressed white scales,
sparser and smaller medially on abdominal sterna V to VII. Abdominal sterna III to VI (especially III and IV) with
individually indistinct to distinct small to large rounded glabrous patches, each with large puncture and single suberect
moderately long hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally)
with at most very slightly-developed median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII
with basal arm long; lateral arms markedly inwardly arcuate at midlength, apical one-half of each arm laterally expanded
(Fig. 165). Stylus moderately large compared to length of gonocoxite II. Male (four examined). Internal sac (Figs. 185a,b)
elongate and low; median dorsal pocket low, with single moderately broad elongate dorsally directed basal lobe (lobe A),
with moderately large median dorsally directed lobe at midlength on dorsal surface (lobe B), with moderately large paired
dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired dorsolaterally directed lobe just
beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin
(lobe F); lobe C absent. Apical pocket not distinct from median dorsal pocket. Ventral median pocket moderately large.
Apical pocket with paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible
apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface of apical pocket with pair of large,
dark, longitudinal sclerites.
Geographic variation. — Elytral and abdominal scale patterns and depth and size of head,
rostral and pronotal punctation vary geographically. Specimens from the western part of the
species range in Arizona, Utah and Idaho have the vittate form of elytral scale pattern. The
ventral surface of the abdomen of these individuals has only small indistinct glabrous patches,
and head, rostral and pronotal punctures are small and shallow. Specimens from the rest of the
species range to the east have a mottled form of elytral scale pattern (Fig. 126), with ventral
abdominal glabrous patches large and distinct, and punctation of the head, rostrum and
pronotum slightly larger and deeper.
The rostrum is also more elongate-narrow in the western form than in the eastern form
based on measurements of rostral width at apex compared to length of rostrum in both male
and female individuals from populations at 6 miles SE. Turkey, Texas; 19 miles SW. Kayenta,
Arizona; and Denver, Colorado. Individuals from the population at Denver have the most
robust rostra (males, X = 0.805 N = 8; females, X = 0.824 N = 8), those from 6 mi. SE. Turkey
have an intermediate sized rostrum (males, X = 0.780 N = 8; females, X = 0.751 N = 7), and
those from 19 mi. SW Kayenta have the least robust rostrum (males, X = 0.673 N=12;
females, X = 0.669 N = 6).
Patterns of variation in all characters in this species parallel those in A. angularis wherein
gene flow appears to be occuring across the continental divide in southern New Mexico and
Arizona, but not to the north, in Colorado and Utah.
Geographic distribution. — This species is distributed in the southcentral United States of
America from extreme southeastern Idaho east to Nebraska, south in the west to southern
Quaest. Ent., 1987, 23 (4)
520
Anderson
Arizona, east to extreme southern Texas (Fig. 227). There is a questionable record from Lassen
National Park in northern California.
Natural history. — This appears to be a species of low to high elevation grassland,
desert-grassland transitional, Texas semi-desert, and Great Basin desert habitats. Plant
associations are few. Adults have been collected only on Symphoricarpos sp. (Caprifoliaceae);
Atriplex canescens (Pursh) Nutt. (Chenopodiaceae); Medicago sativa L. (alfalfa)
(Leguminosae); and Mentzelia sp. (Loasaceae). Definite hosts are not known. Specimens have
been collected in all months of the year at elevations of 200-2868 m (N = 56) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister group of the C. notolomus-C.
collaris-C. infrequens-C. puberulus lineage (Fig. 237).
Cleonidius infrequens Anderson, new species
(Figs. 5, 127, 146, 166, 186, 222)
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “Scott City, Kan./5-29-59/H. Willis”,
“HOLOTYPE/ Cleonidius / infrequens/ Anderson” and with abdomen on card and genitalia in microvial attached to pin
(CWOB). Allotype, female, with a red label “ALLOTYPE”, “TEX. Muleshoe/Bailey Co./16.IV.1971/G.B. Marshall,
“ALLOTYPE/ Cleonidius / infrequens/ Anderson” and with abdomen on card and genitalia in microvial attached to pin
(CWOB). Type locality, Scott City, Kansas.
Paratypes. 1 male, 5 females. MEXICO: San Luis Potosi: San Luis Potosi, 17.VE68, Arsego, IF (USNM).
UNITED STATES OF AMERICA: New Mexico: Fort Sumner, 19. VIII. 51, Robinson, 1M (CWOB); 15mi. n. Las
Cruces, 24. VII. 79, Richmond, IF (RSAN). Texas: 4mi. s. Big Springs, 3. VI. 72, C.W. O’Brien, IF (CWOB); D’Hanis,
23.11.63, Fohn, 1F(TAMU); Van Horn, 24.VI.42, Scullen, 1F(0SUC).
Derivation of specific epithet. — From the Latin “/«” meaning not and ‘ frequens ” meaning
frequent. This name is used in reference to the rarity of individuals of this species.
Problems in recognition. — Individuals of this species are likely to be confused only with
those of C. puberulus and perhaps also C. notolomus. From C. notolomus, specimens are easily
distinguished by their possession of a more robust, sharply and distinctly medially carinate
rostrum, and differently impressed pronotum. Characters allowing for separation of C.
infrequens from C. puberulus are discussed under “Problems in recognition” section for that
species.
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 24. Size. Length, male, 8. 3-8.8 mm;
female, 8.2-10.0 mm. Width, male, 3. 1-3.3 mm; female, 3. 0-3. 8 mm. Head. Eye elongate-oval. Area behind eye with
numerous slight wrinkles of cuticle. Frons with sparse, small, shallow punctures, some longitudinally confluent and
irregularly impressed immediately above eyes, interspersed throughout with minute punctures. Frons also with moderately
dense to dense, elongate-fine to elongate-narrow, white appressed scales medially, suberect to erect immediately above
eyes; otherwise lacking suberect or erect vestiture. Frons distinctly separated from base of rostrum by deep transverse
impression (Fig. 146a). Base of median carina of rostrum with small, shallow fovea. Width of frons less than to subequal to
width at apex of rostrum. Rostrum. Robust, (width at apex 0.78-0.79 times length in males; 0.75-0.92 in females) (Fig.
146). In lateral view straight to very slightly curved downward. In dorsal view with postgenae slightly laterally expanded,
antennal scrobes partially visible. Rostrum slightly medially tumescent from point of antennal insertion to middle of frons;
not laterally expanded at midlength; lateral margins sharp, rostrum flat from immediately laterad of median carina to
lateral margins; with median carina present, very distinct, as sharp, elevated, glabrous shiny line at crest of median
tumescence; also with indistinct low lateral carina at lateral margin in most specimens (Fig. 146b). Dorsal and lateral
punctation moderately dense, moderately large to large, moderately deep and longitudinally confluent (especially so
immediately laterad of median carina); dorsally interspersed with sparse minute punctures. Dorsally lacking suberect or
erect vestiture; with sparse to moderately dense elongate-narrow to robust appressed white scales. Epistoma transversely
swollen at base. Pronotum. Median carina lacking to variously developed in anterior one-half as very low, indistinct, broad
glabrous line. Dorsal and lateral punctation small to moderately large, moderately dense to dense, moderately deep, not
confluent; smaller sparser and shallower apically; areas between large dorsal punctures with minute regularly impressed
punctures; punctation sparser and shallower on flanks. Dorsally with moderately dense, short, suberect hair-like scales,
each situated in large puncture (Fig. 146a). Scales dorsally absent to sparse, small and fine along median line: dense,
New World Cleonini
521
elongate-fine to elongate-narrow, appressed in pair of apically slightly narrowed paramedian stripes; laterally, absent to
sparse, small and fine. Lateral margins with scales dense, elongate-narrow, appressed in moderately broad stripe. Flanks
with scales moderately dense to dense, elongate-fine to elongate-narrow. Median basal area of disk broadly and moderately
deeply impressed. Anterolateral margin with pronotal postocular lobes well-developed; postocular vibrissae of unequal
length, short, greatest length immediately behind base of eye (Fig. 146a). In dorsal view with lateral margins straight and
subparallel to slightly and evenly arcuate from base to apical one-third to one-quarter; constricted at apical one-third to
one-quarter then straight and convergent to apex. Prosternum. With small shallow impression and very slightly to
moderately developed rounded swelling anterior to each procoxal cavity. In lateral view with anterior flat to slightly
angulate portion long, approximately one-half to two-thirds length of posterior more angulate portion; posterior and
anterior portions not or only slightly separated by shallow transverse impression. Elytra. Moderately robust in general
form (width at midlength 0.55-0.61 times length in males; 0.52-0.60 in females) (Fig. 127). In dorsal view with lateral
margins very slightly sinuate and convergent from apical one-third to humerus; moderately and evenly arcuate from apical
one-third to apex. Elytral apices not produced, subacuminate, not divergent. Humerus rounded, indistinct. Dorsally with
very sparse, irregularly distributed very short suberect hair-like scales; with sutural interval and interval 2 with very sparse
to moderately dense, very small and fine to elongate-fine white scales; stria 1 with irregularly distributed patches of larger
and denser white scales; intervals 3 to 5 with uniformly dense, elongate-narrow to moderately robust white scales; intervals
6 to 8 with scales absent to very sparse, very small and fine; stria 6 and 7 with irregularly distributed patches of larger and
denser white scales; intervals 9 to 1 1 with scales uniformly dense, white, elongate-narrow to robust (Fig. 127). Punctures of
elytral striae small, individually distinct, regular rows. Wings. Long (greater than length of elytra [62%, N = 5]) to short
(approximately equal to one-half length of elytra or less [38%, N = 3]). Legs. Foretibia of female with inner margin with
small to large denticles in apical three-quarters; distinct subapical tooth present. Foretibial and mesotibial unci of both
sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-half, slightly divergent, each with basal
internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus of both sexes present as elongate pad on apical
one-third of article 1, as large rounded pad on apical one-half of article 2 and apical three quarters of article 3; of
mesotarsus of both sexes as elongate pad on apical one-quarter to one-third of article 1, as large rounded pad on apical
one-half of article 2 and apical three quarters of article 3; and of metatarsus of both sexes as elongate pad on apical
one-quarter to one-third of article 1, as small rounded to elongate pad on apical one-half of article 2 and as elongate-oval
pad on apical one-half to two-thirds of article 3. Abdomen. Ventral surface with moderately dense to dense,
elongate-narrow to moderately robust appressed white scales, sparser and smaller medially on abdominal sternum VII of
female. Abdominal sterna III to VI (especially III and IV) with individually indistinct to distinct moderately large
rounded glabrous patches, each with large puncture and single suberect to erect, short to moderately long hair-like scale
situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with a slightly-developed median
shiny tubercle. Genitalia. Female (five examined). Abdominal sternum VIII with basal arm long; lateral arms very slightly
inwardly arcuate throughout length, expanded abruptly at apex (Fig. 166). Stylus moderately large compared to length of
gonocoxite II. Male (two examined). Internal sac (Figs. 186a,b) elongate and low; median dorsal pocket low, with single
moderately broad elongate dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on
dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C),
with moderately large paired dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired
dorsolaterally directed lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe
near base near ventral margin (lobe F). Apical pocket and median dorsal pocket individually distinct; apex of dorsal
median pocket rounded in dorsal view. Ventral median pocket moderately large. Apical pocket with paired moderately
large ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple
scythe-like sclerite; adjacent ventral surface and basal portion of sides of apical pocket with pair of large, dark,
longitudinal sclerites.
Geographic distribution. — Very few individuals of this species are known from western
Kansas, southern New Mexico, and western and central Texas, south to San Luis Potosi,
Mexico (Fig. 222).
Natural history. — No definite hosts or host plant associations are known for this species.
Adults have been collected in grassland and desert-grassland transitional habitats from
February to August at elevations of 873-2242 m (N= 7) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. puberulus (Figs. 236
and 237).
Quaest. Ent., 1987, 23 (4)
522
Anderson
Cleonidius puberulus (LeConte)
(Figs. 115, 128, 147, 167, 187, 225)
Cleonus puberulus LeConte 1 876a: 151. Lectotype (here designated), female, one of seven syntypes, labelled “N”, “Type/
5242”, “C. puberulus/ Lee.” and with my designation label “ Cleonus / puberulus/ LeC. LECTOTYPE/ desig.
Anderson” (MCZC). Type locality, Nevada. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Leng 1920
(catal., misinterp. as synonym Cleonidius frontalis.
Cleonus ( Cleonidius ) puberulus ; Casey 1891 (misinterp. as synonym Cleonidius frontalis ). Csiki 1934 (catal., misinterp.
as synonym Cleonidius frontalis).
Cleonus ( Cleonidius ) lobigerinus\ Champion 1902-1906 (in part, distn., misident.).
Cleonis puberulus-, O’Brien and Wibmer 1982 (catal., distn., misinterp. as synonym Cleonidius frontalis ).
Notes about synonymy. — The type locality for this species, given as Nevada (LeConte
1876a), lies outside the geographic range as indicated by specimens I have examined. Perhaps
LeConte (1876a) was in error in that the type locality may have been Nebraska, not Nevada
(presumably an interpretation made by LeConte based on the “N” on the specimen labels).
Problems in recognition.— Individuals of this species are likely to be confused only with
those of C. infrequens. Members of both species possess a very broad rostrum not found in other
Cleonidius species (Figs. 146-147). In C. infrequens however, the median rostral carina is
sharply elevated and distinct, the lateral margins of the rostrum are sharp, the dorsal surface of
the rostrum flat from immediately laterad of the median carina to the lateral margins, the
rostral punctures are deep and large, there are very slight low lateral rostral carinae, ventral
pilose tarsal vestiture is moderately extensive, and both macropterous and brachypterous forms
are known. All of these character states differ from those of C. puberulus individuals. Known
geographic distributions of the two species (Figs. 222, 225) are allopatric, but there may be
sympatry in Colorado or Kansas.
Specimens of C. puberulus may also be confused with small C. poricollis, C. notolomus, and
C. frontalis but can be distinguished by their more robust rostrum (Fig. 147) and lack of the
various diagnostic character states of these other species.
Description. — Specimens examined. 10 males, 26 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 25. Size. Length, male, 5. 1-8.3 mm; female, 6. 4-8. 4 mm. Width, male, 2. 1-3.1 mm; female, 2. 6-3. 3 mm. Head.
Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with sparse, small, shallow punctures,
some longitudinally confluent and irregularly impressed immediately above eyes, interspersed throughout with minute
punctures. Frons also with moderately dense to dense, elongate-fine to elongate-narrow, white appressed scales medially,
suberect to erect immediately above eyes; otherwise lacking suberect or erect vestiture. Frons separated from base of
rostrum by shallow to moderately-deep transverse impression (Fig. 147a). Base of median carina of rostrum with small,
shallow fovea. Width of frons greater than to subequal to width at apex of rostrum. Rostrum. Robust, (width at apex
0.71-0.87 times length in males; 0.74-0.82 in females) (Fig. 147). In lateral view very slightly curved downward. In dorsal
view with postgenae slightly laterally expanded, antennal scrobes partially visible. Rostrum slightly medially tumescent
from point of antennal insertion to middle of frons; not laterally expanded at midlength; lateral margins rounded, rostrum
declivous from immediately laterad of median carina to lateral margins; with median carina present as variously developed
slightly elevated, narrow line, more or less obscured in most specimens by overlying scales. Dorsal and lateral punctation
moderately dense, small to moderately large, shallow, not longitudinally confluent; dorsally interspersed with dense minute
punctures. Dorsally lacking suberect or erect vestiture to with sparse short suberect hair-like scales; moderately dense to
dense, elongate-fine to robust appressed white scales. Epistoma transversely swollen at base. Pronotum. Median carina
lacking to indistinctly developed as very low, broad glabrous line. Dorsal and lateral punctation small, moderately dense to
dense, shallow, not confluent; smaller sparser and shallower apically; areas between large dorsal punctures with minute
regularly impressed punctures; punctation sparser and shallower on flanks. Dorsally with sparse to moderately dense, very
short, suberect hair-like scales, each situated in large puncture (Fig. 147a). Scales dorsally absent to sparse, small and fine
along median line; dense, elongate-fine to elongate-narrow, appressed in pair of apically slightly narrowed paramedian
stripes; laterally, absent to sparse, small and fine. Lateral margins with scales dense, elongate-narrow, appressed in
moderately broad stripe. Flanks with scales moderately dense to dense, elongate-fine to elongate-narrow. Median basal
area of disk broadly and moderately deeply impressed. Anterolateral margin with pronotal postocular lobes well-developed;
postocular vibrissae of unequal length, short, greatest length immediately behind base of eye (Fig. 147a). In dorsal view
with lateral margins straight and subparallel to slightly and evenly arcuate from base to apical one-third to one-quarter;
New World Cleonini
523
constricted at apical one-third to one-quarter then straight and convergent to apex. Prosternum. With small shallow
impression and a very slightly developed rounded swelling anterior to each procoxal cavity. In lateral view with anterior
flat to slightly angulate portion long, approximately one-half to two-thirds length of posterior more angulate portion;
posterior and anterior portions separated by shallow to moderately deep transverse impression. Elytra. Moderately robust
in general form (width at midlength 0.53-0.61 times length in males; 0.54-0.60 in females) (Fig. 128). In dorsal view with
lateral margins very slightly sinuate and convergent from apical one-third to humerus; moderately and evenly arcuate from
apical one-third to apex. Elytral apices not produced, rounded to subacuminate, not to very slightly divergent. Humerus
rounded, indistinct. Dorsally with sparse to moderately dense, very short suberect hair-like scales; with sutural interval and
interval 2 with very sparse to moderately dense, very small and fine to elongate-fine white scales; stria 1 with irregularly
distributed patches of larger and denser white scales; intervals 3 to 5 with uniformly dense, elongate-narrow to moderately
robust white scales; intervals 6 to 8 with scales absent to dense, very small and fine to moderately robust; stria 6 and 7 with
irregularly distributed patches of larger and denser white scales in those specimens with scales of intervals 6 to 8 small and
fine; intervals 9 to 1 1 with scales uniformly dense, white, elongate-narrow to robust (Fig. 148). Punctures of elytral striae
small, individually distinct; arranged in regular rows. Wings. Very short (approximately one-quarter to one-third length of
elytra). Legs. Foretibia of female with inner margin with small denticles in apical three-quarters; no distinct subapical
tooth present. Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate
in basal one-quarter to one-third, slightly divergent, each with basal internal flange moderately to well-developed. Ventral
tarsal pilose vestiture of foretarsus of male present as small elongate pad on apical one-quarter to one-third of article 1, as
small rounded pad on apical one-half of article 2, and as large round pad on apical two-thirds of article 3; of female as
apical tuft of article 1, as small rounded pad on apical one-quarter of article 2, and as large rounded pad on apical one-half
of article 3; of mesotarsus of male as apical tuft to small elongate pad on apical one-quarter of article 1, as apical tuft to
smail elongate pad on apical one-third of article 2, and as small rounded pad on apical one-third of article 3; of female as
apical tuft on articles 1 and 2, and as small rounded pad on apical one-third of article 3; of metatarsus of male lacking
from, to as small elongate pad on apical one-quarter of article 1, as apical tuft to small elongate pad on apical one-third of
article 2, and as small elongate pad on apical one-third of article 3; of female lacking from, to as apical tuft of article 1, as
apical tuft of article 2, and a small elongate pad on apical one-quarter of article 3. Abdomen. Ventral surface with
moderately dense to dense, elongate-narrow to moderately robust appressed white scales, sparser and smaller medially on
abdominal sternum VII of female. Abdominal sterna III to VI (especially III and IV) with individually indistinct to
distinct moderately large rounded glabrous patches, each with large puncture and single suberect, short hair-like scale
situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with slightly to moderately
developed median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm moderately
long; lateral arms slightly inwardly arcuate at mid-length, very slightly expanded at apex (Fig. 167). Stylus moderately
large compared to length of gonocoxite II. Male (three examined). Internal sac (Figs. 187a,b) elongate and low; median
dorsal pocket low, with single moderately broad elongate dorsally directed basal lobe (lobe A), with small median dorsally
directed lobe at midlength on dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal
one-third on dorsal surface (lobe C), with moderately large paired dorsolaterally directed lobe near midlength near dorsal
margin (lobe D), with large paired dorsolaterally directed lobe just beyond midlength at dorsal margin (lobe E), and with
small ventrally directed paired lobe near base near ventral margin (lobe F). Apical pocket and median dorsal pocket
individually distinct; apex of dorsal median pocket rounded in dorsal view. Ventral median pocket moderately large. Apical
pocket with paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible apical
sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface of sides of apical pocket with pair of large,
dark, longitudinal sclerites.
Geographic distribution. — This species is known only from extreme southern Alberta and
Manitoba, south to southern Wyoming and Colorado (Fig. 225).
Natural history. — A single adult has been collected on Lesquerella ludoviciana (Nutt.)
Wats. (Cruciferae) in Kansas. Definite hosts are not known. Adults have been collected from
April to October in grassland habitats at elevations of 1369-2607 m (N = 4) (Fig. 233).
Chorological relationships. — Table V. This species is allopatric with its sister-species C.
infrequens.
Phylogenetic relationships. — This species is the sister-species of C. infrequens (Figs. 236
and 237).
Cleonidius collaris (LeConte)
(Figs. 129, 148, 168, 188,223)
Cleonus collaris LeConte 1876a: 149. Holotype (examined), male, labelled “Col”, “ Cleonus / collaris/ TYPE LeC.”,
“Horn Coll/ H 8517” and with red square label (MCZC). Type locality, Colorado. Henshaw 1881-1882 (check.).
Henshaw 1885 (check.). Wickham 1902 (check.). Leng 1920 (catal.).
Quaest. Ent., 1987, 23 (4)
524
Anderson
Cleonus ( Cleonidius ) collaris\ Casey 1891 (key). Csiki 1934 (catal.).
Cleonis collaris\ O’Brien and Wibmer 1982 (catal., distn.).
Problems in recognition. — The markedly irregularly elevated and sculptured pronotum
with only sparse shallow and indistinct large punctures is characteristic of members this species
and should serve for reliable identification. The pronotum of some specimens of C. notolomus is
similarly broadly and deeply medially impressed at the base and has shallow longitudinal
impressions but is otherwise not as irregularly sculptured as that of C. collaris\ it also has
distinct moderately large, moderately deep dorsal punctation.
Description. — Specimens examined. 7 males, 6 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 26. Size. Length, male, 8.4-10.4 mm; female, 9.3-10.9 mm. Width, male, 3. 3-4. 4 mm; female, 4. 0-4. 6 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with only dense minute
punctures. Frons also with dense, moderately robust, pale brownish appressed scales medially, suberect to erect
immediately above eyes; otherwise lacking suberect or erect vestiture. Frons very slightly separated from base of rostrum
by very shallow transverse impression (Fig. 148a). Width of frons greater than width at apex of rostrum. Rostrum.
Moderately robust (width at apex 0.61-0.73 times length in male; 0.61-0.69 in female) (Fig. 148). In lateral view very
slightly curved downward. In dorsal view with postgenae slightly to moderately laterally expanded, antennal scrobes
partially visible. Rostrum slightly medially tumescent from point of antennal insertion to middle of frons (more markedly
so basally); not laterally expanded at midlength; lateral margins sharp; with median carina present as variously developed
slightly elevated, glabrous, shiny narrow line at crest of median tumescence. Dorsal and lateral punctation sparse, small,
shallow, not longitudinally confluent; dorsally interspersed with dense minute punctures. Dorsally lacking suberect or erect
vestiture; with dense moderately robust to robust appressed white to pale brown scales. Epistoma moderately transversely
swollen at base. Pronotum. Extremely irregulary elevated, markedly constricted dorsolaterally at apical one-quarter,
deeply and broadly impressed medially at base and longitudinally along lateral margins. Median carina variously
developed in anterior one-half from absent to as low, rounded moderately broad glabrous line. Dorsal and lateral
punctation and that of flanks small, very sparse, shallow and indistinct; areas between larger dorsal punctures with dense
minute regularly impressed punctures. Dorsolaterally with sparse, short, suberect hair-like scales, each situated in large
puncture; medially lacking suberect or erect vestiture (Fig. 148a). Scales dorsally absent to sparse, moderately robust
along median line; very dense, moderately robust, appressed in pair of broad apically slightly narrowed paramedian stripes;
laterally, absent to sparse, small and elongate-fine. Lateral margins with scales very dense, moderately robust, appressed to
recumbent. Flanks with scales dense to very dense, moderately robust, appressed. Anterolateral margin with pronotal
postocular lobes well-developed; postocular vibrissae of unequal length, moderately long, greatest length (approximately
one-half width of eye) immediately behind base of eye (Fig. 148a). In dorsal view with lateral margins straight, sinuate or
slightly arcuate from base to apical one-quarter; markedly constricted at apical one-quarter then straight and convergent
to apex. Prosternum. With small shallow impression and slightly developed rounded swelling anterior to each procoxal
cavity. In lateral view with anterior flat to slightly angulate portion long, approximately one-half to two-thirds length of
posterior more angulate portion; posterior and anterior portions slightly to markedly separated by shallow to moderately
deep transverse impression. Elytra. Moderately robust in general form (width at midlength 0.59-0.64 times length in
males; 0.60-0.65 in females) (Fig. 129). In dorsal view with lateral margins very slightly and evenly arcuate, sinuate at
basal one-quarter and convergent from apical one-third to humerus; moderately and evenly arcuate from apical one-third
to apex. Elytral apices not produced, rounded, not to very slightly divergent. Humerus distinct. Dorsally lacking suberect
or erect vestiture; with uniformly dense to very dense, moderately robust appressed white to pale brown scales, absent to
sparse and small in scattered patches on striae 1 and 2, and intervals 9 to 1 1 (Fig. 129). Punctures of elytral striae
individually indistinct, obscured by overlying scales. Wings. Short (approximately one-half length of elytra). Legs.
Foretibia of female with inner margin with small denticles in apical one-half; no distinct subapical tooth present. Foretibial
and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-third, not
divergent, each with basal internal flange slightly developed. Ventral tarsal pilose vestiture of foretarsus of male present as
elongate pad on apical one-third of article 1, as large rounded pad on apical one-half of article 2, and as large round pad on
apical three-quarters of article 3; of female as small elongate pad at apex of article 1, as small rounded pad at apex of
article 2, and as large round pad on apical one-half of article 3; of mesotarsus of male as small elongate pad on apical
one-quarter to one-third of article 1, as moderately large elongate pad on apical one-third to one-half of article 2, and as
large elongate-oval pad on apical two-thirds of article 3; of female as small elongate pad at apex on articles 1 and 2, and as
large rounded pad on apical one-half of article 3; of metatarsus of male as small elongate tuft at apex to on apical
one-quarter of article 1, as small elongate pad on apical one-third to one-half of article 2, and as large elongate-oval pad on
apical one-half of article 3; of female lacking from, to as apical tuft of article 1, as apical tuft on article 2, and as
elongate-oval pad on apical one-half of article 3. Abdomen. Ventral surface with moderately dense to dense, moderately
robust appressed white to pale-brown scales, sparser and smaller medially on abdominal sterna V to VII. Abdominal sterna
III to VI (especially III and IV laterally) with individually distinct large rounded glabrous patches, each with large
puncture and single recumbent to suberect, moderately long hair-like scale situated in or near center. Base of abdominal
sternum VII of female internal y (dorsally) with markedly developed median shiny tubercle. Genitalia. Female (two
New World Cleonini
525
examined). Abdominal sternum VIII with basal arm moderately long; lateral arms slightly inwardly arcuate throughout
length, very slightly expanded at apex (Fig. 168). Stylus moderately large compared to length of gonocoxite II. Male (four
examined). Internal sac (Figs. 188a,b) elongate and low; median dorsal pocket low, with single moderately broad elongate
dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B),
with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), with small paired
dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired dorsolaterally directed lobe just
beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin
(lobe F). Apical pocket and median dorsal pocket individually distinct; apex of dorsal median pocket truncate in dorsal
view. Ventral median pocket moderately large. Apical pocket with paired moderately large ventrally directed lobe near
apical one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent
ventral surface of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is known only from Colorado and adjacent southern
Wyoming, western Kansas and northern New Mexico (Fig. 223).
Natural history. — A single adult specimen from Greeley, Colorado was collected on
Lepidium sp. (pepper-grass; Cruciferae). The very few adults known have been collected in
grassland habitats from March to May at elevations of 1697-2608 m (N = 3) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. notolomus (Fig. 237).
Cleonidius notolomus Anderson, new species
(Figs. 130, 149, 169, 189, 225)
Cleonis frontalis ; O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Type Material. — Holotype, male, with a red label “HOLOTYPE”, “Grant Co NM/ X 1932”, “RT Kellogg”,
“HOLOTYPE/ Cleonidius/ notolomus/ Anderson” and with abdomen on card and genitalia in microvial attached to pin
(USNM). Allotype, female, with a red label “ALLOTYPE”, “New Mex: Hidalgo/Co., lMi.W.Rodeo/VII-31-1971, J.
Doyen & Tschinkel”, “J. Doyen Lot/7 1G20”, “ALLOTYPE/ Cleonidius/ notolomus/ Anderson” and with abdomen on
card and genitalia in microvial attached to pin (UCBC, on indefinite loan to CASC). Type locality, Grant Co., New
Mexico.
Paratypes. 15 males, 16 females. MEXICO: Durango: 26mi. w. Durango, 1 3. VII. 75, L.E. Watrous, 1M (CWOB).
Mexico: Chapingo, 5. VII. 55, Gonzalez, IF (USNM).
UNITED STATES OF AMERICA: Arizona: Cochise County, Guadalupe Canyon, 31. VII. 75, S. McCleve, IF
(SMC); 25mi. n. Douglas, 10.VII.74, S. McCleve, 1M (RSAN); Tucson, XII.27, Downe, IF (UAT); Cochise, 5.VIII.54,
F. Werner, IF (UAT); Douglas, 23. VIII. 27, Jones, 1M (LACM); Chiricahua Mountains, Shake Gulch, 8. VI. 68, Menke,
IF (USNM); Sabino Canyon, 2.VII.52, Kelsey, IF (UAT); Aguirre Lake (lOmi. ne. Sasabe), 2.VIII.78, Hetz, 1M
(UAT); Hereford, 3.III.41, Jones, 1M (USNM); Green Valley, VII.78, R. Lenczy, iM (USNM); Santa Cruz County,
Carmen, 3.VIII.64, Eiland, 1M (USNM); 5mi. s. Cochise, 29.V.61, Statham, IF (AMNH); Sierra Vista, 4.VIII.79, A.E.
Lewis, 1M (CWOB). New Mexico: Grant County, 27. IX. 35, Kellogg, IF (CASC); Silver City, VI. 33, Kellogg, 2F
(OSUC); White Sands National Monument, Lake Lucero, 2. VIII. 74, D. Chandler, 1M (CWOB); White Sands National
Monument, R. Lenczy, IF (USNM), VIII. 78, R. Lenczy, 1M (USNM); 5mi. n. Carlsbad, 21.IX.56, MacSwain, IF
(UCBC); Catron County, Quemado, 24.III.64, Hogg, IF (NMSU). Texas: El Paso, VII, H.F. Wickham, 2M, 4F
(USNM); Van Horn, 24.VI.42, Scullen, 1M (OSUC). Utah: Callao, 8. VI. 22, Spalding, 1M (USNM).
Derivation of specific epithet. — From the Greek “ notos ” meaning south and “ lomus ”
meaning border. This name is used in reference to the distribution of individuals of this species
near the southern border of the United States of America.
Problems in recognition. — Adult individuals of this species are most difficult to separate
from those of C. frontalis . Individuals of both sexes of C. frontalis generally have a
proportionately longer pronotum compared to elytra (see Fig. 200), more elongate-narrow
elytra (see Fig. 197), and more rounded lateral margins of the rostrum. Nearly all C. frontalis
females are easily distinguished from C. notolomus females because the former possess very
large denticles along the inner margin of the fore-tibia (Fig. 144c); these denticles are small in
all C. notolomus. Most C. frontalis individuals have the rostrum very slightly laterally swollen
at midlength and have the submentum slightly ventrally swollen (Figs. 144a,b); this is not so
for C. notolomus. All C. frontalis are also macropterous, whereas C. notolomus are either
Quaest. Ent., 1987, 23 (4)
526
Anderson
macropterous or brachypterous. The two species also are largely allopatric although there is a
zone of sympatry in eastern New Mexico and western Texas.
Individuals of C. notolomus may also be confused with those of C. collaris, C. infrequens, C.
puberulus, C. poricollis, and C. trivittatus. Cleonidius infrequens and C. puberulus individuals
are distinguished by their more robust rostra (Figs. 146-147); C. collaris by the much more
irregularly sculptured and shallowly and indistinctly punctate pronotum, and not distinctly
vittate scale pattern; C. trivittatus by the at most only slightly developed postocular lobes (Fig.
153a); C. poricollis by the curved sulcus behind the eye and the more elongate erect dorsal
vestiture (Fig. 150a).
Description. — Data about variation in LR, WF, WRA, LP, WPB, WE1H, WE1M, LEI, LEy, WEy, WEy/LEy,
WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented in Table 27. Size. Length, male, 7.2-10.8 mm;
female, 8.4-11.4 mm. Width, male, 3. 0-4. 7 mm; female, 3. 6-4. 7 mm. Head. Eye elongate-oval. Area behind eye with
numerous slight wrinkles of cuticle. Frons with sparse to moderately dense, small shallow punctures, interspersed with
minute punctures. Frons also with moderately dense, elongate-narrow appressed white scales medially, suberect to erect
immediately above eyes; otherwise lacking suberect or erect vestiture. Frons slightly separated from base of rostrum by
shallow transverse impression (Fig. 149a). Base of median carina of rostrum with small, shallow fovea. Width of frons
greater than width at apex of rostrum. Rostrum. Moderately robust, (width at apex 0.60-0.77 times length in male;
0.59-0.68 in female) (Fig. 149). In lateral view slightly curved downward. In dorsal view with postgenae at most slightly
laterally expanded, antennal scrobes partially visible. Rostrum slightly medially tumescent from point of antennal insertion
to middle of frons; not laterally expanded at midlength; lateral margins sharp; with median carina as narrow, low, sharp,
and distinct shiny line at crest of median tumescence. Dorsal and lateral punctation moderately dense to dense, small to
moderately large, shallow to moderately deep; smaller and less dense apicad of point of antennal insertion; dorsally
interspersed with dense minute punctures. Dorsally (especially laterally) with sparse to moderately dense, very short,
suberect hair-like scales; also with moderately dense, elongate-narrow, appressed white scales. Epistoma slightly swollen.
Pronotum. Median carina variously developed in anterior one-half from absent to as low, rounded moderately broad
glabrous line. Dorsal and lateral punctation small to moderately large, moderately dense, moderately deep, not confluent;
smaller, sparser and shallower apically and on flanks; areas between larger dorsal punctures with minute regularly
impressed punctures. Dorsally with moderately dense, short, suberect hair-like scales, each situated in large puncture (Fig.
149a). Scales dorsally absent along median line; dense, elongate-narrow, appressed in pair of broad apically slightly
narrowed paramedian stripes; laterally, absent to sparse, small and elongate-fine. Lateral margins with scales dense,
elongate-narrow, appressed to recumbent in moderately broad stripe. Flanks with scales elongate-narrow, slightly less
dense than on lateral margins. Median basal portion of disk deeply and broadly impressed; also longitudinally shallowly
impressed along lateral margins in most specimens. Anterolateral margin with pronotal postocular lobes well-developed;
postocular vibrissae of unequal length, moderately long, greatest length (approximately one-half width of eye)
immediately behind base of eye (Fig. 149a). In dorsal view with lateral margins slightly to moderately arcuate and
convergent to apex; in some specimens, slightly constricted at apical one-quarter then straight and convergent to apex.
Prosternum. With small shallow impression and slightly to moderately developed rounded swelling anterior to each
procoxal cavity. In lateral view with anterior flat to slightly angulate portion long, approximately one-half to subequal in
length to posterior more angulate portion; posterior and anterior portions not or slightly separated by shallow to
moderately deep transverse impression. Elytra. Moderately robust in general form (width at midlength 0.56-0.64 times
length in males; 0.58-0.64 in females) (Fig. 130). In dorsal view with lateral margins straight to very slightly sinuate to
slightly and evenly arcuate and convergent from apical one-third to humerus; moderately and evenly arcuate from apical
one-third to apex. Elytral apices not produced, rounded, very slightly divergent. Humerus distinct to indistinct. Dorsally
with moderately dense, short, suberect to erect hair-like scales; with sutural interval and interval 2 with very sparse to
moderately dense, very small and fine to elongate-fine, golden to white scales; intervals 3 to 5 with uniformly dense,
elongate-narrow to moderately robust white scales; intervals 6 to 8 with scales lacking to very sparse, very small and very
fine, golden to white; intervals 9 to 1 1 with uniformly dense, elongate-narrow to moderately robust white scales; striae 1, 6
and 7 with irregularly distributed patches of larger and denser white scales than on adjacent intervals (Fig. 130).
Punctures of elytral striae small, individually distinct, arranged in regular rows. Wings. Long (greater than elytra in length
[21%, N = 7]), to short (approximately one-half length of elytra or less [79%, N = 26]). Legs. Foretibia of female with
inner margin with small denticles in apical three-quarters; small subapical tooth present, slightly distinct from tibial
denticles. Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in
basal one-third, slightly divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of
foretarsus of male and female present as elongate-narrow pad on apical one-third to one-half of article 1, as moderately
large elongate pad on apical one-third to one-half of article 2, and as large rounded pad on apical two-thirds of article 3; of
mesotarsus of male and female as small elongate pad at apex of article 1, as small rounded pad on apical one-quarter to
one-third of article 2, and as moderately large rounded pad on apical one-third to one-half of article 3; of metatarsus of
male and female as elongate-narrow inner pad on apical one-half and as outer apical tuft of article 1, as elongate-narrow
pad on apical one-half of article 2, and as moderately large elongate-oval pad on apical one-half to two-thirds of article 3.
New World Cleonini
527
Abdomen. Ventral surface with dense, moderately robust appressed white scales, sparser and smaller medially on
abdominal sternum VII of female. Abdominal sterna III to VI (especially III and IV) with individually indistinct to
distinct moderately large rounded glabrous patches, each with large puncture and single suberect to erect, short to
moderately long hair-like scale situated in or near center. Base of abdominal sternum VII of female internally (dorsally)
with slightly to markedly developed median shiny tubercle. Genitalia. Female (five examined). Abdominal sternum VJII
with basal arm moderately long; lateral arms slightly to moderately inwardly arcuate at midlength; slightly expanded at
apex (Fig. 169). Stylus moderately large compared to length of gonocoxite II. Male (two examined). Internal sac (Figs.
189a,b) elongate and low; median dorsal pocket low, with single moderately broad elongate dorsally directed basal lobe
(lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B), with moderately large paired
dorsally directed lobe at basal one-third on dorsal surface (lobe C), with small paired dorsolaterally directed lobe near
midlength near dorsal margin (lobe D), with large paired dorsolaterally directed lobe just beyond midlength at dorsal
margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin (lobe F). Apical pocket and
median dorsal pocket individually distinct; apex of dorsal median pocket truncate in dorsal view. Ventral median pocket
moderately large. Apical pocket with paired moderately large ventrally directed lobe near apical one-third at ventral
margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface and basal portion
of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is known only from southeastern Arizona, southern
New Mexico, and extreme western Texas, south to central Mexico (Fig. 225).
Natural history. — This species appears to be associated with desert-grassland transitional
habitats. There is only a single record of one adult on Atriplex canescens (Pursh) Nutt.
(Chenopodiaceae). No definite hosts are known. Two individuals have been collected at
ultraviolet lights. Adults have been collected from March to December at elevations of
870-2161 m (N= 14) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. collaris (Fig 237).
Cleonidius poricollis species group
Diagnosis. — Size small for Cleonidius (Fig. 197). Dorsal erect or suberect vestiture absent
(Fig. 151a) to present, dense and moderately long (Fig. 150a). Head behind eye with or
without variously developed curved sulcus extended from above eye to area under postocular
lobe (Fig. 150a). Rostrum moderately robust, moderately to markedly medially tumescent, not
to very slightly carinate; lateral margins rounded. Pronotum with postocular lobes
well-developed; postocular vibrissae uniformly short (Figs. 150a- 15 la). Elytra with scale
pattern more or less vittate, intervals other than 9 to 1 1 with white scales equal in size to those
on intervals 9 to 1 1 (Figs. 131-132). Tarsus with ventral pilose vestiture extensive (on greater
part of ventral surface of each article) to slightly reduced in extent (on not less than apical
one-half of articles 2 and 3). Wings present, of variable length (short in only few specimens).
Female with abdominal sternum VIII with basal arm short; lateral arms arcuate (Figs.
170-171). Male with aedeagus with internal sac with apex of dorsal median pocket rounded;
lobes A to F present; dorsal median pocket low in lateral view (Figs. 190-191).
Phylogenetic relationships. — Monophyly of this species group is based only on the presence
of a curved sulcus behind the eye. The C. poricollis group is the sister-group to the C. boucardi
species group, this lineage sister to the C. americanus species group (Fig. 237).
Cleonidius poricollis (Mannerheim)
(Figs. 131, 150, 170, 190, 228)
Cleonis vittatus Kirby 1837:199 [not Zoubkoff 1829:163]. Lectotype (here designated), male, one of two syntypes,
labelled “Co-type”, “namer”, “ Cleonus / vittatus/ Kirby/ Co-type” and with my designation label “LECTOTYPE/
Cleonis vittatus/ Kirby desig./ Anderson” (BMNH). Type locality, North America. O’Brien and Wibmer 1982
Quaest. Ent., 1987,23 (4)
528
Anderson
(catal., distn.).
Lixus poricollis Mannerheim 1843:291. Holotype (examined) female, labelled only with a handwritten label “poricollis" /
Escholtz/ Escholtz (ZMMU). Type locality, California. Gemminger and von Harold 1871 (catal.)3.
Cleonus vittatus ; Melsheimer 1853 (check.). Gemminger and von Harold 1871 (catal.). LeConte 1876a (key). Henshaw
1885 (check.). Fall 1901 (check.). Fletcher 1906. Fall and Cockerell 1907 (check.). Gibson 1914. Leng 1920 (catal.).
Cleonus virgatus LeConte 1876a: 150 [not Gyllenhal 1834:242]. Holotype (examined), male, labelled “Cal”, “C./
virgatus/ TYPE Lee.”, “Horn Coll/ H8521” and a red square label (MCZC). Type locality, Owen‘s Valley,
California. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1889. Leng 1920 (catal., as synonym).
Cleonus ( Cleonidius ) lobigerinus Casey 1891:191. Lectotype (here designated), female, one of eight syntypes, labelled
“Ari”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37281”, “C. Cl./ lobigerinus / Cas” and with my designation label
“ Cleonus / lobigerinus Csy./ LECTOTYPE/ desig. Anderson” (USNM). Type locality, Peach Springs, Arizona.
Champion 1902-1906 (in part; distn., misident. of Cleonidius puberulus). Csiki 1934 (catal.). Hatch 1971 (key,
redesc.).
Cleonus ( Cleonidius ) kirbyi Casey 1891:188 (in key). NEW SYNONYMY Holotype (examined), female, labelled “Vic./
Vane. I.”, “CASLY/ bequest/ 1925”, “TYPE USNM/ 37299”, “C.C/./ kirbyi/ Cas.” (USNM). Type locality,
Victoria, British Columbia. Csiki 1934 (catal.). Hatch 1971 (key, redesc.).
Cleonus ( Cleonidius ) poricollis ; Casey 1891 (key). Csiki 1934 (catal.).
Cleonus ( Cleonidius ) virgatus ; Casey 1891 (key). Csiki 1934 (catal.).
Cleonus ( Cleonidius ) vittatus ; Casey 1891 (key). Hatch 1971 (key, redesc.).
Cleonus lobigerinus-, Wickham 1896 (check.). Fall 1901 (check.). Yothers 1916. Leng 1920 (catal.). Tanner 1966
(redesc., biol.).
Cleonus ( Cleonidius ) boucardi-, Champion 1902-1906 (in part; distn., misident.).
Cleonus kirbyi-, Wilcox et al.. 1934. Wickham 1902 (check.). Leng 1920 (catal.).
Cleonidius vittatus-, Faust 1904 (desig. as type species of Cleonidius).
Cleonus quadrilineatus-, Anderson 1914 (misident.).
Cleonus poricollis-, Leng 1920 (catal.).
Cleonus sparsus-, Essig 1958 (in part; biol., misident.). Arnett et al. 1980 (probable misident.).
Cleonus ( Cleonidius ) quadrilineatus-, Hatch 1971 (key, redesc., misident.).
Cleonus ( Cleonidius ) sparsus-. Hatch 1971 (in part; key, redesc., misident.).
Cleonis frontalis-, O’Brien and Wibmer 1982 (in part; catal., distn., misident.).
Cleonis kirbyi-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis lobigerinus-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis poricollis-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis virgatus-, O’Brien and Wibmer 1982 (catal., distn.).
vittatus, incertae sedis\ Chevrolat 1873.
Notes about synonymy. — There has been much confusion regarding this species and
consequently, numerous published misidentifications. Cleonidius poricollis , as considered here,
includes all those western North American individuals that possess a variously developed,
curved sulcus behind each eye (Fig. 150a). Mixed series are however known that include
specimens with the sulcus well to only slightly developed, to absent. These sympatric specimens
are otherwise indistinguishable and are considered herein to be conspecific. Primary types of
Lixus poricollis Mannerheim, Cleonis vittatus Kirby, Cleonus virgatus LeConte, Cleonus
( Cleonidius ) lobigerinus Casey, and Cleonus (Cleonidius) kirbyi Casey each possess a curved
sulcus behind the eye.
Cleonus (Cleonidius) kirbyi Casey was initially proposed by Casey (1891) as a replacement
name for Cleonus vittatus LeConte, however LeConte (1876a) did not describe a new species,
Cleonus vittatus , but rather was referring to Cleonus vittatus Kirby contrary to assertions of
Casey (1891). Thus Cleonus (Cleonidius) kirbyi Casey should be considered a new species
description and not a replacement name. Casey (1891), based only on the brief and inadequate
published descriptions of Cleonis vittatus Kirby and Lixus poricollis Mannerheim, incorrectly
considered Cleonis vittatus Kirby as a distinct species from Cleonus vittatus Kirby, sensu
LeConte (1876a), but correctly as conspecific with Cleonus poricollis (Mannerheim).
3Leconte 1876a,b (misident. Cleonidius erysimi). Henshaw 1885 (check., misident. Cleonidius erysimi).
New World Cleonini
529
Both Cleonis vittatus Kirby and Cleonus virgatus LeConte are junior homonyms.
Faust (1904) designated Cleonis vittatus Kirby as the type species of Cleonidius. I have
seen a specimen in the British Museum (Natural History), correctly determined by Faust and
labelled in his distinctive handwriting.
Problems in recognition. — Most members of this widespread species are easily recognized
by the presence of a variously developed (usually deep) curved sulcus behind each eye (Fig.
150a). In many specimens the sulcus is present but largely to completely covered by the
postocular lobe. In specimens in which the postocular lobe covers the sulcus, reorientation of the
head exposes the sulcus to view. If this is not done, such individuals prove difficult to separate
from other species, especially C. notolomus and C. frontalis. Specimens lacking a sulcus or
with only a slightly developed sulcus appear confined to Oregon and British Columbia (perhaps
also Washington, although no such specimens have been seen from that area) and are separated
from other species of Cleonidius by the sparse scales of the elytra (forming a faintly vittate
pattern), possession of large pilose ventral tarsal pads, and rostrum with distinct median
tumescence, the base of the rostrum distinctly separated from the frons by a transverse
impression (Fig. 150a).
The eastern C. calandroides is distinguished from the western C. poricollis by the short
suberect dorsal vestiture (Fig. 151a) and less distinctly vittate elytral scale pattern (intervals 6
to 8 with scales as large as those on intervals 9 to 11) (Fig. 132) in the former. The sulcus is
variously developed in individuals of both species but is moderately deep and more distinct in
most C. poricollis and in only few C. calandroides.
Description. — Specimens examined. 387 males, 459 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 28. Size. Length, male, 4. 3-9. 3 mm; female, 5.6-10.3 mm. Width, male, 1.8-3. 6 mm; female, 2. 0-4.0 mm. Head.
Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle or with variously developed (moderately
deeply in most specimens) curved sulcus extended posteroventrally from upper one-half of posterior margin of eye to area
under postocular lobe (sulci continued ventrally and joined at gular suture but covered by prothorax and not visible) (Fig.
150a). Frons with sparse to moderately dense, small shallow punctures, interspersed with minute punctures. Frons also
with very sparse to moderately dense, short to moderately long, suberect to erect hair-like scales, moderately dense
elongate-fine appressed white scales medially, suberect to erect immediately above eyes. Frons distinctly separated from
base of rostrum by moderately deep to deep transverse impression (Fig. 150a). Base of median tumescence of rostrum with
small, shallow to moderately deep fovea. Width of frons less than, to subequal to, width at apex of rostrum. Rostrum.
Moderately robust, (width at apex 0.57-0.71 times length in male; 0.57-0.76 in female) (Fig. 150). In lateral view straight
to very slightly curved downward. In dorsal view with postgenae moderately to markedly laterally expanded, antennal
scrobes distinctly visible. Rostrum moderately to markedly medially tumescent from point of antennal insertion to middle
of frons; not laterally expanded at midlength; lateral margins rounded; with median carina variously developed as narrow
low glabrous shiny line at crest of median tumescence. Dorsal and lateral punctation moderately dense to dense, small,
shallow (many longitudinally confluent); dorsally interspersed with dense minute punctures. Dorsally with very sparse to
moderately dense, short to moderately long, suberect to erect hair-like scales; also with moderately dense, elongate-fine,
appressed to recumbent white scales, sparser medially. Epistoma slightly swollen. Pronotum. Median carina variously
developed in anterior one-half from absent to as low, rounded moderately broad glabrous line. Dorsal and lateral
punctation moderately large, moderately dense to dense, moderately deep to deep, not confluent; smaller, sparser and
shallower apically and on flanks; areas between larger dorsal punctures with or without minute regularly impressed
punctures. Dorsally with moderately dense, short to long, erect hair-like scales, each in large puncture (Fig. 150a). Scales
dorsally absent to small fine and sparse along median line; moderately dense to dense, elongate-fine to elongate-narrow,
appressed in pair of broad apically slightly narrowed paramedian stripes; laterally, absent to sparse, small and
elongate-fine. Lateral margins with scales moderately dense to dense, elongate-fine to elongate-narrow, appressed to
recumbent in moderately broad stripe. Flanks with scales elongate-fine to elongate-narrow, sparse to moderately dense.
Median basal portion of disk shallowly to deeply and narrowly impressed. Anterolateral margin with pronotal postocular
lobes well-developed; postocular vibrissae uniformly short (Fig. 150a). In dorsal view with lateral margins slightly arcuate
and convergent from base to apical one-quarter, slightly constricted at apical one-quarter then straight and convergent to
apex. Prosternum. With small shallow to moderately deep impression and moderately to markedly developed rounded
swelling anterior to each procoxal cavity. In lateral view with anterior flat to slightly angulate portion long, approximately
two-thirds to subequal in length to, posterior more angulate portion; posterior and anterior portions not or slightly
separated by shallow transverse impression. Elytra. Elongate-narrow in general form (width at midlength 0.50-0.63 times
Quaest. Ent., 1987, 23 (4)
530
Anderson
length in males; 0.50-0.64 in females) (Fig. 131). In dorsal view with lateral margins straight to very slightly sinuate and
slightly convergent from apical one-third to humerus; moderately and evenly arcuate from apical one-third to apex. Elytral
apices not produced, rounded, slightly divergent. Humerus distinct. Dorsally with suberect or erect vestiture lacking to
dense, moderately long, erect hair-like scales; with sutural interval with dense, very small and fine to elongate-fine, golden
to white scales, interspersed with scattered patches of larger white scales, to with scales uniformly white, dense, small and
elongate-fine; intervals 2 with scales golden to white, dense, very small and fine, with scattered patches of larger white
scales; intervals 3 to 5 with dense, small and fine to elongate-narrow white scales; intervals 6 to 8 with scales dense, very
small and very fine, golden to white, with scattered patches of larger white scales; intervals 9 to 1 1 with uniformly dense,
small to large, elongate-fine to elongate-narrow white scales (Fig. 131). Punctures of elytral striae small, individually
distinct, arranged in regular rows. Wings. Long (greater than elytra in length [99%, N = 838]), to short (approximately
one-half length of elytra or less [1%, N = 8]). Legs. Foretibia of female with inner margin with small denticles in apical
one-half; subapical tooth indistinct from tibial denticles. Foretibial uncus of both sexes moderately large, mesotibial and
metatibial unci small. Tarsal claws connate in basal one-third, not divergent, each with basal internal flange slightly
developed. Ventral tarsal pilose vestiture of foretarsus, mesotarsus, and metatarsus of male and female present as
elongate-oval pad on apical one-half to three-quarters of article 1, as moderately large rounded pad on apical one-half of
article 2, and as large round pad on apical three-quarters of article 3 in male, apical one-half to three-quarters in female.
Abdomen. Ventral surface with moderately dense, elongate-narrow to moderately robust appressed white scales.
Abdominal sterna III to VI (especially III and IV) with individually indistinct to distinct moderately large rounded
glabrous patches, each with large puncture and single suberect to erect, moderately long hair-like scale situated in or near
center. Base of abdominal sternum VII of female internally (dorsally) with slightly to well-developed median shiny
tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm short; lateral arms slightly inwardly
arcuate throughout length; very slightly expanded at apex (Fig. 170). Stylus moderately large compared to length of
gonocoxite II. Male (four examined). Internal sac (Figs. 190a, b) elongate and low; median dorsal pocket low, with single
moderately broad elongate dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on
dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C),
with moderately large paired dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired
dorsolaterally directed lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe
near base near ventral margin (lobe F). Apical pocket and median dorsal pocket individually distinct; apex of dorsal
median pocket rounded in dorsal view. Ventral median pocket moderately large. Apical pocket with paired moderately
large ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple
scythe-like sclerite; adjacent ventral surface of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic variation. — Specimens from coastal localities in Oregon and British Columbia
(likely also Washington) differ in a number of respects from individuals from throughout the
rest of the species range, exhibiting: generally slightly larger prosternal swellings; generally
deeper and denser punctation of head, rostrum and pronotum; smaller elytral scales, the pattern
only faintly vittate; lack of sulcus behind the eye; lack of or else very short indistinct dorsal
suberect or erect vestiture; and broad foretarsal articles with more extensive ventral pilosity.
Not all individuals however possess all of these character states; mixed series are known in
which some individuals possess various combinations of the above character states but
otherwise possess the typical C. poricollis states. As such, no consistent set of character states
can be used to justify separation of these Pacific Northwest individuals as a distinct taxonomic
entity. A large number of these specimens have been reared from Erysimum sp. (wallflower;
Cruciferae at Victoria, British Columbia, but other typical C. poricollis have also been reared
from various Cruciferae throughout the species range. C. calandroides, sister-species of C.
poricollis also occurs on Cruciferae.
Individuals from a series of specimens from San Francisco, California, have the elytra
uniformly covered with moderately dense, elongate-fine white scales; the elytra do not appear
distinctly vittate. Otherwise these specimens are typical C. poricollis and in my view do not
warrant separate status.
Geographic distribution. — This is the most widely distributed species of Cleonidius,
ranging throughout western North America across southern British Columbia, Alberta and
Saskatchewan in Canada into the adjacent United States of America east to Michigan in the
north, south along the west coast to California in the west and south to western Texas and
central Mexico in the east (Fig. 228).
New World Cleonini
531
Smith (1957) discusses taxa with populations in prairie habitats in Michigan that are
disjunct from populations in the western prairie. The distribution of C. poricollis in the
northern United States appears to fit this pattern.
Natural history. — This species is associated with a wide variety of habitats: grassland,
desert-grassland transitional, Pacific semi-desert, Great Basin desert, Sonoran desert, Mojave
desert, and western xerophytic evergreen forest. Adults of this species have been collected on a
wide taxonomic range of plants as follows: Asclepias sp. (Asclepiadaceae); Atriplex sp., A.
nutalli Wats., A. rosea L., Beta vulgaris L. (beet), Eurotia lanata (Pursh) Moq., Salsola kali
L. (all Chenopodiaceae); Ambrosia sp., Franseria confertiflora (DC.) Rydb., Chrysothamnus
sp., C. nauseosus (Pall.) Britton, C. viscidiflorus (Hook.) Nutt. var. typicus, Gutierrezia
californica (DC.) Torr. and Gray, Aplopappus acradenius (Greene) Blake, A. venetus Blake
ssp. vernonioides (Nutt.) Hall, Hymenoclea monogyra T. and G., Zinnia sp. (all Compositae);
Sisymbrium altissimum L., “S. pestifer ”, Erysimum sp. (wallflower), Descurainia sophia. ( L.)
Webb., Brassica campestris L. (mustard), B. oleracea L. (broccoli) (all Cruciferae); Salvia sp.
(Labiatae); Lupinus sp.. Trifolium sp., Melilotus albus Desr., Medicago sativa L. (all
Leguminosae); Fragaria sp., Rubus sp., Purshia tridentata (Pursh) DC. (all Rosaceae). Adults
have been reared or collected from the roots of Chrysothamnus sp., Gutierrezia sp., G.
microcephala (DC.) Gray, Aplopappus sp., A. venetus (Hbk.) Blake (all Compositae);
Brassica sp., Raphanus sativus L. (radish), Erysimum sp., Descurainia pinnata (Walt.)
Britton (all Cruciferae). Tanner (1966) reports collection of adults from a Grayia-Lycium
community, a Salsola community, and on Atriplex canescens (Pursh) Nutt.. Hatch (1971)
notes larvae of this species injuring radishes at Corvallis, Oregon (as Cleonus sparsus) and as
attacking wallflower roots on a seed farm on Vancouver Island, British Columbia (as Cleonus
kirbyi).
This species has been collected on various types of vegetation primarily in dry washes and
stream beds throughout the species range. Individuals have also been collected on sand dunes or
other sandy habitats.
I have seen a single specimen collected from the stomach of Bufo sp. (Amphibia).
Adults have been collected from June to November at elevations from sea level to 4004 m
(N = 132) (Fig. 233).
Chorological relationships. — Table V. C. poricollis is allopatric to its sister-species C.
calandroides.
Phylogenetic relationships. — This species is the sister-species of C. calandroides , the only
other species in the C. poricollis species group (Fig. 236).
Cleonidius calandroides (Randall)
(Figs. 132, 151, 171, 191,229)
Lixus calandroides Randall 1838:42. Type(s) lost. Gemminger and von Harold 1871 (catal.). Sprague and Austin 1875.
LeConte 1876a (misinterpret., as synonym of Lixus musculus).
Cleonus calandroides ; LeConte 1876b (diagnosis). Henshaw 1885 (check.). Casey 1891 (key). Ely 1913 (biol.). Blatchley
and Leng 1916 (key, redesc., biol.). Leng 1920 (catal.). Leonard 1926 (check., biol.). Csiki 1934 (catal.). Bruhn 1947
(morphol.). Sanders 1960 (morphol.). Kissinger 1964 (biol.).
Cleonus ( Cleonidius ) calandroides-, Casey 1891 (key). Csiki 1934 (catal.).
Cleonis calandroides-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — Lixus calandroides was described from an unspecified number of
specimens from Chelsea Beach, Massachusetts (Randall 1838). Unfortunately type material of
Randall has been lost (Sprague and Austin 1875). Following reccomendations of the
Quaest. Ent., 1987, 23 (4)
532
Anderson
International Code of Zoological Nomenclature, no neotype designation is required, because
there has been no recent confusion regarding identity of this species. After identification of
specimens as this species by Sprague and Austin (1875), LeConte (1876b), based on
examination of specimens sent to him by Austin, transferred the species to Cleonus and
removed the name from his previously proposed synonymy with Lixus musculus Say (LeConte
1876a).
Problems in recognition. — This is one of only two species of Cleonidius restricted to eastern
North America (Figs. 215, 229). These two differ markedly in structural features and should
not be confused. The eastern C. calandroides is distinguished from the western C. poricollis by
the short suberect dorsal vestiture (Fig. 151a) and less distinctly vittate elytral scale pattern
(intervals 6 to 8 with scales as large as those on intervals 9 to 11) (Fig. 132) in the former. The
sulcus behind the eye is variously developed in individuals of both species but is moderately
deep and distinct in most C. poricollis and in only few C. calandroides (see descriptions).
Description. — Specimens examined. 65 males, 63 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 29. Size. Length, male, 8.0-9. 3 mm; female, 5. 5-9.6 mm. Width, male, 3.0-3.6 mm; female, 2. 1-3.9 mm. Head.
Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle or with variously developed (moderately
deeply in only few specimens) curved sulcus extended posteroventrally from upper one-half of posterior margin of eye to
area under postocular lobe (sulci continued ventrally to juncture at gular suture but covered by prothoracic structures and
not visible) (as in Fig. 150a). Frons with only scattered small shallow punctures, interspersed with minute punctures. Frons
also with moderately dense, short, suberect hair-like scales, sparse to moderately dense elongate-fine appressed white
scales medially, suberect to erect immediately above eyes. Frons distinctly separated from base of rostrum by moderately
deep to deep transverse impression (Fig. 151a). Base of median tumescence of rostrum with small, shallow to moderately
deep fovea. Width of frons greater than to subequal to width at apex of rostrum. Rostrum. Moderately robust, slightly less
so in females (width at apex 0.60-0.68 times length in male; 0.56-0.65 in female) (Fig. 151). In lateral view straight to very
slightly curved downward. In dorsal view with postgenae moderately laterally expanded, antennal scrobes partially visible.
Rostrum medially tumescent from point of antennal insertion to middle of frons; not laterally expanded at midlength;
lateral margins rounded; with median carina variously developed as narrow low glabrous shiny line at crest of median
tumescence. Dorsal and lateral punctation moderately dense, small, shallow (many longitudinally confluent); dorsally
interspersed with dense minute punctures. Dorsally with very sparse to moderately dense, short, suberect hair-like scales
(especiallly laterally); also with moderately dense, elongate-fine appressed to recumbent white scales, sparser medially.
Epistoma slightly swollen. Pronotum. Median carina variously developed in anterior one-half from absent to as low,
rounded narrow glabrous line. Dorsal and lateral punctation moderately large, moderately dense to dense, shallow, not
confluent; smaller apically and on flanks; areas between larger dorsal punctures with or without minute regularly
impressed punctures. Dorsally with moderately dense, very short, suberect hair-like scales, each situated in large puncture
(Fig. 151a). Scales dorsally absent to small fine and sparse along median line; moderately dense, elongate-narrow,
appressed in pair of broad apically slightly narrowed paramedian stripes; laterally, absent to sparse, small and
elongate-fine. Lateral margins with scales dense, elongate-narrow, appressed in moderately broad stripe. Flanks with
scales dense, elongate-fine to elongate-narrow. Median basal portion of disk shallowly to deeply and narrowly impressed.
Anterolateral margin with pronotal postocular lobes well-developed; postocular vibrissae uniformly short (Fig. 151a). In
dorsal view with lateral margins slightly arcuate and slightly convergent from base to apical one-quarter, slightly
constricted at apical one-quarter then straight and convergent to apex. Prosternum. With small shallow impression and
slight to moderately developed rounded swelling anterior to each procoxal cavity. In lateral view with anterior flat to
slightly angulate portion long, subequal in length to posterior more angulate portion; posterior and anterior portions
slightly separated by shallow transverse impression. Elytra. Moderately robust in general form (width at midlength
0.57-0.60 times length in males; 0.55-0.63 in females) (Fig. 132). In dorsal view with lateral margins straight, subparallel
to slightly convergent from apical one-third to humerus; moderately and evenly arcuate from apical one-third to apex.
Elytral apices not produced, rounded, slightly divergent. Humerus distinct. Dorsally with moderately dense, very short,
suberect, indistinct hair-like scales; scales more or less uniformly dense, white; with sutural interval and interval 2 with
scales small and fine; intervals 3 to 5 and 9 to 1 1 with scales large and elongate-narrow; scales small and fine to large and
elongate-narrow on intervals 6 to 8 (Fig. 132). Punctures of elytral striae small, individually distinct, arranged in regular
rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with small denticles in
apical one-half; subapical tooth indistinct from tibial denticles. Foretibial and mesotibial unci of both sexes moderately
large, metatibial uncus small. Tarsal claws connate in basal one-half, slightly divergent, each with basal internal flange
slightly developed. Ventral tarsal pilose vestiture of foretarsus of male present as large pads on more or less entire ventral
surface of articles 2 and 3 and apical one-half of article 1; of female, as elongate pad on apical one-half of article 1, as
large oval pad on apical three-quarters of article 2, and as large round pad on more or less entire ventral surface of article
New World Cleonini
533
3; of mesotarsus of male and female as small elongate-narrow pad on apical one-half of article 1, as large elongate pad on
more or less entire ventral surface on articles 2 and 3; of metatarsus of male and female as elongate pad on apical
three-quarters of article 1, as large elongate-oval pad on apical one-half of article 2, and as large oval pad on apical
three-quarters of article 3. Abdomen. Ventral surface with moderately dense to dense, elongate-narrow appressed white
scales. Abdominal sterna III to VI (especially III and IV) with individually indistinct to distinct moderately large rounded
glabrous patches, each with large puncture and single appressed hair-like scale situated in or near center. Base of
abdominal sternum VII of female internally (dorsally) with moderately developed median shiny tubercle. Genitalia.
Female (two examined). Abdominal sternum VIII with basal arm short; lateral arms slightly inwardly arcuate throughout
length; very slightly expanded at apex (Fig. 171). Stylus moderately large compared to length of gonocoxite II. Male (four
examined). Internal sac (Figs. 191 a,b) elongate and low; median dorsal pocket low, with single moderately broad elongate
dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B),
with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), with moderately large
paired dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired dorsolaterally directed
lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near ventral
margin (lobe F). Apical pocket and median dorsal pocket individually distinct; apex of dorsal median pocket rounded in
dorsal view. Ventral median pocket moderately large. Apical pocket with paired moderately large ventrally directed lobe
near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent
ventral surface of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is distributed along the northern Atlantic Coast
from Long Island, New York, north to New Hampshire and New Brunswick (Fig. 229).
Natural history. — This species breeds in the sea-rocket, Cakile edentula (Bigel.) Hook.
(Cruciferae). Ely (1913) collected various life stages from larvae to adults in the roots of this
plant in mid-August. He reports that larvae live entirely within the root and that the portion of
the root living within two or three inches of the surface of the ground is most commonly
attacked. Pupation takes place in a coccoon constructed of small shreds of plant material built
up in a longitudinal excavation in one side of the root. He further states that the beetles must be
very common as virtually all plants examined showed evidence of feeding by larvae. Blatchley
and Leng (1916) also state that larvae breed in Cakile edentula and other specimens I have
examined bear label data indicating they were reared from this plant species. No other plant
associations are known. Adults have been collected only on sea beaches from April to October.
Chorological relationships. — Table V. Cleonidius calandroides is sympatric only with
northern populations of C. subcylindricus.
Phylogenetic relationships. — This species is the sister-species of C. poricollis, the only
other species in the C. poricollis species group (Fig. 236).
Cleonidius boucardi species group
Diagnosis — Size moderately large to large for Cleonidius (Fig. 197). Dorsal erect or
suberect vestiture absent (Fig. 152a) to present, moderately dense and moderately long (Fig.
155a). Rostrum moderately robust, not (Fig. 152a) to moderately (Fig. 155a) medially
tumescent, not to variously carinate; lateral margins rounded. Pronotum with postocular lobes
absent (Fig. 155a) to very slightly developed (Fig. 152a); postocular vibrissae moderately long
to long, of unequal length and longest behind base of eye. Elytra with scale pattern more or less
vittate, intervals other than 9 to 1 1 with white scales equal in size to those on intervals 9 to 1 1
(Figs. 133-136). Tarsus with ventral pilose vestiture extensive (on greater part of ventral
surface of each article) to slightly reduced in extent (on not less than apical one-half of articles
2 and 3). Wings present, long. Female with abdominal sternum VIII with basal arm short to
moderate in length; lateral arms arcuate (Figs. 172-175). Male with aedeagus with internal sac
with apex of dorsal median pocket variously emarginate or sinuate (Figs. 192a,b-195a,b); lobes
A to F present; dorsal median pocket low in lateral view.
Quaest. Ent., 1987, 23 (4)
534
Anderson
Phylogenetic relationships. — Primary characters show this species group to be
monophyletic and a member of the unresolved heptachotomy also including the C. front alis-C.
americanus lineage, the C. infrequens-C. puberulus lineage, C. canescens , C. collaris, C.
notolomus (all of the C. americanus group), and the C. poricollis species group (Fig. 236).
Secondary characters further support the monophyly of the group and indicate that the C.
boucardi group is the sister-group to the C. poricollis species group, this lineage sister to the C.
americanus species group (Fig. 237).
Cleonidius boucardi (Chevrolat)
(Figs. 133, 152, 172, 192, 221)
Apleurus boucardi Chevrolat 1873:79. Holotype (examined), male, labelled “Puebla”, “[letter illegible] Boucard”, pale
blue square label, “Typus”, “41”, “471/ 85”, “Riksmuseum/ Stockholm” and with my label ” Apleurus boucardi/
Chevrolat/ HOLOTYPE/ lab. Anderson” (NRS). Type locality, Puebla, Mexico.
Cleonus carinicollis LeConte 1876a: 152 [not Gyllenhal 1834:241], NEW SYNONYMY. Holotype (examined), female,
labelled with a pale green circle ( = Wyoming Territory), "Type/ 5241”, “C. carinicollis/ Lee.” (MCZC). Type
locality, Colorado. Henshaw 1881-1882 (check.). Henshaw 1885 (check.). Wickham 1902 (check.). Blatchley and
Leng 1916 (key, redesc.). Leng 1920 (catal.). Brimley 1938 (check.). Arnett et al. 1980.
Cleonus ( Cleonidius ) boucardi-, Casey 1891 (key). Champion 1902-1906 (in part; distn., misident of Cleonidius
poricollis). Csiki 1934 (catal.).
Cleonus ( Cleonidius ) carinicollis-, Casey 1891 (key, misident. of Cleonidius trivittatus Say).
Cleonus ( Cleonidius ) lecontei Casey 1891:190. NEW SYNONYMY Holotype (examined), female, labelled “Ari”,
“CASEY/ bequest/ 1925”, “TYPE USNM/ 37280”, “C. Cl./ lecontei/ Cas.”(USNM). Type locality, Arizona. Csiki
1934 (catal.).
Cleonus lecontei-, Leng 1920 (catal.).
Cleonus ( Cleonidius ) lecontellus Csiki 1934:64. NEW SYNONYMY New name for Cleonus carinicollis LeConte.
Blackwelder 1939 (check.).
Cleonus boucardi-, Blackwelder 1947 (check.).
Cleonis boucardi ; O’Brien and Wibmer 1982 (catal., distn.).
Cleonis carinicollis-, O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Cleonis lecontei-, O’Brien and Wibmer 1982 (catal., distn.).
Cleonis lecontellus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — Chevrolat (1873), in describing C. boucardi noted the similarity
to C. trivittatus (Say). Although most individuals of the two species are quite distinct, some
individuals are difficult to reliably assign as belonging to either species. One such specimen is
the holotype of Cleonus carinicollis LeConte, which has dense, extensive ventral tarsal pilose
pads, moderately developed postocular lobes (both character states of C. boucardi ), but also
has a rostrum that is medially tumescent and has a low median carina, and an elytral scale
pattern which is not that of most C. boucardi (the latter, all states of C. trivittatus). The type
locality is “Colorado Territory” and therefore within the ranges of both species. I have chosen
to regard this individual as conspecific with C. boucardi and accordingly emphasize the
taxonomic significance of the characters of extent of ventral tarsal vestiture and development of
postocular lobes as diagnostic for these two species.
Cleonidius lecontellus Csiki is a typical C. boucardi , differing from the latter, as noted by
Casey (1891), only in details of the elytral scale pattern.
Problems in recognition. — Most individuals of this species are likely to be confused only
with C. trivittatus and C. quadrilineatus. They are however, easily separated by the rostrum
not or only slightly dorsally medially tumescent (Fig. 152a), postocular lobes slightly to
moderately developed (Fig. 152a), and with all tarsal articles with extensive ventral pilose pads
(on more or less entire ventral surface of each article) in C. boucardi specimens. In both C.
trivittatus and C. quadrilineatus the rostrum is medially dorsally tumescent and carinate (Figs.
153a, 155a), the postocular lobes are not to moderately developed (but only in few specimens of
New World Cleonini
535
C. trivittatus, especially in northern part of the species range in southern Canada) (Figs. 153a,
155a), and the ventral tarsal vestiture is less extensive. A few C. boucardi specimens from
Kansas and Texas are very similar to C. trivittatus in elytral scale pattern and in rostral
characters but can be separated by their extensive ventral tarsal pilosity and moderately
developed postocular lobes.
Cleonidius boucardi may also be confused with some C. poricollis that have a similar
faintly vittate elytral scale pattern and lack the curved sulcus behind the eye. Although C.
boucardi and C. poricollis are sympatric over part of their ranges, those individuals of C.
poricollis that are most likely to be confused with C. boucardi are found only in the Pacific
Northwest; C. boucardi does not occur in this or adjacent areas.
This is the only species of Cleonidius that has been extensively collected in Mexico (Fig.
221).
Description. — Specimens examined. 73 males, 82 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 30. Size. Length, male, 7.5-14.7 mm; female, 8.4-18.6 mm. Width, male, 3. 2-5. 9 mm; female, 3. 4-7.0 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with sparse to moderately dense,
small to large, shallow to deep punctures, largest, deepest and longitudinally confluent in many specimens immediately
above eyes; interspersed with sparse minute punctures. Frons also with sparse, short, suberect hair-like scales, sparse to
moderately dense elongate-fine, appressed to recumbent white scales medially, suberect to erect immediately above eyes.
Frons continuous with base of rostrum or only very slightly separated from base of rostrum by very shallow transverse
impression (Fig. 152a). Base of median tumescence of rostrum with small, shallow to moderately deep fovea. Width of
frons subequal to width at apex of rostrum. Rostrum. Moderately robust, (width at apex 0.57-0.67 times length in male;
0.59-0.66 in female) (Fig. 152). In lateral view straight to very slightly curved downward. In dorsal view with postgenae
slightly laterally expanded, antennal scrobes distinctly visible. Rostrum not to very slightly medially tumescent from point
of antennal insertion to middle of frons; not laterally expanded at midlength; lateral margins rounded; with median carina
variously developed as narrow to broad low glabrous shiny line. Dorsal and lateral punctation sparse to dense, small to
large, moderately deep to deep (dense and large in most specimens with many punctures longitudinally confluent);
punctures smaller and shallower apically; dorsally interspersed with dense minute punctures. Dorsally with scattered,
short, suberect hair-like scales in some specimens; also with sparse to moderately dense, elongate-fine to elongate-narrow,
appressed to recumbent white scales. Epistoma very slightly swollen. Pronotum. Median carina variously developed in
anterior one-half from as minutely punctate slightly elevated broad line to elevated sharp narrow glabrous shiny line,
well-developed and distinct in most specimens. Dorsal and lateral punctation small, moderately dense, shallow, not
confluent; smaller apically and on flanks; areas between larger dorsal punctures with dense minute regularly impressed
punctures. Lacking dorsal suberect or erect vestiture (Fig. 152a). Scales dorsally absent in moderately broad median
longitudinal line; moderately dense, elongate-fine to elongate-narrow, appressed in pair of narrow to broad apically slightly
narrowed paramedian stripes; laterally, scales absent to very sparse, small and fine. Lateral margins with scales
moderately dense, elongate-narrow, appressed in moderately broad stripe. Flanks with scales sparse to moderately dense,
elongate-fine. Median basal portion of disk shallowly to moderately deeply and broadly impressed. Anterolateral margin
with pronotal postocular lobes very slightly to moderately developed; postocular vibrissae of unequal length, moderately
long to long, greatest length (from one-half to two-thirds width of eye) behind base of eye (Fig. 152a). In dorsal view with
lateral margins uniformly slightly arcuate and convergent from base to apex, or slightly constricted at apical one-quarter,
then straight and convergent to apex. Prosternum. With small very shallow indistinct impression and lacking or with at
most only very slight rounded swelling anterior to each procoxal cavity. In lateral view with anterior flat to slightly
angulate portion short, approximately equal to one-half length of posterior more angulate portion; posterior and anterior
portions not to slightly differentiated by shallow transverse impression. Elytra. Moderately robust in general form (width
at midlength 0.57-0.62 times length in males; 0.54-0.61 in females) (Fig. 133). In dorsal view with lateral margins
straight, to very slightly sinuate and slightly convergent from apical one-third to humerus; moderately and evenly arcuate
from apical one-third to apex. Elytral apices not to slightly produced, rounded to subacuminate, not to slightly divergent.
Humerus distinct. Dorsally lacking suberect or erect dorsal vestiture; sutural interval with moderately dense, small fine
white scales; interval 2 with scales absent to sparse small and fine; interval 3 with scales moderately dense large
elongate-narrow; intervals 4 and 5 with scales absent to moderately dense, moderately large and elongate-fine; intervals 6
to 8 with scales absent to sparse small and fine; intervals 9 to 1 1 with scales moderately dense, large elongate-narrow;
striae 1, 6 and 7 with irregularly distributed patches of larger and denser white scales (Fig. 133). Punctures of elytral
striae small, individually distinct, arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of
female with inner margin with small to large denticles in apical three-quarters; subapical tooth large, distinct from apical
denticles. Foretibial and mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in
basal one-third to one-half, not to slightly divergent, each with basal internal flange well-developed. Ventral tarsal pilose
vestiture of all tarsal articles present as large pads on more or less entire ventral surface of articles 2 and 3 and apical
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536
Anderson
one-half to three-quarters of article 1. Abdomen. Ventral surface with sparse to moderately dense, elongate-fine to
elongate-narrow appressed white scales. Abdominal sterna III to VI lacking glabrous patches or with sterna III and IV
with individually indistinct small rounded glabrous patches, each with large puncture and single appressed hair-like scale
situated in or near center. Base of abdominal sternum VII of female internally (dorsally) with very slightly developed
median shiny tubercle. Genitalia. Female (four examined). Abdominal sternum VIII with basal arm short; lateral arms
moderately inwardly arcuate throughout length; slightly expanded at apex (Fig. 172). Stylus moderately large compared
to length of gonocoxite II. Male (three examined). Internal sac (Figs. 192a, b) elongate and low; median dorsal pocket low,
with single narrow elongate dorsally directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on
dorsal surface (lobe B), with moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C),
with moderately large paired markedly dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with
small paired dorsolaterally directed lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally
directed paired lobe near base near ventral margin (lobe F). Apical pocket and median dorsal pocket individually distinct;
apex of dorsal median pocket slightly emarginate medially in dorsal view (Fig. 192b). Ventral median pocket large. Apical
pocket with paired moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible apical
sclerite complex with paired simple scythe-like sclerite; adjacent ventral surface and basal portion of sides of apical pocket
with pair of large, dark, longitudinal sclerites.
Geographic variation. — Specimens from Kansas and Texas in the northern part of the
species range have a rostrum that is slightly medially tumescent with smaller dorsal punctation,
and an elytral scale pattern that is more distinctly vittate.
Geographic distribution. — This species is found from extreme southern California and
Arizona in the west, and Kansas and northern Texas in the east, south through central Mexico
to Honduras (Fig. 221).
Natural history.— This species lives in mesquite-grassland and grassland habitat
throughout Mexico and the southern United States. Plant associations are not known for adults
of this species, which have been collected from March to September at elevations of 255-3385
m (N = 29) (Fig. 233).
Chorological relationships.— Table V.
Phylogenetic relationships. — This species is the sister-species of the C. trivittatus-C.
quadrilineatus-C. placidus lineage in the C. boucardi group (Fig. 237).
Cleonidius trivittatus (Say)
(Figs. 134, 153, 173, 193, 220)
Cleonus trivittatus Say 1831:10. Neotype (here designated, from the LeConte Collection), female, labelled with a pale
green circle ( = Wyoming Territory), “1847”, “J. LECONTE/ COLLECTION”, “C/./ trivittatus / Say” and with my
designation label “ Cleonus trivittatus/ Say NEOTYPE/ desig. Anderson” (MCZC). Type locality, Colorado.
Melsheimer 1853 (check.). LeConte 1859b (error as C. vittatus Say). Gemminger and von Harold 1871 (catal.).
LeConte 1876a (key). Henshaw 1885 (check.). Wickham 1902 (check.). Fall and Cockerell 1907 (check.). Leng 1920
(catal.). Blackwelder 1947 (check.). Essig 1958 (biol.). Kumar et al. 1976 (in part; biol.).
Rhynchophorus praepotens Say 1831:21 . Type destroyed.
Lixus praepotens-, Boheman 1836 (redescr.).
Apleurus trivittatus-, Chevrolat 1873.
Cleonus inornatus LeConte 1876a: 149. NEW SYNONYMY Holotype (examined), female, labelled “Cal”, “C./
inornatus/ TYPE LeC”, “Horn Coll/ H 8519” (MCZC). Type locality, Owen‘s Valley, California. Henshaw
1881-1882 (check.). Henshaw 1885 (check.). Fall 1901 (check.). Leng 1920 (catal.).
Cleonus ( Cleonidius ) bicarinatus Casey 1891:190 [not Gebler 1830:158]. NEW SYNONYMY Holotype (examined),
male, labelled “Tex”, “CASEY/ bequest/ 1925”, “TYPE USNM/ 37279”, “C. Cl./ bicarinatus/ Cas” (USNM). Type
locality, near Austin, Texas. Csiki 1934 (catal.).
Cleonus ( Cleonidius ) carinicollis-, Casey 1891 (key, misident.).
Cleonus ( Cleonidius ) inornatus-, Casey 1891 (key). Csiki 1934 (catal.).
Cleonus ( Cleonidius ) praepotens-, Casey 1891 (as synonym).
Cleonus ( Cleonidius ) trivittatus-, Casey 1891 (key). Csiki 1934 (catal.).
Cleonus bicarinatus-, Leng 1920 (catal.).
Cleonus quadrilineatus-. Pierce 1907 (biol., misident.). Chittenden 1911 (biol., misident.). Blatchley and Leng 1916
(figure, misident.). Yothers 1916 (misident., in part). Essig 1958 (biol., misident.). Arnett et al. 1980 (key, redesc.,
biol., misident.).
New World Cleonini
537
Cleonis bicarinatus\ O'Brien and Wibmer 1982 (catal., distn.).
Cleonis inornatus\ O’Brien and Wibmer 1982 (catal., distn.).
Cleonis praepotens, O’Brien and Wibmer 1982 (catal., distn. as synonym).
Cleonis trivittatus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — The type series of this species has been lost. I have selected and
designated as neotype, a specimen from the LeConte collection, because it is generally
considered that LeConte compared his material with that of Say before the loss of the latter.
This collection thus represents one of the best sources of reliably identified specimens of Say
species. The type locality of the original type series is given as “Arkansaw” (Say 1831) but it is
later stated that he (Say) “obtained two or three specimens near the Rocky Mountains”. The
neotype selected is from “Colorado Territory” which probably refers to the same general area
as that intended by the use of “Arkansaw” by Say (1831), which, as indicated by the reference
to the Rocky Mountains, undoubtedly refers to more than what is presently the state of
Arkansas.
Rhynchophorus praepotens Say (type lost) was regarded by LeConte (1859b) as conspecific
with Cleonus trivittatus Say. This synonymy has since been generally followed and thus no
neotype designation is required. LeConte (1859b) also states that Lixus praepotens (Say) of
Boheman (1836) is evidently this same species. Lixus praepotens (Say) of Boheman (1836)
therefore represents only a reassignment from Rhynchophorus to Lixus and a species
redescription, not description of a new species as indicated by Csiki (1934) and O’Brien and
Wibmer (1982).
The holotype of Cleonus inornatus LeConte from Owen’s Valley, California, as noted under
“Notes about synonymy” section for C. placidus, although possessing a similar elytral scale
pattern to C. placidus individuals, is here considered conspecific with C. trivittatus. This
decision is based on the character states of the reduced extent of tarsal vestiture, presence of
slight postocular lobes, and short dorsal erect vestiture on the pronotum shared with C.
trivittatus and not C. placidus. Cleonus (Cleonidius) bicarinatus Casey is a typical, but
abraded C. trivittatus.
Problems in recognition.— Specimens of C. trivittatus are most likely to be confused with
those of C. quadrilineatus, C. placidus and C. boucardi. Discussions of characters allowing for
separation of the former are given under the “Problems in recognition” section for each of the
latter three species.
Description. — Specimens examined. 128 males, 159 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 31. Size. Length, male, 9.4-17.3 mm; female, 8.2-18.4 mm. Width, male, 3. 8-9. 4 mm; female, 3. 0-8. 2 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with punctures sparse to
moderately dense, small, shallow, some large and longitudinally confluent immediately above eyes; interspersed with dense
minute punctures. Frons also with sparse to moderately dense, elongate-narrow appressed to recumbent white scales
medially, erect immediately above eyes. Frons separated from base of rostrum by shallow to moderately deep transverse
impression (Fig. 153a). Base of median tumescence of rostrum with small, shallow fovea. Width of frons greater than to
subequal to width at apex of rostrum. Rostrum. Moderately robust, (width at apex 0.58-0.73 times length in male;
0.57-0.80 in female) (Fig. 153). In lateral view very slightly curved downward. In dorsal view with postgenae slightly
laterally expanded, antennal scrobes partially visible. Rostrum slightly to markedly medially tumescent from point of
antennal insertion to middle of frons (especially basally) (Fig. 153a); not laterally expanded at midlength; lateral margins
rounded; with median carina variously developed as narrow low glabrous shiny line at crest of median tumescence. Dorsal
and lateral punctation sparse to dense, small, shallow; dorsally interspersed with dense minute punctures. Dorsally either
lacking suberect or erect vestiture or with at most sparse to moderately dense, short, suberect hair-like scales; also with
moderately dense to dense, elongate-narrow, appressed to recumbent white scales laterally, scales sparser medially and
apically. Epistoma not to very slightly transversely swollen at base. Pronotum. Median carina variously developed in
anterior one-half from as minutely punctate, slightly elevated broad glabrous line to slightly elevated narrow glabrous
shiny line. Dorsal and lateral punctation small to moderately large, sparse to moderately dense, shallow to moderately
deep; smaller and shallower apically; sparser and shallower on flanks; areas between larger dorsal punctures with dense
Quaest. Ent., 1987, 23 (4)
538
Anderson
minute regularly impressed punctures. Lacking dorsal suberect or erect vestiture or with sparse to moderately dense, short,
suberect hair-like scales each situated in large puncture (Fig. 153a). Scales dorsally absent to sparse small and fine along
median line; moderately dense to dense, elongate-narrow, appressed in pair of narrow to broad apically slightly narrowed
paramedian stripes; laterally, scales absent to sparse, small and fine. Lateral margins with scales moderately dense to
dense, elongate-narrow, appressed in moderately broad stripe. Flanks with scales moderately dense to dense,
elongate-narrow. Median basal portion of disk shallowly to moderately deeply and broadly impressed. Anterolateral
margin with pronotal postocular lobes lacking (most specimens) to moderately developed (few specimens); postocular
vibrissae of unequal length, moderately long to very long, greatest length (from one-half to slightly greater than two-thirds
width of eye) behind base of eye (Fig. 153a). In dorsal view with lateral margins uniformly slightly arcuate to straight and
convergent from base to apical one-quarter to one-third, slightly constricted at apical one-quarter to one-third, then
straight and convergent to apex. Prosternum. With small shallow to moderately deep impression and lacking or with at
most only very slight rounded swelling anterior to each procoxal cavity. In lateral view with anterior flat to slightly
angulate portion very short to short, approximately one-half length of posterior more angulate portion or less; posterior and
anterior portions not to slightly differentiated by a shallow transverse impression. Elytra. Moderately robust in general
form (width at midlength 0.55-0.62 times length in males; 0.52-0.61 in females) (Fig. 134). In dorsal view with lateral
margins very slightly sinuate to moderately and evenly arcuate and slightly convergent from apical one-third to humerus;
moderately and evenly arcuate from apical one-third to apex. Elytral apices not to slightly produced, rounded to
subacuminate, not to slightly divergent. Humerus distinct. Dorsally lacking suberect or erect dorsal vestiture or with at
most sparse, short suberect hair-like scales on declivity; sutural interval with dense, small, fine to elongate-fine white
(rarely golden) scales; interval 2 with scales very sparse to dense small and fine; intervals 3 to 5 with scales dense large
elongate-narrow; intervals 6 to 8 with scales absent to moderately dense small and fine to elongate-fine; intervals 9 to 1 1
with scales dense, large elongate-narrow; striae 1, 6 and 7 with irregularly distributed patches of larger and denser white
scales (Fig. 134). Punctures of elytral striae small, individually distinct, arranged in regular rows. Wings. Long (greater
than elytra in length). Legs. Foretibia of female with inner margin with small to large denticles in apical three-quarters;
subapical tooth small to large, not to distinct from tibial denticles. Foretibial and mesotibial unci of both sexes moderately
large, metatibial uncus small. Tarsal claws connate in basal one-third, slightly divergent, each with basal internal flange
well-developed. Ventral tarsal pilose vestiture of foretarsus and mesotarsus of male present as small elongate pads on
apical one-third of article 1, as rounded pads on apical one-half of article 2, and as large rounded pads on apical
three-quarters of article 3; of foretarsus and mesotarsus of female as apical tufts to small elongate-narrow pads on apical
one-third of article 1, as rounded pads on apical one-quarter to one-half of article 2, and as large rounded pads on apical
one-half to two-thirds of article 3; of metatarsus of male as elongate-narrow pad on apical one-quarter on inner lobe and as
apical tuft on outer lobe of article 1, as elongate pads on apical one-half of article 2, as moderately large elongate-oval pads
on apical one-half to three-quarters of article 3; of metatarsus of female as apical tuft to elongate-narrow pad on apical
one-quarter on inner lobe and as elongate-narrow pad on apical one-half on outer lobe of article 1, as elongate pads on
apical one-quarter to three-quarters of article 2, as elongate-oval pads on apical one-third to one-half of article 3.
Abdomen. Ventral surface with moderately dense, elongate-narrow to moderately robust appressed white scales.
Abdominal sterna III to VI (especially sterna III and IV) with distinct rounded glabrous patches, each with large puncture
and single appressed to suberect moderately long hair-like scale situated in or near center. Base of abdominal sternum VII
of female internally (dorsally) with very slightly developed median shiny tubercle. Genitalia. Female (six examined).
Abdominal sternum VIII with basal arm short; lateral arms moderately inwardly arcuate throughout length; not to slightly
expanded at apex (Fig. 173). Stylus moderately large compared to length of gonocoxite II. Male (six examined). Internal
sac (Figs. 193a,b) elongate and low; median dorsal pocket low, with single moderately broad dorsally directed basal lobe
(lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B), with moderately large paired
dorsally directed lobe at basal one-third on dorsal surface (lobe C), with moderately large paired markedly dorsolaterally
directed lobe near midlength near dorsal margin (lobe D), with small paired dorsolaterally directed lobe just beyond
midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin (lobe F).
Apical pocket and median dorsal pocket individually distinct; apex of dorsal median pocket slightly emarginate medially in
dorsal view (Fig. 193b). Ventral median pocket large. Apical pocket with paired moderately large ventrally directed lobe
near apical one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent
ventral surface and basal portion of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic variation. — Individuals from the northern part of the species range (especially
Alberta, Canada) possess moderately developed postocular lobes and correspondingly shorter
postocular vibrissae. Throughout the rest of the species range, lobes are lacking or only slightly
developed and postocular vibrissae are longer.
In the western part of the species range in California and Nevada (especially the former),
individuals possess short suberect dorsal vestiture on the head and pronotum. Throughout the
rest of the species range, dorsal suberect or erect dorsal vestiture is lacking or at most very short
and indistinct.
New World Cleonini
539
Geographic distribution. — This species is widespread in the western United States of
America and Canada from extreme southern British Columbia south to southern California in
the west, and from southern Manitoba, south through eastern South Dakota, Kansas to western
Texas (Fig. 220).
Natural history. — This species appears to be associated primarily with grassland and
desert-grassland transitional habitats throughout the species range. Adults have been collected
exclusively on Leguminosae as follows; Astragalus spp. (as “loco” or “locoweed”), A.
bisulcatus (Hook.) Gray var. haydenianus, A. earlei Greene ex. Rydb., A. flavus Nutt., A.
utahensis Torr. and Gray, A. wootoni Sheldon, Oxytropis lambertii Pursh, and O. sericea
Nutt. Adults have been reared from Astragalus sp. (as “locoweed”). Adults have been collected
throughout the year at elevations of from 131-3640 m (N = 42) (Fig. 233).
A single specimen was found in the stomach contents of Bufo sp. (Amphibia).
Chorological relationships. — Table V.
Phylogenetic relationships. — Primary characters indicate only that this species is a
member of an unresolved quadrichotomy composed of the four species in the C. boucardi group
(Fig. 236). Distribution of the apotypic state of the secondary character of length of dorsal
vestiture place this species as the sister-species to the C. quadrilineatus-C. placidus lineage in
the C. boucardi group (Fig. 237).
Cleonidius placidus Csiki
(Figs. 135, 154, 174, 194, 222)
Cleonus pacificus Fall 1901:260 [not Olivier 1807:268]. Holotype (examined), male, labelled “Redondo/ Cal. 4. 7. 98”,
“TYPE/ pacificus ”, “M.C.Z./ Type/ 25194”, “H.C.FALL/ COLLECTION”, “Cleonus/ pacificus/ Fall” (MCZC).
Type locality, Redondo, California. Leng 1920 (catal.).
Cleonus ( Cleonidius ) placidus Csiki 1934:65. New name for Cleonus pacificus Fall. Blackwelder 1939 (check.).
Cleonis pacificus; O’Brien and Wibmer 1982 (catal., distn., as jr. homonym).
Cleonis placidus; O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — This species is commonly known as C. pacificus (Fall). This
name is a junior homonym and C. placidus Csiki is the valid name. The holotype of Cleonus
inornatus LeConte from Owen‘s Valley, California has an elytral scale pattern similar to that
of C. placidus (Fig. 135) but differs in other structural features (possession of short dorsal
suberect vestiture of the head and pronotum, slightly developed postocular lobes, and less
extensive ventral tarsal pilosity) that are found in Californian C. trivittatus. Cleonus inornatus
herein regarded as conspecific with C. trivittatus (see also “Notes about synonymy” section
under C. trivittatus).
Problems in recognition. — Most individuals of this species are easily recognized by the lack
of, or at most only very slightly developed postocular lobes (Fig. 154a), moderately long dorsal
erect vestiture of the head and pronotum (Fig. 154a), and distinctive elytral scale pattern (Fig.
135). From C. trivittatus , most individuals are separated by the elytral scale pattern and also
by the presence of moderately long dorsal erect vestiture on the head and pronotum in C.
placidus ; in C. trivittatus , dorsal erect or suberect vestiture is lacking or at most only suberect
and short (Fig. 153a). The two species are also largely allopatric in their distributions; C.
placidus is known only from California (Fig. 222), whereas C. trivittatus is widely distributed
in western North America, but known from only a few localities in California (Fig. 220). These
few C. trivittatus from California are difficult to separate from C. placidus , however, most of
them possess short erect dorsal vestiture, whereas this vestiture is slightly longer on the head
and pronotum of most C. placidus ; they also possess less extensive ventral tarsal pilosity,
Quaest. Ent., 1987, 23 (4)
540
Anderson
slightly developed postocular lobes, and lack the distinctive elytral scale pattern of C. placidus.
Both species are known only from Leguminosae.
Cleonidius placidus individuals may also be confused with individuals of C. quadrilineatus ,
but as for C. trivittatus specimens, the distinctive scale pattern of C. placidus should serve to
separate the two (Fig. 135). Otherwise, C. placidus is known only from Leguminosae and is
found in California, whereas C. quadrilineatus is known from Rosaceae and Rhamnaceae
(rarely from Leguminosae) and is widespread in southwestern North America (including
California). Most individuals of C. placidus are also slightly larger than are most C.
quadrilineatus (see Fig. 197).
Description. — Specimens examined. 57 males, 49 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 32. Size. Length, male, 9.9-17.1 mm; female, 12.6-17.6 mm. Width, male, 3. 7-6. 2 mm; female, 4. 3-6. 7 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with punctures sparse to
moderately dense, small to moderately large, shallow, some longitudinally confluent immediately above eyes; interspersed
with dense minute punctures. Frons also with moderately dense, moderately long, erect hair-like scales and sparse to dense,
elongate-narrow appressed white scales medially, erect immediately above eyes. Frons continuous with base of rostrum or
only slightly separated by shallow transverse impression (Fig. 154a). Base of median tumescence of rostrum with small,
shallow fovea. Width of frons greater than width at apex of rostrum. Rostrum. Moderately robust, (width at apex
0.59-0.65 times length in male; 0.57-0.62 in female) (Fig. 154). In lateral view very slightly curved downward. In dorsal
view with postgenae slightly laterally expanded, antennal scrobes visible. Rostrum slightly to moderately medially
tumescent from point of antennal insertion to middle of frons (Fig. 154a); not laterally expanded at midlength; lateral
margins rounded; with median carina variously developed as rounded to sharp, narrow low glabrous shiny line at crest of
median tumescence. Dorsal and lateral punctation sparse to moderately dense, small to moderately large, shallow to
moderately deep; punctures smaller and irregularly impressed apically, in some specimens dorsally interspersed with dense
minute punctures. Dorsally with moderately dense, moderately long to long, erect hair-like scales; also with sparse,
elongate-narrow, recumbent white scales laterally, scales sparser medially and apically. Epistoma not to very slightly
transversely swollen at base. Pronotum. Median carina variously developed in anterior one-half from as minutely punctate,
slightly elevated broad glabrous line to slightly elevated narrow glabrous shiny line, well-developed and distinct in most
specimens. Dorsal and lateral punctation moderately large, moderately dense to dense, moderately deep; smaller and
shallower apically; sparser, smaller and shallower on flanks; areas between larger dorsal punctures with dense minute
regularly impressed punctures. Dorsally, laterally and medially with moderately dense, moderately long to long, erect
hair-like scales each situated in large puncture (Fig. 154a). Scales dorsally absent along broad median line; moderately
dense to dense, elongate-narrow, appressed in pair of narrow apically convergent and slightly narrowed paramedian
stripes; laterally, scales absent. Lateral margins and flanks with scales moderately dense to dense, elongate-narrow.
Median basal portion of disk moderately deeply to deeply and broadly impressed. Anterolateral margin with pronotal
postocular lobes lacking (most specimens) to only very slightly developed (few specimens); postocular vibrissae of unequal
length, very long, greatest length (slightly greater than two-thirds width of eye) behind base of eye (Fig. 154a). In dorsal
view with lateral margins uniformly very slightly arcuate and convergent from base to apical one-quarter, slightly
constricted at apical one-quarter, then straight and convergent to apex. Prosternum. With small shallow to moderately
deep impression and lacking or with at most only very slight rounded swelling anterior to each procoxal cavity. In lateral
view with anterior flat to slightly angulate portion very short, indistinct. Elytra. Moderately elongate-narrow in general
form (width at midlength 0.51-0.56 times length in males; 0.50-0.57 in females) (Fig. 135). In dorsal view with lateral
margins straight to very slightly sinuate and slightly convergent from apical one-third to humerus; moderately and evenly
arcuate from apical one-third to apex. Elytral apices not to slightly produced, rounded to subacuminate, not to slightly
divergent. Humerus distinct. Dorsally with sparse, short suberect hair-like scales (most evident on declivity); sutural
interval with sparse to moderately dense, small fine white scales; interval 2 with irregularly distributed patches of dense,
elongate-fine white scales, otherwise with scales absent to sparse, small and fine; intervals 3 to 1 1 with scales uniformly
moderately dense to dense, elongate-fine to elongate-narrow (some specimens with irregularly distributed patches of even
larger denser, white scales, especially on intervals 9 to 11) (Fig. 135). Punctures of elytral striae small, individually
distinct, arranged in regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner
margin with small denticles in apical three-quarters; subapical tooth not distinct from tibial denticles. Foretibial and
mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-third, slightly
divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus and mesotarsus of
male present as elongate pads on apical two-thirds of article 1, as large rounded pads on apical two-thirds of article 2, and
as large rounded pads on more or less entire ventral surface of article 3; of foretarsus of female as small elongate pads on
apical one-third of article 1, as rounded pads on apical one-half of article 2, as large rounded pads on apical three-quarters
of article 3; of mesotarsus of female as small elongate pads on apical one-quarter to one-third of article 1, as rounded pads
on apical one-half of article 2, as large rounded pads on apical two-thirds of article 3; of metatarsus of male as elongate
pads on apical one-half to two-thirds of article 1, as large rounded pads on apical two-thirds of article 2, as large rounded
New World Cleonini
541
pads on more or less entire ventral surface of article 3; of metatarsus of female as elongate-narrow pads on apical
one-quarter to two-thirds of article 1, as elongate pads on apical one-half to two-thirds of article 2, as large rounded pads
on apical two-thirds of article 3. Abdomen. Ventral surface with moderately dense to dense, elongate-narrow to moderately
robust appressed white scales; scales sparser along midline on sternum VII of female. Abdominal sterna III to VI
(especially sterna III and IV) with distinct large rounded glabrous patches, each with large puncture and single recumbent
to suberect moderately long hair-like scale situated in or near center. Base of abdominal sternum VII of female internally
(dorsally) with very slightly developed median shiny tubercle. Genitalia. Female (two examined). Abdominal sternum
VIII with basal arm moderate in length; lateral arms moderately inwardly arcuate at midlength; slightly expanded at apex
(Fig. 174). Stylus moderately large compared to length of gonocoxite II. Male (two examined). Internal sac (Figs. 194a, b)
elongate and low; median dorsal pocket low, with single moderately broad dorsally directed basal lobe (lobe A), with small
median dorsally directed lobe at midlength on dorsal surface (lobe B), with moderately large paired dorsally directed lobe
at basal one-third on dorsal surface (lobe C), with moderately large paired dorsolaterally directed lobe near midlength
near dorsal margin (lobe D), with moderately large paired dorsolaterally directed lobe just beyond midlength at dorsal
margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin (lobe F). Apical pocket and
median dorsal pocket individually distinct; apex of dorsal median pocket bisinuate medially in dorsal view (Fig. 194b).
Ventral median pocket moderately large. Apical pocket with paired moderately large ventrally directed lobe near apical
one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral
surface and basal portion of sides of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — This species is known only from California in the coastal and
central valleys from San Francisco south to San Diego (Fig. 222).
Natural history. — This species is found only in the Pacific semi-desert region. Adults have
been collected almost exclusively from Astragalus spp. (Leguminosae). A single specimen was
collected on parsnip, Pastinaca sativa L. (Umbelliferae). Adults have also been reared from
roots of Astragalus sp. Adults have been collected throughout the year at elevations of from
9-1747 m(N= 10) (Fig. 233).
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. quadrilineatus, the two
species forming a lineage that is the sister group of C. trivittatus (Fig. 237).
Cleonidius quadrilineatus (Chevrolat)
(Figs. 136, 155, 175, 195, 224)
Apleurus quadrilineatus Chevrolat 1873:80. Holotype (examined), male, labelled “66”, “ex coll./ Salle”, “MUSEUM
PARIS/ 1952/ COLL R OBERTHUR”, ''apleurus/ quadrilineatus/ Chev. type texas”, and with an inverted label
“ Lixus / praepotens/ Boh. Texas”, and with a label indicating it as the holotype “ Apleurus / quadrilineatus / Chev.
HOLOTYPE/ labelled Anderson” (MNHP). Type locality, Texas.
Cleonus quadrilineatus-, LeConte 1876a (key, diag.). Henshaw 1885 (check.). Wickham 1889. Wickham 1896 (check.).
Fall 1897. Fall 1901. Wickham 1902 (check.). Fall and Cockerell 1907 (check.). Pierce 1907 (biol., misident. of
Cleonidius trivittatus). Chittenden 1911 (biol., misident. of Cleonidius trivittatus). Anderson 1914 (misident. of
Cleonidius poricollis). Blatchley and Leng 1916 (figure, misident. of Cleonidius trivittatus). Yothers 1916 (prob.
misident. of Cleonidius trivittatus, in part). Leng 1920 (catal.). Essig 1958 (biol., misident. of Cleonidius trivittatus).
Arnett et al. 1980 (key, redesc., biol., misident. of Cleonidius trivittatus).
Lixus texanus\ LeConte 1876a (in part, mixed type series).
Cleonus ( Cleonidius ) quadrilineatus-, Casey 1891 (key). Csiki 1934 (catal.). Hatch 1971 (key, redesc., misident. of
Cleonidius poricollis).
Cleonis quadrilineatus-, O’Brien and Wibmer 1982 (catal., distn.).
Notes about synonymy. — The holotype of C. quadrilineatus is the typical elongate-narrow
Rosaceae form. Many published references to C. trivittatus are to this species.
Problems in recognition. — Specimens of C. quadrilineatus are very likely to be commonly
confused with those of the broadly sympatric C. trivittatus. Cleonidius quadrilineatus
specimens are separated from the latter by their more elongate-narrow elytral form (Fig. 136)
and by presence of moderately long erect hair-like scales on the pronotal disk (Fig. 155a),
which in C. trivittatus are at most only short, suberect and indistinct (Fig. 153a). Most C.
quadrilineatus are also associated with Rosaceae or Rhamnaceae, although rarely also with
Quaest. Ent., 1987,23 (4)
542
Anderson
Leguminosae, whereas all C. trivittatus are associated with Leguminosae. Some C. trivittatus
have slightly to moderately developed postocular lobes (especially from the northern part of the
range) whereas all C. quadrilineatus lack postocular lobes entirely (Fig. 155a).
In California, specimens of C. quadrilineatus may be confused with C. placidus, especially
specimens of the latter in which the distinctive elytral scale pattern (Fig. 135) is abraded.
Individuals of C. placidus have moderately long, erect hair-like scales on the pronotum (Fig.
154a), as do C. quadrilineatus , but are associated exclusively with Leguminosae. C. placidus
individuals are also slightly larger than are those of C. quadrilineatus (Fig. 197) and have
slightly more extensive ventral tarsal pilosity.
Description. — Specimens examined. 108 males, 107 females. Data about variation in LR, WF, WRA, LP, WPB,
WE1H, WE1M, LEI, LEy, WEy, WEy/LEy, WPB/LP, WRA/LR, WRA/WF, WE1M/LE1, and LP/LE1 are presented
in Table 33. Size. Length, male, 8.3-14.3 mm; female, 9.3-13.1 mm. Width, male, 2.9-5. 3 mm; female, 3. 2-4. 8 mm.
Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with punctures sparse to
moderately dense, small to moderately large, shallow, some large and longitudinally confluent immediately above eyes;
interspersed with minute punctures. Frons also with sparse to moderately dense, elongate-narrow appressed white scales
medially, erect immediately above eyes. Frons continuous with base of rostrum or separated from base of rostrum by
shallow transverse impression (Fig. 155a). Base of median tumescence of rostrum with small, shallow fovea. Width of
frons greater than to subequal to width at apex of rostrum. Rostrum. Moderately robust, (width at apex 0.60-0.70 times
length in male; 0.58-0.69 in female) (Fig. 155). In lateral view very slightly curved downward. In dorsal view with
postgenae slightly laterally expanded, antennal scrobes partially visible. Rostrum moderately medially tumescent from
point of antennal insertion to middle of frons (especially basally) (Fig. 155a); not laterally expanded at midlength; lateral
margins rounded; with median carina variously developed as narrow low glabrous shiny line at crest of median tumescence.
Dorsal and lateral punctation sparse to dense, small to moderately large, shallow; punctures smaller apically; dorsally
interspersed with dense minute punctures. Dorsally with moderately dense, moderately long, suberect to erect, hair-like
scales (especially laterally); also with moderately dense, elongate-narrow, recumbent white scales laterally, scales absent to
sparse medially and apically. Epistoma not to very slightly transversely swollen at base. Pronotum. Median carina
variously developed in anterior one-half from as minutely punctate, slightly elevated broad glabrous line to slightly
elevated narrow glabrous shiny distinct line. Dorsal and lateral punctation moderately large, moderately dense, moderately
deep; smaller and shallower apically; sparser and shallower on flanks; areas between larger dorsal punctures with dense
minute regularly impressed punctures. Dorsally, laterally and medially, with moderately dense, moderately long, erect
hair-like scales each situated in large puncture (Fig. 155a). Scales dorsally absent to sparse small and fine along median
line; moderately dense, elongate-fine to elongate-narrow, appressed in pair of narrow apically slightly narrowed
paramedian stripes; laterally, scales absent to sparse, small and fine. Lateral margins with scales moderately dense to
dense, elongate-narrow to moderately robust, appressed in moderately broad stripe. Flanks with scales moderately dense,
elongate-fine to elongate-narrow. Median basal portion of disk shallowly to moderately deeply and broadly impressed.
Anterolateral margin with pronotal postocular lobes lacking; postocular vibrissae of unequal length, very long, greatest
length (greater than two-thirds width of eye) behind base of eye (Fig. 155a). In dorsal view with lateral margins uniformly
slightly arcuate and convergent from base to apex, to straight and convergent from base to apical one-quarter to one-third,
slightly constricted at apical one-quarter to one-third, then straight and convergent to apex. Prosternum. With small
shallow to moderately deep impression and slight rounded swelling anterior to each procoxal cavity. In lateral view with
anterior flat to slightly angulate portion very short, indistinct. Elytra. Elongate-narrow in general form (width at
midlength 0.48-0.55 times length in males; 0.48-0.52 in females) (Fig. 136). In dorsal view with lateral margins straight to
very slightly sinuate slightly convergent from apical one-third to humerus; moderately and evenly arcuate from apical
one-third to apex. Elytral apices not to slightly produced, rounded to subacuminate, not to slightly divergent. Humerus
distinct. Dorsally with sparse, short suberect hair-like scales (most evident on declivity); sutural interval with dense, small
fine golden to white scales; interval 2 with scales very sparse to dense small and fine, white; intervals 3 to 5 with scales
dense large elongate-narrow, white; intervals 6 to 8 with scales sparse to moderately dense, small and fine to elongate-fine,
white; intervals 9 to 11 with scales dense, large elongate-narrow, white; striae 1, 6 and 7 with irregularly distributed
patches of larger and denser white scales (Fig. 136). Punctures of elytral striae small, individually distinct, arranged in
regular rows. Wings. Long (greater than elytra in length). Legs. Foretibia of female with inner margin with small to
moderately large denticles in apical three-quarters; subapical tooth indistinct from tibial denticles. Foretibial and
mesotibial unci of both sexes moderately large, metatibial uncus small. Tarsal claws connate in basal one-third, slightly
divergent, each with basal internal flange well-developed. Ventral tarsal pilose vestiture of foretarsus and mesotarsus of
male present as small elongate pads on apical one-half of article 1, as large, elongate-oval pads on apical two-thirds of
article 2, and as large rounded pads on apical three-quarters of article 3; of foretarsus and mesotarsus of female as as small
elongate-narrow pads on apical one-quarter of article 1, as small rounded pads on apical one-third of article 2, and as
moderately large rounded pads on apical one-half to two-thirds of article 3; of metatarsus of male as elongate-narrow pad
on apical one-half on inner lobe and apical one-third on outer lobe of article 1, as elongate pads on apical one-half to
two-thirds of article 2, as large rounded pads on apical three-quarters of article 3; of metatarsus of female as
New World Cleonini
543
elongate-very-narrow pad on apical one-half on inner lobe and apical one-third on outer lobe of article 1, as elongate pads
on apical one-third to one-half of article 2, as moderately large elongate-oval pads on apical one-half to two-thirds of
article 3. Abdomen. Ventral surface with moderately dense, elongate-narrow to moderately robust appressed white scales.
Abdominal sterna III to VI (especially sterna III and IV) with distinct rounded glabrous patches, each with large puncture
and single appressed to erect moderately long hair-like scale situated in or near center. Base of abdominal sternum VII of
female internally (dorsally) with slightly to moderately developed median shiny tubercle. Genitalia. Female (four
examined). Abdominal sternum VIII with basal arm short; lateral arms moderately inwardly arcuate throughout length;
not to slightly expanded at apex (Fig. 175). Stylus moderately large compared to length of gonocoxite II. Male (four
examined). Internal sac (Figs. 195a,b) elongate and low; median dorsal pocket low, with single moderately broad dorsally
directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B), with
moderately large paired dorsally directed lobe at basal one-third on dorsal surface (lobe C), with moderately large paired
markedly dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with small paired dorsolaterally
directed lobe just beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near
ventral margin (lobe F). Apical pocket and median dorsal pocket individually distinct; apex of dorsal median pocket
slightly emarginate medially in dorsal view (Fig. 195b). Ventral median pocket large. Apical pocket with paired
moderately large ventrally directed lobe near apical one-third at ventral margin. Eversible apical sclerite complex with
paired simple scythe-like sclerite; adjacent ventral surface and basal portion of sides of apical pocket with pair of large,
dark, longitudinal sclerites.
Geographic distribution .— - This species is found from California east to eastern Nebraska,
Oklahoma, and central Texas (Fig. 224). There is a single Mexican record from near
Zacatecas.
Natural history. — This species is found primarily in the western xerophytic evergreen
forest region but also in desert-grassland transitional habitats. Many adults have been collected
primarily on Rosaceae or Rhamnaceae, especially in oak-pinyon-juniper woodlands throughout
the species range; a few have been found on Leguminosae in Arizona. Adults have been
collected on Astragalus wootoni Sheldon, Lupinus sp. (Leguminosae); Ceanothus sp.
(Rhamnaceae); Cercocarpus montanus Raf., Rosa sp. (Rosaceae). Adults have been reared
only from Astragalus sp. roots at Sulphur Springs Valley, Arizona. I suspect that a variety of
Rosaceae or Rhamnaceae serve as hosts at higher altitudes throughout the species range.
Adults have been collected from January to October at elevations from 15-4186 m (N = 63)
(Fig. 233).
Adults have been found in the stomach contents of Geococcyx californicus' (roadrunner;
Aves) at Carlsbad, New Mexico.
Other individuals and 1 have collected this species and C. texanus together on various
Rosaceae and Rhamnaceae in Texas and Arizona.
Chorological relationships. — Table V.
Phylogenetic relationships. — This species is the sister-species of C. placidus, the two
species forming a lineage that is the sister group of C. trivittatus (Fig. 237).
Incertae sedis
This category is used herein for a single species of uncertain placement, Cleonidius vibex
(Pallas). Although adults of C. vibex have a vittate elytral scale pattern, lack extensive ventral
tarsal pilose pads, possess a moderately broad rostrum, and are brachypterous, all of which
suggest inclusion in the C. americanus group, the nature of formation of the vittate scale
pattern is fundamentally different from that in other Cleonidius and thus not homologous. In
other Cleonidius the vittate pattern is produced as a result of differing sizes and densities of
scales on the various elytral intervals; in C. vibex, the pattern is apparently produced as a result
of presence or absence of white pigment in the individual scales. This suggests independent
derivation of a vittate elytal pattern and perhaps of states of other characters as well.
Quaest. Ent., 1987, 23 (4)
544
Anderson
Placement of C. vibex as sister to the remainder of Cleonidius can only be substantiated by
an adaptive argument concerning the low likelihood of evolution of the fundamentally different
vittate elytral scale pattern of that species from the scale pattern of other Cleonidius , which is
formed in a very different non-homologous manner.
Cleonidius vibex (Pallas), new combination
(Figs. 156, 176, 196)
Curculio vibex Pallas 1781:32. Types not examined. Herbst 1795.
Lixus denudatus Zoubkoff 1833:319. Types not examined. Faust 1890. Petri 1912.
Lixus karelini Boheman 1836:22. Types not examined. Faust 1890. Petri 1912.
Lixus vibex ; Boheman 1836. Capiomont and Leprieur 1874. Faust 1890. Petri 1912. Lukjanovitsh 1926.
Lixus vibex scutellaris Petri 1905:103. Types not examined.
Lixus (Lixestus) vibex\ Reitter 1916. Csiki 1934.
Lixus (Lixesthus) vibex ; Ter-Minasyan 1978.
Notes about synonymy. — No types associated with the names recorded above have been
examined by me. Synonymy presented here follows Petri (1912) and Csiki (1934).
Two additional species have been placed in Lixus ( Lixestus ) by Ter-Minasyan (1978).
These are L. pallasi Faust and L. meles Boheman. These species will likely also prove to be
Cleonidius but representatives have not been examined by me.
Problems in recognition. — Individuals of this species are easily distinguished from Nearctic
Cleonidius by the widely divergent tarsal claws each lacking a basal internal flange, and the
large unci especially on the fore- and mesotibiae. The rostrum of this species has uniformly very
dense and small punctures, but no or else very few larger punctures typical of Nearctic species
of Cleonidius. Individuals also differ from those of Nearctic species in that the contrasting dark
and light areas of the pronotum and elytra are a result of apparently having white scales, such
that the underlying dark cuticle is obscured, or unpigmented, such that the dark cuticle is not
obscured. Individuals of Nearctic species on the other hand, have a vittate elytral scale pattern
and a pronotal scale pattern that is a result of differing sizes of scales; small scales do not
obscure the dark cuticle, whereas larger scales do.
The robust, downwardly curved rostrum and elytral scale pattern of individuals of this
species result in superficial resemblance to C. canescens.
Description. — Specimens examined. 1 male, 1 female. Size. Length, male, 7.3 mm; female, 7.1 mm. Width, male,
2.7 mm; female, 2.6 mm. Head. Eye elongate-oval. Area behind eye with numerous slight wrinkles of cuticle. Frons with
only uniformly very dense minute punctures. Frons also with very dense, truncate grey-brown appressed scales. Frons
continuous with base of rostrum (Fig. 156a). Base of median tumescence or carina of rostrum (if present) with small,
shallow fovea. Width of frons greater than width at apex of rostrum. Rostrum. Robust, (width at apex 0.73 times length in
male; 0.75 in female) (Fig. 156). In lateral view moderately curved downward. In dorsal view with postgenae very slightly
laterally expanded, antennal scrobes distinctly visible. Rostrum markedly medially tumescent from point of antennal
insertion to middle of frons (Fig. 156a); not laterally expanded at midlength; lateral margins more or less rounded, not
sharp; with median carina absent. Dorsal and lateral punctation very sparse, small, shallow; sparser apically and medially;
dorsally interspersed with minute punctures. Dorsally lacking suberect or erect vestiture, with only uniformly very dense,
appressed truncate grey-brown scales (punctation almost completely obscured), sparser apically and medially. Epistoma
not swollen. Pronotum. Median carina lacking. Dorsal and lateral punctation sparse, small, shallow, interspersed with very
dense minute regularly impressed punctures. Suberect or erect vestiture lacking. Scales dorsally uniformly dense small
truncate appressed medially; laterally transparent in pair of narrow stripes (apparently black). Lateral margins and flanks
with scales uniformly dense, small, truncate and appressed, punctation almost completely obscured. Median basal area of
disk broadly and very shallowly impressed. Anterolateral margin with pronotal postocular lobes well-developed; postocular
vibrissae short, of uniform length (Fig. 156a). In dorsal view with lateral margins straight and convergent from apical
one-third to base; slightly arcuate from apical one-third to apex; subapical constriction indistinct. Prosternum. With
moderately large, moderately deep impression anterior to each procoxal cavity; swelling lacking. In lateral view with
anterior flat to slightly angulate portion moderately long, approximately one-half length of posterior more angulate
portion; anterior and posterior portions slightly differentiated by shallow transverse impression. Elytra. Moderately
New World Cleonini
545
elongate-narrow in general form (width at midlength 0.52 times length in male; 0.53 in female). In dorsal view with lateral
margins very slightly sinuate and slightly convergent from apical one-third to humerus; moderately arcuate at apical
one-third then straight and convergent to apex. Elytral apices not produced, rounded, slightly divergent. Humerus
rounded, indistinct. Dorsally lacking suberect or erect vestiture. Scales uniformly very dense, small, robust, appressed
grey-brown on sutural interval and intervals 2 to 5 and 8 to 11; intervals 6 and 7 with scales truncate, transparent,
underlying dark cuticle visible through scales. Punctures of elytral striae small, individually distinct; arranged in regular
rows. Wings. Moderate (slightly shorter than length of elytra). Legs. Foretibia of female with inner margin with small
denticles in apical one-half; no distinct subapical tooth present. Foretibial and mesotibial unci of both sexes very large,
metatibial uncus moderately large. Tarsal claws widely divergent, each lacking distinct basal internal flange. Ventral
tarsal pilose vestiture of foretarsus and mesotarsus of both sexes lacking from articles 1 and 2, present as small rounded
apical tuft of article 3; of metatarsus of both sexes, lacking from articles 1 and 2, present as minute apical tuft of article 3.
Abdomen. Ventral surface with dense, truncate, small appressed grey-brown scales. Abdominal sterna III to VI sparse
moderately large shallow punctures, no distinct glabrous patches or suberect or erect vestiture present. Base of abdominal
sternum VII of female internally (dorsally) with at most moderately-developed median shiny tubercle. Genitalia. Female
(one examined). Abdominal sternum VIII with basal arm long; lateral arms straight and divergent from base to
approximately apical one-third; moderately inwardly arcuate at apical one-third and slightly convergent to apex; apices
moderately expanded (Fig. 176). Stylus moderately large compared to length of gonocoxite II. Male (one examined).
Internal sac (Figs. 196a,b) elongate and low; median dorsal pocket low, with single moderately broad elongate dorsally
directed basal lobe (lobe A), with small median dorsally directed lobe at midlength on dorsal surface (lobe B), with
moderately large paired dorsally directed lobe at basal one-third at dorsal margin (lobe C), with moderately large paired
dorsolaterally directed lobe near midlength near dorsal margin (lobe D), with large paired dorsolaterally directed lobe just
beyond midlength at dorsal margin (lobe E), and with small ventrally directed paired lobe near base near ventral margin
(lobe F). Apical pocket individually distinct from median dorsal pocket; apex of median dorsal pocket rounded in dorsal
view. Ventral median pocket large. Apical pocket with paired moderately large ventrally directed lobe near apical
one-third at ventral margin. Eversible apical sclerite complex with paired simple scythe-like sclerite; adjacent ventral
surface of apical pocket with pair of large, dark, longitudinal sclerites.
Geographic distribution. — Southern Russia, Persia, Siberia and Mongolia (Csiki 1934).
Natural history. — No information is available about the natural history of this species.
Based on its general geographic distribution I suspect that it is a grassland or semi-desert
inhabiting species.
Chorological relationships. — This is the only Palearctic species of Cleonidius I have seen.
Phylogenetic relationships. — See “ Incertae sedis ” section preceding.
Nomen dubium
Lixus californicus Motschulsky is herein treated as a nomen dubium. Motschulsky
(1845:378) in his original description of this species noted only that its type was smaller, more
elongate, and whiter than specimens of either Lixus poricollis Mannerheim and L. modestus
Mannerheim. He further stated that he thought it likely that it was a male of L. modestus.
Based upon examination of the holotype (on loan from Zoological Museum of the Moscow
Lomonosov State University) Lixus modestus Mannerheim is, however, not a Cleonidius nor is
it conspecific with any North American Lixus species known to me. Furthermore, L. modestus
does not possess character states of the New World Lixus species component and thus I suspect
has been erroneously labelled as having been collected in California. This notion of a
misidentified type locality is supported by Mannerheim (1843: 291), who stated that
Eschscholtz (in litteris ) had referred to L. modestus or Lixus adspersus (a species described by
Boheman (1836) from the Republic of South Africa).
In view of this confusion regarding the type locality of Lixus modestus Mannerheim, the
statement by Motschulsky (1845) that he suspects L. californicus Motschulsky is a male of L.
modestus (the type of L. modestus incidentally is a male), and the inadequate characterization
(and the apparent loss of type material) of L. californicus, I consider recognition of Lixus
californicus Motschulsky as a nomen dubium to be in the best interests of a stable
nomenclature.
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ADVENTITIOUS TAXA
During the course of study, individuals of two adventitious species were encountered from
North American localities. These are Stephanocleonus glaucus labelled “S.C.” (1), and
“Ames, Iowa” (1); and, an unidentified species of Tetragonothorax from “Mobile, Alabama”
(1). These species are considered not to be established in North America.
EXTANT TAXA ERRONEOUSLY PLACED IN CLEONINAE
Cleonis chilensis Blanchard 1851:326 is listed in both Cleonus (Incertae Sedis) and
Adioristus in the Coleopterorum Catalogus (Csiki 1934; Schenkling and Marshall 1931) and
also in Blackwelder (1947). Kuschel (1949:31) subsequently placed it in Scotoeborus
(Leptopiinae [now Entiminae], Cylydrorhinini). I have examined the type of this species
(deposited in MHNP) and agree that it is not cleonine.
Argentinorhynchus breyeri Brethes 1910:211 was described from Argentina and originally
noted as “Cleonidarum”. It has since been regarded as cleonine by (Csiki 1934) who placed the
genus as a subgenus of Cleonus. Likewise, and probably following Csiki, Blackwelder (1947)
lists the species in Cleonus. Kuschel (1950:112) subsequently returned Argentinorhynchus to
generic status and placed it in Erirhininae. The holotype of this species is located in the Museo
Argentino de Ciencias Naturales “Bernardino Rivadavia” but has not been examined by me
(Bachman in lift., 1985).
The genus Lepyrus is also not Cleoninae and has been removed to Molytinae (Hylobiinae of
authors) (Solari 1941; Ter-Minasyan 1963).
FOSSIL MATERIAL
Various Tertiary coleopterous fossils have been identified as Cleoninae but, preservation is
such that most of these are actually not reliably identifiable to subfamily, according to
Kingsolver (1961) who has examined much of the material of Scudder (1893) and Wickham
(1911, 1912) deposited at the Museum of Comparative Zoology, Harvard University.
Matthews (1977) also notes a Cleonus sp. from the Tertiary of Banks Island in the Canadian
Arctic but this identification is based only on elytral fragments. I have not examined any of
these specimens.
Sleeper (1969) described two species of Cleonidius (as Cleonus ) from Pleistocene Rita
Blanca Lake deposits in Texas. Unfortunately I have not seen the specimens, and descriptions
and figures do not provide information permitting meaningful comparison with extant species.
Sleeper (1969) states that Cleonidius ritablancensis is near Cleonidius sparsus and that
Cleonidius channingensis is near Cleonidius circumductus , but, the history of misidentification
in Cleonidius and subtle differences between many of the species necessitates re-examination.
Unfortunately location of the specimen depository was not stated by Sleeper (1969).
Fossils of species identified as the genus Cleonus are frequently encountered in late
Pleistocene deposits in northern North America (Matthews, 1974, 1975; Hughes et al ., 1981;
Mott et al ., 1981; Morlan and Matthews, 1983). Some of these have been tentatively identified
as Cleonus plumbeus, but as noted by Morgan et al. (1985), they may actually represent one or
more Palearctic species or even one of the herein-newly-described Nearctic species. All of this
northern Pleistocene material actually belongs in the genus Stephanocleonus , and the present
New World Cleonini
547
revision of North American members of that genus reveals that taxa traditionally regarded as
S. plumbeus are a complex of six species, four of which are newly described herein. All of this
fossil material is deposited with John V. Matthews, Jr., Geological Survey of Canada, Ottawa;
none of it has been examined by me.
EVOLUTIONARY ANALYSIS: RECONSTRUCTED PHYLOGENY
My profession embodies one theme even more inclusive than evolution— the nature and meaning of history. History
employs evolution to structure biological events in time. History subverts the stereotype of science as a precise,
heartless enterprise that strips the uniqueness from any complexity and reduces everything to timeless, repeatable,
controlled experiments in a laboratory. Historical sciences are different, not lesser. Their methods are comparative,
not always experimental; they explain, but usually do not try to predict; they recognize the irreducible quirkiness
that history entails, and acknowledge the limited power of present circumstances to impose or elicit optimal
solutions.
Steven Jay Gould, Prologue to “The Flamingo's Smile”, 1985.
Inquiry in historical science proceeds primarily by investigation of relationships between
pattern and process. Factual data are assembled and patterns are searched for and recognized
in these data. Recognition of a pattern, especially if repeated within the study group or in other
groups, begs explanation by proposal of a common causal mechanism in the form of a testable
hypothesis. Proposal and subsequent testing of this hypothesis, the former inductive, the latter
hypothetico-deductive, are the cornerstones of science in general and systematic research in
particular. Comparison of these approaches in systematics is adequately treated by Wiley
(1981) and application of the hypothetico-deductive method to systematics by Gaffney (1979).
In this study, aspects of both inductive and hypothetico-deductive science are employed and
applied. Although some recent authors advocate only the hypothetico-deductive approach, I
believe that much useful information and advances in biological knowledge have been and
should continue to be, achieved through use of both approaches, but that they operate best at
different stages in the process of scientific inquiry; induction for hypothesis formulation,
deduction for hypothesis testing.
Herein, an inductive approach is used primarily to attempt to formulate explanations of
details of the evolutionary and biogeographic history of the weevil subfamily Cleoninae,
especially species of the genera Cleonidius and Apleurus in North America. From these
analyses, evident patterns are identified and more general hypotheses concerning biogeographic
patterns and evolutionary trends in arid-land-adapted organisms are proposed, to be available
for subsequent testing in future studies for their applicability and predictive value. In addition,
some competing and previously proposed hypotheses concerning general evolutionary or
biogeographic patterns are tested by application of the appropriate results of this study of
North American Cleoninae.
Methods of phylogeny reconstruction
Since the publication of Hennig (1966), there has been an expanding emphasis on both
theoretical and practical aspects of phylogenetic and biogeographic studies. While the pages of
Systematic Zoology clearly document an intense interest in theoretical aspects, increased
attention to practical aspects and applications is evidenced simply by the numbers of
revisionary works that deal extensively with reconstruction of phylogeny and its applicability to
character evolution, classification and biogeography. The appeal of the phylogenetic approach
(Hennig, 1966) to phylogeny reconstruction over various other approaches (Wiley, 1981) is
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primarily in the potential for critical assessment of resultant hypotheses of relationship, their
logical predictive nature and their general applicability. The approach I have taken herein is
phylogenetic (Hennig, 1966). Character states are determined as apotypic (derived) or
plesiotypic (primitive) and only distributions of apotypic states are used in the analysis.
Of paramount importance in reconstructing phylogeny are: one, determination of polarity of
character states; and two, recognition of homoplasy or simply, determination of which
characters actually reflect descent from a common ancestor, not other processes.
Out-group criteria were used to polarize character states of all primary characters (Watrous
and Wheeler, 1981). For secondary characters, out-group criteria were employed where
possible although in-group or local parsimony (Maddison et al., 1984), or the functional
out-group of Watrous and Wheeler (1981), were also used, especially in instances where more
than one character state occurs in the out-groups or where, although both out-groups have the
same state, there is incompatibility with the reconstructed phylogeny as determined by primary
characters.
Character weighting was employed. Characters used in the phylogenetic analysis of species
in Cleonidius and Apleurus are considered at two levels as indicated above. Primary characters
are those which are deemed of high weight because there is no ambiguity in determining
polarity of the various character states based on an out-group analysis (Watrous and Wheeler,
1981; Maddison et al ., 1984), and the apotypic state, so determined, is not widely distributed
elsewhere in Cleoninae. Characters meeting these criteria are given further support as primary
characters should they, as a group, result in a reconstructed phylogeny of the genus in question
in which the distributions of apotypic states of the various characters are compatible, with the
result that there is minimal homoplasy. Characters that exhibit more than a little homoplasy
elsewhere in Cleoninae may be regarded as primary should the distribution of their apotypic
states be entirely compatible with the distribution of apotypic states of numerous other primary
characters.
Secondary characters are those considered to be of lesser weight because of widespread
presence of the apotypic state elsewhere in Cleoninae. Lack of complete compatibility with
primary characters, and/or problems in determining polarity of the character states based
strictly on an out-group analysis necessitating reference to in-group relationships previously
resolved on the basis of the distribution of states of primary characters also results in such
characters being considered at a secondary level. Distribution of apotypic states of secondary
characters may or may not be compatible with distributions of apotypic states of primary or
other secondary characters with the result that there is generally more homoplasy in characters
included in this more resolved reconstructed phylogeny of the genus is question.
Characters examined but not included in the phylogenetic analysis are those that exhibit
extensive homoplasy elsewhere in Cleoninae and are not compatible with, or, when compared
with other potential secondary characters, exhibit minimal compatibility with, distributions of
apotypic states of primary characters.
Primary characters are the characters used to establish a basic set of relationships that may
or may not be totally dichotomously resolved, but in which confidence can generally be
regarded as high. This set of relationships is then accepted a priori and the distribution of states
of secondary characters considered with reference to this reconstructed phylogeny. Thus
secondary characters can only be used to reinforce previously established relationships or to
further resolve the reconstructed phylogeny by providing additional characters in which
confidence is low.
New World Cleonini
549
I do not use a complex system of weighting characters such as that of Hecht and Edwards
(1976, 1977). Instead, I use this simple two level weighting system which depends solely upon
the distributions of character states, and makes no inferences or assumptions about complexity,
function or adaptive value which may not have a logically sound basis. As such, I think this
system better reflects the confidence level for a given set of relationships especially when these
are depicted on successively more resolved cladograms.
Phylogenetic position of Cleoninae
The detailed phylogenetic position of Cleoninae is unclear although relationships at some
level appear to be with Molytinae, specifically the Hylobiinae, of authors. Taxonomy of these
two subfamilies has a history of repeated misclassification and a repeated concern for adequate
distinction (Aslam, 1963; Marshall, 1932; Solari, 1941; Ter-Minasyan, 1963). In addition,
Aslam (1961) has noted that among Curculionidae, the Molytinae and Cleoninae are unique in
their possession of free-lobed testes, and, along with a few other subfamilies, both have the vasa
efferentia free from the testicular lobes and each of the latter with numerous follicles. These
states however, all appear to be symplesiotypic for they are all found in primitive
Curculionoidea (Morimoto, 1962) and cannot be considered evidence for a sister-group
relationship but rather, as will be noted forthwith, for a primitive position for Cleoninae with
respect to other Curculionidae. Although unjustified by synapotypy as sister-taxon to
Cleoninae, certain Molytinae are employed herein as the out-group for the purpose of tentative
polarization of character states in genera of Cleoninae.
Evidence that Cleoninae are a very primitive lineage within Curculionidae is extensive.
Indeed they may prove sister to the rest of the Curculionidae based on the large number of
plesiotypic character states that they possess.
Buchner (1933, 1953) and Scheinert (1933) discuss at length the mechanism by which adult
female Cleoninae pass on symbiotic bacteria to the egg and site of localization of these
symbionts in the larvae. In Cleoninae, symbiont pouches are paired and attach to the vagina
near the base of gonocoxite II. Bacteria develop in these pouches embedded in a secretion
originating from the epithelial cells of the pouch. As eggs pass through the vagina they are
coated with bacteria which are then ingested by young larvae. In the larvae, there are four
variously shaped evaginations in the anterior part of the midgut where the bacteria are
localized. Both of these states are unique within Curculionidae to Cleoninae and are very
similar to those of Anobiidae, Lagriidae and Cerambycidae (Buchner, 1953). Recently, Mann
and Crowson (1984) noted that similar pouches to those in adult female Cleoninae also occur in
various Chrysomelidae, a constituent taxon of the presumed sister-lineage of Curculionoidea;
site of localization in larvae was not examined. Although Curculionoidea generally regarded to
be primitive have not been examined for these characters, the widespread distribution of the
states in related groups suggests that these states are plesiotypic in Curculionidae and that
Cleoninae is a very primitive lineage within that family.
Wing structure is also primitive (Kingsolver, 1961), because in at least some Cleoninae, the
branches of the second anal vein are joined at the base and the basal one-half of the first anal
vein is present. These features appear to be unique to Cleoninae within Curculionidae and are
undoubtedly plesiotypic because they are found in primitive Curculionoidea.
Ting (1936) has also noted that in Curculionidae, Cleoninae are the only subfamily in which
adults have the labial palpi ventral in their site of attachment to the labium. Although Ting
(1936) suggests this to be the autapotypic state, the fact that the same state is found in
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Anderson
primitive Curculionoidea suggests that it is plesiotypic.
Further evidence for a primitive phylogenetic position for the subfamily comes from the
structure of the testes (as noted previously), and generalized stem or root mining habits of
larvae of the subfamily.
Monophyly of Cleoninae is indicated by structure of the labial palpi, which, although
described as of one article (Aslam, 1963; Ter-Minasyan, 1963), are of three very small and
telescoped articles (Figs. 61-63). This character state appears to be unique to Cleoninae within
Curculionoidea. Most other Curculionoidea possess larger three-segmented labial palpi; some
lack palpi entirely or have the palpi variously modified, but these latter states are clearly
autapotypic. At present, this is the only autapotypic character state supporting monophyly of
Cleoninae.
Phylogenetic relationships among tribes of Cleoninae
Within Cleoninae, Csiki (1934) recognized four tribes: Lepyrini, Cleonini, Rhinocyllini, and
Lixini. Lepyrini, has since been removed from Cleoninae and placed in Molytinae (Solari,
1941; Ter-Minasyan, 1963). Since then, Aslam (1963) and Ter-Minasyan (1978) have
recognized only two tribes, Cleonini and Lixini; O’Brien and Wibmer (1982) continue to
recognize three tribes, as above, excluding Lepyrini.
In general, the tribe Lixini are all those Cleoninae whose adults have a more
elongate-narrow cylindrical rostrum and elongate-narrow body form. The tribe Cleonini on the
other hand, includes taxa whose adults have a more robust rostrum and body form. Cleonini are
generally associated with arid habitats; Lixini, with mesic habitats. Rhinocyllini, when used, is
reserved for the genera Rhinocyllus and Bangasternus which are otherwise placed as Lixini.
No satisfactory suprageneric classification has been proposed for Cleoninae, because the
phylogenetic relationships of the world genera must be assessed before such can be realized. No
tribal placement is therefore herein attempted although generic relationships of North
American taxa, traditionally placed as Cleonini, are discussed and represent a start at a
reconstructed phylogeny of genera and revised tribal classification.
Phylogenetic relationships of genera of New World Cleonini
In an attempt to determine the relative position of Apleurus and Cleonidius, a survey of
representatives of 50 genera of world Cleoninae was undertaken (Appendix 2). In instances
where type species have not been designated for a given genus, the species considered
representative of that genus is one originally included and which could subsequently be
designated as the type species. Where possible, relationships were inferred on the basis of
shared, presumably derived character states, preferably unique to those taxa or found in few
other Cleoninae (preferably those taxa, which for reasons of distribution of other more
widespread and presumably apotypic character states, are not considered closely related to the
genus under study). Inferred monophyly of this grouping was then corroborated by examination
of distribution of either apotypic or plesiotypic states of other characters (with more widespread
distributions) for concordance.
Confidence in phylogenetic relationships of Apleurus is high because a large number of
representatives of genus-group names in Cleonini were examined and considered in the search
for out-groups. Unfortunately, confidence in out-group relationships of Cleonidius is low,
largely because few representatives of genus-group names of Lixini were examined. Although
Cleonidius has traditionally been considered a member of Cleonini (the initial seach for an
New World Cleonini
551
out-group accepted that premise), indications are that Cleonidius (and Cylindropterus) is more
closely related to Lixini than to any traditional Cleonini. In fact, as noted elsewhere, some
Lixus species proved to belong in Cleonidius. A revision of this large and structurally divergent
genus is required on a world basis to better formulate the phylogenetic relationships of
Cleonidius.
For Apleurus, out-groups Cnemodontus and Chromoderus, together with Apleurus itself,
are inferred to compose a monophyletic group based primarily upon presence of a variously
developed acute pronotal postocular projection fringed with anteriorly directed, long, postocular
vibrissae of unequal length (Figs. 81-88). This state is otherwise found only in species of Lixus
(Fig. 4), Lixocleonus and Microlarinus (Fig. 1), taxa that on the basis of distributions of states
of other characters are not considered closely related to Apleurus.
Recognition of this group as monophyletic is supported by the distributions of the following
character states, which are shared, except as noted, by members of the three genera and in most
instances, by a variable number of other genera as well. Polarity of the states of these
characters may or may not have been determined.
1 . Body form more or less robust. The three taxa under consideration possess this state as do
numerous other Cleoninae. Molytinae of phylogenetic interest are also more or less robust in
body form; therefore, this state is likely plesiotypic.
2. Pronotum with scale pattern with large white scales present in a lateral stripe of variable
width, small and fine in a moderately broad to very broad apically narrowed median stripe;
median area largely black in color, the underlying dark cuticle not obscured by the overlying
scales (Figs. 24, 68-80). This state is present in Apleurus and Chromoderus , as well as
numerous other genera of Cleoninae. A distinct pronotal scale pattern is not evident in
Cnemodontus. Polarity of the states of this character has not been determined.
3. Eye like an inverted, elongate teardrop in shape. This state is found in the three genera in
question as well as in numerous other genera of Cleoninae. Polarity of the states of this
character has not been determined.
4. Eye prominent and convex in dorsal view. This state is found in Apleurus and
Cnemodontus (but not Chromoderus ), as well as in various Lixus species and Lixocleonus.
Polarity of the states of this character has not been determined.
5. Female with abdominal sternum VIII with basal arm absent (Figs. 56e-57e, 89a-96a).
This state is found in all three genera as well as in numerous other genera of Cleoninae.
Molytinae possess a long basal arm and thus lack of the arm is likely apotypic but also
homoplasious.
6. Tarsi with articles moderately wide, not markedly elongate-narrow. This state is found in
Chromoderus , Cnemodontus and some Apleurus species as well as numerous other Cleoninae.
Molytinae of phylogenetic interest also possess this state and thus it is likely plesiotypic.
7. Tarsi with ventral pilose pads present, large and distinctly delimited. This state is found in
Chromoderus , Cnemodontus and some Apleurus species as well as numerous other Cleoninae.
Molytinae of phylogenetic interest also possess this state and thus it is likely plesiotypic.
8. Pronotum with dorsal surface punctate, not tuberculate. This state is found in
Chromoderus , Cnemodontus and Apleurus species as well as numerous other Cleoninae.
Polarity of the states of this character has not been determined.
9. Female with spermathecal gland spherical, not variously elongate and cylindrical. This
state is found in Chromoderus , Cnemodontus and Apleurus species as well as numerous other
Cleoninae. Polarity of the states of this character has not been determined.
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10. Rostrum moderately elongate-narrow to robust. This state is found in the three taxa in
question as well as various other Cleoninae. Polarity of the states of this character has not been
determined.
11. Tibiae with corbel ridge sharp (as in Fig. 6). This state is found in Chromoderus,
Cnemodontus and Apleurus species as well as numerous other Cleoninae. Polarity of the states
of this character has not been determined.
12. Ventral surface of male lacking extensive long erect vestiture. This state is found in
Chromoderus , Cnemodontus and Apleurus species as well as numerous other Cleoninae.
Molytinae of phylogenetic interest also possess this state and thus it is likely plesiotypic.
13. Female with gonocoxite II triangular in shape (Figs. 89b-96b), apex not prolonged into a
marked lobe (Fig. 54). This state is found in Chromoderus, Cnemodontus and Apleurus species
as well as numerous other Cleoninae. Molytinae of phylogenetic interest also possess this state
and thus it is likely plesiotypic.
14. Antenna with apical three articles with placoidal sensilla. This state is found in
Chromoderus, Cnemodontus and Apleurus species as well as various other Cleoninae. Polarity
of the states of this character has not been determined and not all Cleoninae have been
examined in detail for this character.
For Cleonidius, out-groups Lixus ( Lixoglyptus ) and Cylindropterus were selected largely
because of their close resemblance to Cleonidius species, yet lack of the internal swelling at the
base of abdominal sternum VII in the female which characterizes the latter. No synapotypies
were identified to unite the three taxa as a monophyletic group and indeed, the group may
prove to be paraphyletic. Distributions of the states of the following characters support a close
but unresolved relationship among these three taxa.
1. Body form more or less elongate-narrow, cylindrical. The three taxa under consideration
possess this state as do Mecaspis, Lixocleonus, and most Lixus. Molytinae of phylogenetic
interest are more robust in body form therefore elongate-narrow form is likely apotypic.
2. Pronotum with scale pattern slightly to distinctly vittate, with white scales largest and/or
densest immediately laterad of midline, smaller and/or sparser laterally; largely whitish in
color, the underlying dark cuticle largely obscured by the white scales (Figs. 118-136). This
state is present in Lixoglyptus and Cleonidius species as well as Pseudocleonus, Mecaspis, and
various other species of Lixus. A distinct pronotal scale pattern is lacking from many genera
including Cylindropterus.
3. Eye elongate-oval in shape. This state is found in the three genera in question as well as
numerous other genera of Cleoninae. Polarity of the states of this character has not been
determined.
4. Eye more or less flat in dorsal view. This state is found in the three taxa in question as
well as numerous other Cleoninae. Polarity of the states of this character has not been
determined.
5. Tibiae with corbel ridge rounded (as in Fig. 5). This state is present in the three taxa in
question, Lixocleonus, Menocleonus, Epirhynchus, and all Lixus species. Polarity of the states
of this character has not been determined.
6. Tarsi with articles moderately wide, not markedly elongate-narrow. This state is found in
the three taxa in question as well as numerous other Cleoninae. Molytinae of interest also
possess this state and thus it is likely plesiotypic.
7. Tarsi with ventral pilose pads present, large and distinctly delimited. This state is found in
the three taxa in question as well as numerous other Cleoninae. Molytinae of interest also
New World Cleonini
553
possess this state and thus it is likely plesiotypic.
8. Pronotum with dorsal surface punctate, not tuberculate. This state is found in the taxa in
question as well as numerous other Cleoninae. Polarity of the states of this character has not
been determined.
9. Female with spermathecal gland spherical, not elongate and cylindrical. This state is
found in the three taxa in question as well as numerous other Cleoninae. Polarity of the states
of this character has not been determined.
10. Rostrum moderately to very elongate-narrow. This state is found in the three taxa in
question (although not all species of Cleonidius ) as well as various other Cleoninae. Polarity of
the states of this character has not been determined.
11. Pronotum with postocular lobes present. This state is found in the three taxa in question
(although not in all species of Cleonidius ) as well as numerous other Cleoninae. Molytinae of
interest also possess this state and thus it is likely plesiotypic.
12. Female with abdominal sternum VIII with basal arm present, short to long (Figs.
113-114, 157-176). This state is found in the three taxa in question as well as in numerous
other genera of Cleoninae. Molytinae of interest possess a long basal arm and thus this state is
likely plesiotypic with a successively shorter arm increasingly apotypic.
13. Antenna with apical three articles with placoidal sensillae lacking or else very few in
number. This state is found in the three genera in question as well as Lixocleonus and Lixus
species. Polarity of states of this character has not been determined and not all Cleoninae have
been examined in detail for this character.
14. Prosternum with swelling in front of fore-coxal cavity. This state is found in the three
taxa in question and Cosmogaster, Mecaspis, Lixocleonus, Microcleonus and Lixus. Polarity
of states of this character has not been determined.
Genus Apleurus Chevrolat
Character evolution
Character state 1 for each character is considered plesiotypic; other states are considered
apotypic at various levels (see Table I).
Primary characters
Character 1. Epistoma, apical margin .— Two states of this character are recognized:
1. emarginate (Fig. 81b);
2. rounded and produced (Figs. 82b-88b).
Out-groups, most Cleoninae, and Molytinae possess an emarginate epistoma. An apically
rounded epistoma is otherwise known only in Gonocleonus , Leucochromus, Eurycleonus, and
Koenigius.
Character 2. Prementum, setae. — Two arbitrarily defined states of this character are
recognized:
1. lacking or with only one large seta per side (Figs. 61-62);
2. with two or more large setae per side (Fig. 63).
Out-groups Cnemodontus and Chromoderus lack and possess one large seta per side
respectively. Other Cleoninae were not examined in detail for this character, however, at least
some species of Lixus and Cleonidius possess two or more long setae on each side of the
prementum.
Quaest. Ent., 1987, 23 (4)
554
Anderson
Character 3. Pronotum, tubercles, development and shape of anterior angles .— Three
states are recognized:
1 . absent (Figs. 24, 68-69);
2. present, anterior angles obtuse (Figs. 75-80);
3. present, anterior angles acute (Figs. 70-74).
Out-groups lack distinct tubercles but the pronota are slightly constricted at the apical
one-third to one-quarter, the position of the tubercles if developed. This and lack of
acutely-angled tubercles in other Cleoninae which possess variously developed pronotal
tubercles, justify recognition of obtusely angled tubercles as plesiotypic.
Character 4. Tarsus, ventral pilose vestiture. — Three arbitrarily defined states are
recognized:
1. extensive;
2. moderately reduced in extent;
3. markedly reduced in extent, or absent.
Out-groups each possess extensive ventral tarsal pilose vestiture. This state is considered
plesiotypic and successive stages in reduced extent of pilose vestiture increasingly apotypic.
Molytinae possess extensive ventral pilose vestiture. Extent of ventral tarsal pilose vestiture
varies extensively in Cleoninae and on that basis alone this character should perhaps best be
regarded as of a secondary nature. However, compatibility of successively apotypic states of
this character with numerous other primary characters results in increased confidence that it
also be considered primary.
Character 5. Female, stylus, size. — Three states are recognized:
1. large (Figs. 89b-91b);
2. small (Figs. 92b-94b);
3. absent (Figs. 95b-96b).
Out-groups and all Cleoninae examined (except for Brachycleonus in which a stylus is
absent) possess a moderately large stylus. Decreased size is considered increasingly apotypic.
Character 6. Female, stylus, position. — Three states are recognized:
1. apical (Figs. 89b-91b);
2. anteapical (Figs. 92b-94b);
3. absent, anteapical sclerotized ridge present (Figs. 95b-96b).
Out-groups and all Cleoninae examined in which a stylus is present, possess an apically
situated stylus. The anteapical sclerotized ridge is regarded as positionally homologous with the
stylus in other Apleurus species and thus state 3 is not considered as increasingly apotypic.
Character 7. Male, abdominal sternum VII, dorsally directed tooth at apical margin. —
Two states are recognized:
1. absent;
2. present.
Out-groups and all Cleoninae except Brachycleonus and Centrocleonus lack a dorsally
directed tooth on the apical margin of abdominal sternum VII.
Character 8. Male, abdominal sternum VIII, interior angles of each sternite. — Three states
are recognized:
1. basal projection absent (Fig. 97f);
2. basal projection slightly to moderately developed (Fig. 101 e);
3. basal projection present, markedly developed (Fig. 103e).
New World Cleonini
555
Both out-groups lack a basal projection. Extent of development from slight to marked
represents successively apotypic states. Other Cleoninae have not been examined in detail for
this character although a distinct basal projection is lacking from both Cleonidius and
Stephanocleonus.
Secondary characters
Character 9. Size, length of elytra. — Three arbitrarily defined states are recognized (Fig.
105).
1. X<7.0 mm;
2. 7.0 mm<X<9.0 mm;
3. X>9.0 mm.
Large size (state 3) is widespread in Cleoninae but the out-groups Chromoderus and
Cnemodontus possess state 1 and state 2 respectively. Small size of Chromoderus and primitive
Apleurus species, however, suggests that the common ancestor of Chromoderus and Apleurus
was also of small size, and that medium to large sizes (states 2 and 3) in Apleurus are
successively apotypic. Whether the medium size of Chromoderus is then primitive or represents
an independent increase in size is not known.
Character 10. Eye, prominence and convexity. — Three arbitrarily defined states are
recognized:
1. markedly convex and prominent;
2. slightly to moderately prominent and convex;
3. flat.
Flat, non-prominent eyes are known in virtually all Cleoninae with the exception of
Lixocleonus , Microlarinus , and various Lixus species. Out-groups Cnemodontus and
Chromoderus have markedly convex and flat eyes respectively. As for character 9, occurrence
of markedly convex eyes in one of the out-groups and in primitive species of Apleurus , suggests
that this state is plesiotypic. Decreased prominence and convexity of eyes is thus considered
apotypic.
Character 11. Suberect or erect vestiture, length. — Three states are recognized:
1 . absent or indistinct;
2. short;
3. long.
Out-groups lack distinct erect vestiture. Presence and subsequently increased length of
vestiture are considered successively apotypic. Long suberect or erect vestiture is known in
numerous genera of Cleoninae and other Curculionidae ( e.g ., Trigonoscuta, Miloderes, etc.)
and appears correlated with life in sand substrates.
Character 12. Rostrum, carina. — Two states are recognized:
1. present (Figs. 84-86);
2. absent (Figs. 81-83, 87-88).
Both out-groups possess a rostral carina. However, primitive Apleurus species lack a carina.
This suggests that lack of a carina is apotypic for Apleurus but that presence of a carina in
certain species of Apleurus is also apotypic.
Character 13. Rostrum, declivity at apex. — Two states are recognized:
1. flat to at most slightly declivous (Figs. 81a-83a);
2. moderately to steeply declivous (Figs. 84a-88a).
Quaest. Ent., 1987, 23 (4)
556
Anderson
Out-groups possess a flat to slightly declivous rostral apex. A rostrum with a steeply
declivous apex also occurs in many other Cleoninae.
Character 14. Maxillary palpus, setae. — Two states are recognized:
1 . one large seta on palpifer, one on stipes (Fig. 64);
2. one large seta on palpifer (Fig. 65).
Out-groups possess large setae on both palpifer and stipes. Whether this seta is lacking or
simply reduced in size was not determined. Other Cleoninae were not examined in detail for
this character.
Character 15. Pronotum, width. — Two states are recognized (see Fig. 109):
1. width greatest at base to width at base more or less subequal to width at tubercles;
2. width distinctly greatest at tubercles.
Out-groups have pronota that are widest at the base. Pronota that are distinctly widest at
the tubercles are also known in Tetragonothorax and Gonocleonus.
Character 16. Pronotum, punctation. — Two states are recognized:
1. small and shallow;
2. large and deep.
Out-groups have pronota that have small shallow punctures. No other Cleoninae examined
have similarly large and deep pronotal punctures.
Character 17. Pronotum, median carina. — Two states are recognized:
1. absent;
2. present, of variable extent.
Out-groups lack a pronotal carina. Presence of a carina is widespread and common in most
genera of Cleoninae.
Character 18. Prosternum, impressions and swellings. — Four arbitrarily defined states are
recognized in this lineage:
1. impression anterolaterad of fore-coxal cavity present; no swelling present;
2. impression anterolaterad of fore-coxal cavity present; swelling immediately anterior to
impression present, low and rounded (Figs. 82a-83a);
3. impression anterolaterad of fore-coxal cavity present; swelling immediately anterior to
impression present, high (Figs. 81a, 84a-88a);
4. impression anterolaterad of fore-coxal cavity present; swelling immediately anterior to
impression present, low and rounded; low rounded swelling also present immediately in front of
fore-coxal cavity.
All Cleoninae possess an impression, although various in depth and extent, immediately
anterolaterad of each fore-coxal cavity. Swellings, if present, are found in two locations, one,
immediately in front of the fore-coxal cavity, adjacent to the impression, or two, immediately in
front of the impression. Out-group Cnemodontus lacks swellings and Chromoderus is unique in
Cleoninae examined in that it has swellings in front of both the fore-coxal cavity and the
impression. Primitive Apleurus have low rounded swellings anterior to the impression, thus this
state is judged plesiotypic for Apleurus ; high swellings in front of the prosternal impressions are
considered apotypic. Swellings in front of the impression are widespread in Cleoninae but are
generally low and rounded except in Stephanocleonus and Conorhynchus . They are however
absent from most genera. Cosmogaster, Mecaspis, Lixocleonus, Microcleonus, Lixus and
Cleonidius all have a swelling of variable extent in front of the fore-coxal cavity.
Character 19. Elytra, elevation of intervals. — Two states are recognized;
1. all intervals equally flat or only slightly elevated and convex;
New World Cleonini
557
2. sutural interval and intervals 3, 5, 7, and 9 slightly to markedly elevated and convex.
Out-groups possess state 1. The apotypic state occurs in many genera of Cleoninae and other
Curculionidae. Development of elevated and convex elytral intervals appears to be correlated
with reduction in size or absence of the metathoracic wings.
Character 20. Fore-tibia of female, apex. — Two states are recognized:
1. uncus and large subapical tooth present;
2. uncus and at most only small subapical tooth present.
Out-groups each possess a large tooth in addition to the uncus. Both states are widespread in
Cleoninae.
Character 21. Fore-tibia, inner margins. — Two states are recognized:
1 . large denticles present;
2. with at most only small denticles present.
Cnemodontus individuals possess large fore-tibial denticles; Chromoderus individuals, only
small denticles. Both states are widespread in other Cleoninae.
Character 22. Female abdominal sternum VIII, lateral arms. — Two states are recognized:
1. more or less straight throughout length (Figs. 49e-50e, 90b-91b, 95b-96b);
2. arcuate (Figs. 89b, 92b-94b).
Abdominal sternum VIII in females of both out-groups has lateral arms that are more or
less straight. Both states are widespread in Cleoninae.
Character 23. Female abdominal tergum VII, median longitudinal carina. — Three states
are recognized:
1. absent;
2. slightly developed;
3. markedly developed.
Cnemodontus lacks a longitudinal carina whereas a markedly developed carina is present in
Chromoderus. Other Cleoninae examined lack a carina.
Character 24. Female abdominal tergum VIII, median longitudinal carina. — Three states
are recognized:
1. absent;
2. slightly developed;
3. markedly developed.
Cnemodontus lacks a longitudinal carina whereas a slightly developed carina is present in
Chromoderus. Other Cleoninae examined lack a carina.
Character 25. Male, aedeagus, curvature. — Two states are recognized:
1. markedly arcuate at base (Figs. 56a-57a);
2. more or less evenly arcuate throughout length (Figs. 97d, 98c-104c).
Both out-groups possess an aedeagus that is markedly arcuate at the base. Both states occur
in Stephanocleonus (Ter-Minasyan 1979). The aedeagus of all Cleonidius species, Lixoglyptus
and Cylindropterus is evenly arcuate throughout its length. Other Cleoninae have not been
examined in detail for this character.
Character 26. Male, aedeagus, apex. — Two states are recognized:
1. spatulate (Figs. 56a-57a);
2. not spatulate (Figs. 97d, 98c-104c).
Both out-groups possess an aedeagus with a spatulate apex. Both states are widely
distributed in Cleoninae.
Quaest. Ent., 1987, 23 (4)
558
Anderson
Character 21 . Male, aedeagus, internal sac, paired lateral sclerite at midlength. — Two
states are recognized:
1. present (Fig. 57b);
2. absent (Figs. 56b, 97a-104a).
Cnemodontus males possess a distinct paired lateral sclerite at the midlength of the internal
sac; Chromoderus males lack such sclerites. Stephanocleonus , Prionorhinus, and Lixomorphus
males also possess such sclerites, and another unpaired sclerite, situated in a ventral position
closer to the apex of the internal sac, is present in Menocleonus, Pleurocleonus, and
Xanthochelus. This latter sclerite, however, is questionably homologous with the paired sclerite
at midlength.
Character 28. Male, aedeagus, internal sac, shape. — Two states are recognized:
1 . dorsal median pocket low and with two pairs of lobes (Figs. 56b-57b, 97a-99a);
2. dorsal median pocket high and with one pair of lobes (Figs. 100a- 104a).
Cnemodontus males possess an internal sac with state 1; Chromoderus males possess an
internal sac with state 2. The shape of the internal sac varies in other Cleoninae examined, but
only in some species of Cleonidius is it similarly high and possessing of only a single pair of
lobes.
Character 29. Male, aedeagus, internal sac, lobe A. — Two states are recognized:
1. present (Figs. 56b,c-57b,c);
2. absent (Figs. 97a,b-104a,b).
Males of each out-group possess lobe A. This character has not been examined in detail in
other Cleoninae. However, lobe A is present in at least Cleonidius , Lixoglyptus,
Cylindropterus, Adosomus, Cleonis and Cyphocleonus. It is absent from those
Stephanocleonus examined.
Character 30. Male, aedeagus, internal sac, apical sclerite complex, median projection. —
Two states are recognized:
1. long (Figs. 56d-57d, 103d-104d);
2. absent or present, but short and indistinct (Figs. 97c, 98d- 101 d) .
Both Cnemodontus and Chromoderus males possess apical sclerite complexes with a
well-developed median projection. Presence of state 2 in primitive Apleurus species suggests
presence of state 1 within Apleurus is secondarily apotypic. This character has not been
examined in detail in other Cleoninae.
Character 31. Male, aedeagus, internal sac, apical sclerite complex, sclerotization of base
and sides of the apical pocket. — Two states are recognized:
1. well-sclerotized, dark brown in color;
2. lightly-sclerotized, pale brown in color.
Males of each out-group have the base and sides of the apical pocket markedly sclerotized.
This character has not been examined in detail in other Cleoninae.
New World Cleonini
559
Table I. Plesiotypic and apotypic states of characters used in phylogenetic analysis of Apleurus
species and related taxa
Character
Plesiotypic
Apotypic
1 . Epistoma, apical margin
2. Prementum, setae
3. Pronotum, tubercles,
development and shape of
anterior angles
4. Tarsus, ventral pilose
vestiture
5. Female, stylus, size
7. Male, abdominal
sternum VII,
dorsally directed
tooth at apical margin
8. Male, abdominal
sternum VIII,
interior angles
of each sternite
9. Size, length of elytra
10. Eye, prominence and
convexity
1 1 . Suberect or erect
vestiture, length
12. Rostrum, carina
13. Rostrum, declivity
at apex
14. Maxillary palpus, setae
15. Pronotum, width
— emarginate
— setae lacking or only one
large seta per side
— tubercles absent
— present, extensive
— present, large
— absent
— basal projection absent
— X < 7.0 mm
— markedly convex
and prominent
— absent or indistinct
— present
— flat to slightly declivous
— one large seta on palpifer,
one on stipes
— greatest at base to subequal
at base and tubercles
— rounded and produced
— 2 or more large setae per side
— tubercles present, anterior
angles obtuse (2)
— tubercles present, anterior
angles acute (3)
— present, moderately reduced
in extent (2)
— present, markedly reduced
in extent or absent (3)
— present small (2)
— absent (3)
— present, anteapical (2)
— absent, anteapical sclerotized
ridge present (3)
— present
— basal projection present,
slightly to moderately
developed (2)
— basal projection present,
markedly developed (3)
— 7.0 mm < X < 9.0 mm (2)
— X > 9.0 mm (3)
— slightly to moderately
prominent and convex (2)
—flat (3)
— present, short (2)
— present, long (3)
— absent
— moderately to steeply
declivous
— one large seta on palpifer
— greatest at tubercles
6. Female, stylus, position — present, apical
(continued on next page)
560
Anderson
Table I (continued)
Character
Plesiotypic
Apotypic
16. Pronotum, punctation
17. Pronotum, median carina
18. Prosternum, impressions
and swellings
19. Elytra, elevation of
intervals
20. Fore-tibia of female, apex
21. Fore-tibia, inner margins
22. Female abdominal
sternum VIII, lateral arms
23. Female abdominal
tergum VII, median
longitudinal carina
24. Female abdominal
tergum VIII, median
longitudinal carina
25. Male, aedeagus,
curvature
26. Male, aedeagus, apex
— small and shallow
— absent
— impression anterolaterad of
fore-coxal cavity;
no swelling present
— all intervals equally flat or
slightly elevated and convex
— uncus and large subapical
tooth present
— large denticles present
— more or less straight
throughout length
— absent
— absent
— markedly arcuate at base
— spatulate
— large and deep
— present, various in extent
— impression anterolaterad of
fore-coxal cavity;
low rounded swelling
present immediately
anterior to impression (2)
— impression anterolaterad of
fore-coxal cavity;
high sharp swelling
present immediately
anterior to impression (3)
— impression anterolaterad
of fore-coxal cavity;
low rounded swelling
present immediately
anterior to impression and
low rounded swelling
present immediately in
front of fore-coxal
cavity (4)
— sutural and intervals 3, 5, 7,
and 9 slightly to markedly
elevated and convex
— uncus and at most small
subapical tooth present
— small denticles present
— arcuate
— present, slightly
developed (2)
— present, markedly
developed (3)
— present, slightly
developed (2)
— present, markedly
developed (3)
— more or less evenly arcuate
throughout length
— not spatulate
(continued on next page)
New World Cleonini
561
Table I (continued)
Anderson
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Phylogenetic relationships of included species
Distribution of character states is shown in Table II.
Confidence in phylogenetic relationships of species of Apleurus (Figs. 234-235) is high;
most primary characters exhibit very little homoplasy elsewhere in Cleoninae and the
distribution of apotypic states is concordant with distributions of apotypic states in most
secondary characters. Furthermore, primary characters are employed to define most major
lineages and are more general in their applicability (Fig. 234) (i.e., not all primary characters
are used to define the same or few lineages). Because of this, confidence in recognizing
homoplasy in secondary characters is enhanced and their applicability amplified.
Relationships among Chromoderus, Cnemodontus and Apleurus based on primary
characters are unclear (Fig. 234). Apotypic states of secondary characters of size of female
fore-tibial denticles, and presence or absence of a paired lateral sclerite at midlength in the
internal sac of the aedeagus of males, suggests Chromoderus is the sister group to Apleurus ,
with Cnemodontus as sister to the Chromoderus- Apleurus lineage (Fig. 235). Flat eyes in
Chromoderus and most other Cleoninae, and shared presence of swollen and convex eyes in
Cnemodontus and most Apleurus is the only evidence that Cnemodontus is sister to Apleurus.
However, I favour consideration of flat eyes as independently derived in Chromoderus and
swollen and convex eyes as plesiotypic for the lineage.
Monophyly of Apleurus is based upon distribution of apotypic states of a number of
characters, both primary and secondary. All Apleurus have ventral tarsal pilose vestiture
moderately to markedly reduced in extent, the aedeagus of males is evenly arcuate throughout
its length, the apex of the aedeagus is not spatulate, and the internal sac of the aedeagus lacks
lobe A. These apotypic states all occur elsewhere in Cleoninae but not in those taxa believed
closely related to Apleurus. Although there is homoplasy within Apleurus , distributions of
apotypic states of other characters also support monophyly of the genus. These states are lack
of a rostral carina (secondarily derived in A. angularis-A. jacobinus-A. porosus lineage),
fore-tibia of female with small subapical second spur (secondarily derived in A. aztecus ), and
internal sac of aedeagus of male with the median projection of the apical sclerite complex
absent or short and indistinct (secondarily derived in A. albovestitus-A. saginatus lineage).
Apleurus (Gibbostethus) hystrix is hypothesized to be the most primitive species of
Apleurus (Figs. 234-235). It is the only species of Apleurus which has an emarginate
epistoma; all other Apleurus species have the epistoma rounded. Apleurus hystrix is
autapotypic in a number of characters (see species treatment of A. hystrix ), and also shares,
with various other species of Apleurus , a number of apotypic states which are best regarded as
homoplasious. All of these apotypic states are of secondary characters and are otherwise found
only in the A. angularis-A. jacobinus-A. porosus lineage or the common ancestor of this
lineage and the A. albovestitus-A. saginatus lineage. This suggests that A. hystrix might be
better placed with these species, but distribution of apotypic states of all primary characters in
the analysis argues against this proposal (Fig. 234).
Apleurus aztecus is the next most primitive species and is sister to the remainder of the
subgenus Apleurus (Figs. 234-235). Phylogenetic position of this species is based upon
possession of a rounded epistoma, lack of a dorsally directed tooth at the apical margin of
abdominal sternum VII in males, and lack of basal projections on the interior angles of each
sternite of abdominal sternum VIII also in males.
Apleurus lutulentus is sister to the ancestor of the A. angularis-A. jacobinus-A. porosus
and A. albovestitus-A. saginatus lineages (Figs. 234-235). This position is due to distributions
Quaest. Ent., 1987, 23 (4)
564
Anderson
of apotypic states of numerous primary as well as secondary characters. These latter five
species comprise a monophyletic group divisible into two lineages as noted above. Monophyly of
the A. angularis-A. jacobinus-A. porosus lineage is based upon possession of acute-angled
pronotal tubercles, presence of rostral carina, maxillary palpus with only one large seta on
palpifer, large and deep pronotal punctation, alternate elytral intervals elevated and convex,
female abdominal sternum VIII with arcuate lateral arms, female abdominal tergum VIII with
slight median longitudinal carina, and internal sac of aedeagus of male with sides and base of
apical pocket only lightly sclerotized. Within this group of species, long erect dorsal vestiture
and relative width of the basal and apical portions of the pronotum suggests an A. angularis-A.
jacobinus sister species relationship. Widely divergent tarsal claws in all A. angularis and some
A. porosus are likely a result of independent evolution or of hybridization.
The sister species status of A. albovestitus and A. saginatus (Figs. 234-235) is strongly
supported by the lack of a stylus on gonocoxite II of the female and by markedly developed
basal projections on the interior angles of each sternite of abdominal sternum VIII in males.
Distribution of apotypic states of numerous secondary characters supports this relationship.
Alternative hypotheses
The only apparent possible alternative hypothesis of phylogenetic relationship to that
presented in Figure 235 concerns the placement of Apleurus hystrix. Although primary
characters indicate a primitive phylogenetic position for this species, there are a number of
secondary character states homoplasious with the A. porosus-A. jacobinus-A. angularis lineage
which could be interpreted as suggesting inclusion in that lineage.
Genus Cleonidius Casey
Character evolution
Character state 1 for each character is considered plesiotypic; other states are considered
apotypic at various levels (see Table III).
Primary characters
Character 1. Submentum, degree of ventral swelling. — Two states of this character are
recognized:
1. flat, not ventrally swollen (Figs. 137a-142a, 145a-156a);
2. swollen ventrally (Figs. 143a- 144a).
Out-groups and all Cleoninae examined possess a submentum that is not swollen ventrally.
Character 2. Rostrum, ratio of length to width in females. — Four arbitrarily defined states
are recognized (see Fig. 198):
-1. very elongate, X<0.45 (Fig. 141);
1. moderately elongate, 0.45<X<0.65 (Figs. 137-140, 142-144, 148-155);
2. short, robust, 0.65<X<0.77 (Figs. 145, 156);
3. very short, very robust, X>0.77 (Figs. 146-147).
Out-groups possess a rostrum that is moderately elongate. Successively shorter and more
robust rostra are considered increasingly apotypic. A very elongate rostrum is also considered
apotypic. States of this character are widely distributed in Cleoninae although in general
members of the tribe Lixini possess a more elongate, narrow rostrum than do members of the
Cleonini. Rostral length in Molytinae of phylogenetic interest varies from moderately to very
elongate.
New World Cleonini
565
Character 3. Rostrum, degree of lateral swelling.— Two states are recognized:
1 . not swollen at midlength in dorsal view (Figs. 1 37b- 1 42b, 1 45b- 1 56b);
2. swollen at midlength in dorsal view (Figs. 1 43b- 1 44b) .
Out-groups possess state 1. No other Cleoninae or Molytinae of phylogenetic concern
examined possess a similarly laterally expanded rostrum.
Character 4. Frons/vertex, surface sculpture. — Three states of this character are
recognized:
1. area behind eye with wrinkles of cuticle;
2. area behind eye with a curved sulcus (Fig. 150a);
3. area behind eye with a deep, straight sulcus (Fig. 139).
Other than various punctures, out-groups and most Cleoninae possess at most only a few
wrinkles of the cuticle behind the eye. Some African Lixus possess a very deep elongate
impression above the eye. State 3 is unlikely homologous with states 1 and 2.
Character 5. Elytra, scale pattern.— Three states of this character are recognized:
1. marginal; intervals 9 to 11 with dense, moderately large, white scales, otherwise moderately
densely and uniformly covered with fine white scales which are smaller than those on intervals
9 to 11 (Figs. 118-123);
2. vittate; intervals other than 9 to 1 1 with scales as large as those on intervals 9 to 1 1 (Figs.
127-128, 130-136);
3. mottled; with irregularly scattered and sized patches of large white scales (Figs. 124-126,
129).
Of the out-groups, Cylindropterus lacks a distinct scale pattern whereas Lixoglyptus species
both have elytral scale patterns that are of the marginal type. Most Lixus species possess
uniformly dense small fine scales on the elytra although elytra of individuals of a few species
are distinctly vittate. Members of the Cleonini have various elytral scale patterns, but these are
not recognizable as any of the states discussed herein for Cleonidius. State 1 is considered
plesiotypic based largely on its occurrence in Lixoglyptus species. This state is assumed to
represent an intermediate stage between the uniformly distributed small fine scales of most
Lixus and the vittate state of certain Cleonidius species. The mottled state is then assumed to
be a result of decreased scale size at various positions of the vittate pattern and is homoplasious
within Cleonidius although restricted to a single species group.
Character 6. Abdomen of female, ornamentation of abdominal sternum VII. — Two states
of this character are recognized:
1. basal margin uniformly flat;
2. basal margin at middle internally (dorsally) with variously developed glabrous shiny
tubercle.
Out-groups and all other Cleoninae examined possess state 1. Within Cleonidius there is
variation within and between species in the degree of development of the tubercle. Very slightly
developed tubercles or glabrous patches are also present at the basal margin of abdominal
sterna V and VI in some specimens, but in these individuals, the most markedly developed
tubercle is that on sternum VII.
Character 7. Male, aedeagus, internal sac, height of dorsal median pocket. — Two states of
this character are recognized:
l.low (Figs. 11 la-1 12a, 177a-179a, 183a-196a);
2. high (Figs. 180a-182a).
Quaest. Ent., 1987, 23 (4)
566
Anderson
Out-groups possess a low dorsal median pocket. Both states occur elsewhere in Cleoninae
but the character has not been examined in detail.
Character 8. Male, aedeagus, internal sac, apex of dorsal median pocket. — Two states of
this character are recognized:
1. rounded or truncate in dorsal view (Figs. 11 lb- 112b, 1 77b- 191b, 196b);
2. emarginate or sinuate in dorsal view (Figs. 192b- 195b).
Out-groups possess a dorsal median pocket with a rounded to truncate apical margin. Other
Cleoninae have not been examined in detail for this character.
Character 9. Male, aedeagus, internal sac, lobe B. — Two states of this character are
recognized:
1 . present (Figs. 1 1 la-1 12a, 177a- 179a, 183a- 196a);
2. absent (Figs. 180a- 182a).
Out-groups possess a lobe B on the internal sac. Other Cleoninae have not been examined in
detail for this character although both states are found in Apleurus species, Cleonis and
Cnemodontus possess a lobe B, and the lobe is lacking from Stephanocleonus species examined
and Chromoderus.
Character 10. Male, aedeagus, internal sac, lobe D. — Two states of this character are
recognized:
1 . present (Figs. 1 1 la-1 12a, 177a-179a, 183a-196a);
2. absent (Figs. 180a- 182a).
The internal sac of each out-group possesses a lobe D, however, the lobe is small in L.
spartii. Lobe D is present in all species of Apleurus , Cleonis , Stephanocleonus , Chromoderus ,
and Cnemodontus examined.
Secondary characters
Character 11. Size, length of elytra. — Two arbitrarily defined states of this character are
recognized (see Fig. 197):
1 . large, X>9mm;
2. small, X<9mm.
Individuals of both out-groups are large in size but distribution of states of primary
characters within Cleonidius suggests large size of some Cleonidius species is secondarily
apotypic. Within and between species variation in size is extensive in most Cleoninae.
Character 12. Rostrum, punctation. — Two states are recognized:
1 . apical portion with punctures moderately dense and moderately deep;
2. apical portion with punctures sparse, small and shallow.
Out-groups each possess state 1. State 2 occurs in most species of Lixus and appears
associated with an elongate-narrow rostrum. The apotypic state of this character is
homoplasious within Cleonidius.
Character 13. Rostrum, median tumescence. — Three states are recognized:
1. absent (Figs. 139b, 141b-142b, 152b);
2. present, slightly developed (Figs. 140b, 1 43b- 1 44b, 1 46b- 1 49b);
3. present, markedly developed (Figs. 1 37b- 138b, 145b, 1 50b- 151b, 1 53b- 1 56b).
Out-groups each possess state 1 . Increased degree of development of the median tumescence
is considered increasingly apotypic. There is extensive homoplasy in this character.
Character 14. Rostrum, lateral margins. — Two states are recognized:
1. rounded;
New World Cleonini
567
2. sharp.
Out-groups each possess state 1. Species of Lixus and Larinus examined possess rounded
lateral margins; species of Cleonini have either state.
Character 15. Pronotum, anterolateral margins. — Three states are recognized:
1. postocular lobe rounded, well-developed (Figs. 137a-139a, 141a-144a, 1 46a- 151a);
2. postocular lobe rounded, slightly developed (Figs. 140a, 152a);
3. postocular lobe absent, margin straight (Figs. 145a, 153a-155a).
Out-groups and most other Cleoninae possess well-developed postocular lobes. Successively
less markedly developed postocular lobes are increasingly apotypic. Acute pronotal postocular
projections, although present in some Cleoninae which lack rounded postocular lobes (e.g.,
Lixus , Apleurus), are not present in those Cleonidius species that lack a postocular lobe. This
character is homoplasious in Cleonidius.
Character 16. Pronotum, impressions. — Two states are recognized:
1. basal median area shallowly to moderately deeply impressed;
2. basal median area deeply impressed, lateral margins variously longitudinally impressed.
State 1 occurs in both out-groups. Similar sculpture is not present in other Cleoninae
examined.
Character 17. Suberect or erect vestiture, length. — Three states are recognized:
1. absent (Figs. 140a, 152a, 156a);
2. present, short to moderate in length (Figs. 138a-139a, 145a-149a, 151a, 153a);
3. present, long (Figs. 137a, 141a-144a, 150a, 154a-155a).
Both out-groups and many Lixus lack suberect or erect vestiture. Increased length of
vestiture is considered as increasingly apotypic. There is extensive homoplasy in this character
in Cleonidius and in Cleoninae in general.
Character 18. Tarsus, ventral pilose vestiture. — Three states are recognized:
1. extensive;
2. moderately reduced in extent;
3. markedly reduced in extent, to absent.
Both out-groups, all Lixus species examined, many Cleonini, and Molytinae possess
extensive ventral pilose vestiture. Successive stages in reduction of vestiture are considered as
increasingly apotypic. Reduced ventral pilose vestiture is otherwise confined to members of
Cleonini. This character is homoplasious in Cleonidius.
Character 19. Foretibia of female, inner margin. — Two states are recognized:
1. large denticles present (as in Fig. 144c);
2. only small denticles present (as in Fig. 143c).
Both out-groups possess large denticles on the inner margin of the foretibia. Distribution of
states of primary characters within Cleonidius suggests presence of large denticles in some
species of Cleonidius is secondarily apotypic and homoplasious. Both states of this character
occur in other Cleoninae.
Character 20. Female abdominal sternum VIII, length of basal arm. — Two states are
recognized:
1. long (Figs. 165, 167-169, 174);
2. short (Figs. 157-164, 166, 170-174, 175-176).
Out-groups each possess a long basal arm. Both states of this character are widely
distributed in Cleoninae, but as Molytinae examined possess a long basal arm, this state is
likely plesiotypic for Cleoninae. This character is homoplasious in Cleonidius.
Quaest. Ent., 1987, 23 (4)
568
Anderson
Character 21. Female abdominal sternum VIII, shape of lateral arms. — Two states are
recognized:
1. arcuate (Figs. 163, 165-175);
2. more or less straight, divergent (Figs. 157-162, 164, 176).
Out-groups each possess state 1. This character is homoplasious in Cleonidius. Both states
occur in other Cleoninae.
Character 22. Male, aedeagus, internal sac, lobe E. — Two states are recognized:
1 . present (Figs. 1 1 la, 177a-179a, 1 83a- 1 96a);
2. absent (Figs. 112a, 180a- 182a).
The internal sac of Cylindropterus possesses a lobe E; that of Lixoglyptus lacks lobe E.
Lobe E is present in species of Cleonis, Cnemodontus, some Apleurus and at least some
Stephanocleonus. Distribution of states of primary characters within Cleonidius suggests
presence of lobe E be considered plesiotypic.
New World Cleonini
569
Table III. Plesiotypic and apotypic states of characters used in phylogenetic analysis of
Cleonidius species and related taxa
Character
Plesiotypic
Apotypic
1 . Submentum, degree of
ventral swelling
2. Rostrum, ratio of length
to width
— flat
— moderately elongate,
0.45 < X < 0.65
— swollen
— very elongate, X < 0.45 (-1)
—short, robust,
0.65 < X < 0.77 (2)
3. Rostrum, degree of
lateral swelling
4. Frons/vertex, surface
sculpture
5. Elytra, scale pattern
6. Abdomen of female,
ornamentation
of abdominal
sternum VII
7. Male, aedeagus,
internal sac, height of
dorsal median pocket
8. Male, aedeagus,
internal sac, apex of dorsal
median pocket
9. Male, aedeagus,
internal sac, lobe B
10. Male, aedeagus,
internal sac, lobe D
1 1. Size, length of elytra
12. Rostrum, punctation
13. Rostrum, median
tumescence
14. Rostrum, lateral margins
— not swollen at midlength
- — area behind eye with
wrinkles of cuticle
— marginal (see text)
— basal margin uniformly flat
— low
— very short, robust,
X > 0.77 (3)
— swollen at midlength
— area behind eye with curved
sulcus (2)
— area behind eye with deep,
straight sulcus (3)
— vittate (see text) (2)
— mottled (see text) (3)
— basal margin at middle
internally with variously
developed glabrous shiny
tubercles
—high
-rounded or truncate in
dorsal view
-emarginate or sinuate
in dorsal view
— present
— absent
— present
— absent
-X > 9 mm —X < 9 mm
-apical portion with punctures — apical portion with punctures
moderately large, dense sparse, small and shallow
and moderately deep
-absent — present, slightly
developed (2)
— present, markedly
developed (3)
-rounded — sharp
(continued on next page)
570
Anderson
Table III (continued)
New World Cleonini
571
Quaest. Ent., 1987,23 (4)
Cyl i ndropterus
572
Anderson
Phylogenetic relationships of included species
Distribution of character states is shown in Table IV.
In contrast to Apleurus, confidence in phylogenetic relationships in Cleonidius is low, due to
the low numbers of primary characters as compared to the number of species in the genus, but
especially the level at which the primary characters are applicable. Seven of the ten primary
characters apply to lineages of only 2 or 3 species; only 3 apply at more general levels, one of
which is used to define the monophyly of Cleonidius (Fig. 236). This leaves a large number of
relationships subject to resolution only on the basis of secondary characters (Fig. 237).
Unfortunately, secondary characters containing any phylogenetic information are also not
numerous, few are applicable at a general level, and many, especially the latter, are subject to
marked homoplasy within Cleonidius (Fig. 237).
Basal relationships among Cleonidius , Cylindropterus and Lixoglyptus are very unclear
and, as discussed elsewhere (see “Phylogenetic relationships of genera of New World
Cleonini”) are left as unresolved (Fig. 237). Monophyly of Cleonidius however, is strongly
supported by the presence of a variously internally (dorsally) developed glabrous shiny median
tubercle at the base of abdominal sternum VII of the female. All secondary characters used in
support of monophyly of Cleonidius involve homoplasy within Cleonidius (Fig. 237).
Relationships of the Palearctic species component of Cleonidius are herein left as unresolved
because of unavailability of series of specimens of C. vibex and lack of specimens of other
Palearctic species that are potentially Cleonidius. Further examination of Palearctic taxa
should resolve the placement of the Palearctic species component of Cleonidius.
Among species groups, monophyly of each of the C. boucardi and C. poricollis groups is
substantiated by a single primary character (Fig. 236) and also in the C. boucardi group by an
additional two secondary characters (Fig. 237). Other species groups are defined as
monophyletic strictly on the basis of secondary characters, some of which involve extensive
homoplasy. Monophyly of the C. poricollis group, which contains only two species, is based
upon the primary character of presence or absence of a curved sulcus behind the eye. The
apotypic state is found in both C. poricollis and C. calandroides , but, not in all individuals of
either species. Monophyly of the C. boucardi group is based upon the emarginate or sinuate
apex of the dorsal median pocket of the internal sac of males, larger size (secondarily reduced
in C. quadrilineatus ), and reduced pronotal postocular lobes (homoplasious in C. canescens). In
the C. americanus group, monophyly is based upon the ventral tarsal pilose pads being reduced
in extent (homoplasious in other Cleonidius and other Cleoninae). High incidence of
brachyptery and of a mottled elytral scale pattern may further support monophyly of this
group. Monophyly of the C. erysimi group is based upon apotypic states of two secondary
characters of abdominal sternum VIII of female; lateral arms straight; and basal arm short.
Both of these characters exhibit homoplasy elsewhere in Cleonidius.
Within the C. erysimi group, three primary characters indicate that C. subcylindricus-C.
texanus-C. longinasus comprise a monophyletic group. Relationships of this lineage to the rest
of the species group, however, are unclear (Fig. 237). Cleonidius erysimi and C.
eustictorrhinus are considered sister-species, and this lineage along with C. pleuralis and the C.
subcylindricus-C. texanus-C. longinasus lineage, are left as elements of an unresolved
trichotomy. Cleonidius texanus and C. longinasus are regarded as sister-species because they
lack large punctures apically on the rostrum and have long dorsal erect vestiture. Cleonidius
subcylindricus is therefore sister to the C. texanus-C. longinasus lineage.
New World Cleonini
573
In the C. americanus group, primary characters support only the sister-species status of
each of C. americanus and C. frontalis , and C. infrequens and C. puberulus (Fig. 236); all
other relationships are based on secondary characters (Fig. 237). Cleonidius notolomus and C.
collaris are regarded as sister-species based upon pronotal sculpture although the state in C.
collaris is much more extreme in its irregularity than that in C. notolomus. A moderately
broad rostrum is used to propose monophyly of the C. canescens-C. infrequens-C. puberulus-C.
notolomus-C. collaris lineage, and sharp lateral margins of the rostrum, the sister group status
of the C. infrequens-C. puberulus and C. notolomus-C. collaris lineages. Acceptance of the C.
notolomus-C. collaris lineage as having a secondarily plesiotypic, more elongate-narrow
rostrum and of C. puberulus having secondarily rounded lateral rostral margins is necessitated
by the sister-species status of C. puberulus and C. infrequens. Other interpretations require
that these two not be sister-species and are in my view less parsimonious than that presented
above. Cleonidius canescens is thus the sister-species of the C. infrequens-C. puberulus-C.
notolomus-C. collaris lineage and the C. americanus-C. frontalis lineage sister to the
remainder of species in the C. americanus group.
In the C. boucardi group, C. boucardi is considered sister to the remaining three species in
the group based upon the ventral tarsal pilose pads being reduced in extent and the pronotal
postocular lobes also reduced (both homoplasious in Cleonidius) in the latter three species. C.
quadrilineatus and C. placidus are considered sister-species, with C. trivittatus as sister to that
lineage, because of long dorsal erect vestiture in the former two species.
In the reconstructed phylogeny of Cleonidius , great emphasis is placed on scale pattern
because this is the only primary character applicable in a broad or general sense within this
genus (Fig. 236). As a result, misinterpretation of distributions of states of other characters are
influenced by the topology established on the basis of this one character. If this character is
misinterpreted, radical changes in the reconstructed phylogeny would result.
Alternative hypotheses
Alternative hypotheses of phylogeny in Cleonidius mainly concern phylogenetic
relationships of the species in the C. boucardi group. In all other species groups, hypothesized
phylogenetic relationships are largely consistent with specific instances of character state
intergradation between geographically approximate species (i.e., instances of intergradation are
between geographically approximate sister-species) and with geographical distribution of
lineages in general (i.e., sister lineages are largely geographically approximate). In the C.
boucardi group however, this is not so because character state intergradation is marked
between each of C. boucardi, C. trivittatus , and C. placidus, none of which are hypothesized as
sister-species of any one of the others (see Fig. 237). To be consistent with patterns of
intergradation, C. quadrilineatus would have to be the sister-species of a lineage composed of
C. boucardi, C. trivittatus, and C. placidus, an unfavourable revision in the hypothesized
phylogeny that would require additional and unsatisfying homoplasy in the characters of form
of the anterolateral margins of the pronotum, and extent of ventral pilose tarsal vestiture, and,
reconsideration of long suberect or erect body vestiture as plesiotypic for the species group.
Ability to interbreed, presumably the likely cause of the intergradation, is a plesiotypic feature
and thus cannot be used as evidence of common ancestry although it may be tempting to do so.
Alternatively, the observed intergradation in character states among C. boucardi, C.
trivittatus, and C. placidus, geographically approximate distributions of the three, and
association with Leguminosae for at least C. trivittatus and C. placidus (plant relationships are
Quaest. Ent., 1987, 23 (4)
574
Anderson
unknown in C. boucardi ), could warrant recognition of only a single species composed of three
geographically distinct forms.
I do not favour either of these approaches, the former because it necessitates additional
homoplasy, and the latter because it necessitates extensive polymorphism in a number of
otherwise phylogenetically informative characters. I prefer recognition of three species, and
attribute the geographical inconsistency to the host shift from Leguminosae to
Roseaceae/Rhamnaceae in C. quadrilineatus, which has likely promoted structural
differentiation in that species and which has permitted redispersal such that C. quadrilineatus
is now largely sympatric with other members of the C. boucardi group.
EVOLUTIONARY ANALYSIS: BIOGEOGRAPHY
Methods of Biogeographic Analysis
Historical and ecological factors combine to produce the observed distributions of plants and
animals, the explanation of which is the goal of the science of biogeography. Unfortunately
many past studies of biogeography attempt to explain the distributions observed in terms of
only one of ecology or history (Endler, 1982). Distinguishing the relative roles of these factors
must at least be considered although most agree it is very difficult to separate the effects of the
two and to estimate their relative roles (Endler, 1982; Vuilleumier and Simberloff, 1980). The
present study is no exception and my approach, although attempting to encompass aspects of
both historical and ecological biogeography, will be largely historical in its focus.
Knowledge of distribution and aspects of natural history combine with knowledge of
phylogenetic relationships and of earth history to allow for discussion of the historical
biogeography and ultimately, the evolutionary history of the taxon in question. Ecological
biogeography on the other hand does not require knowledge of phylogeny and earth history, but
rather ecological theory and principles. Systematists and ecologists are the proponents of the
historical and ecological approaches respectively. Rarely are the two approaches combined in a
single study and relative roles assessed. This is brought about simply by the nature of the data
bases because systematists frequently have not carried out the detailed sampling necessary for
the application of ecological theory. Similarly, more often than not, ecologists do not have the
knowledge of phylogeny that predicates all historical biogeographic analyses.
In ecological biogeography, the units selected for study are usually communities or various
guilds of taxa that share association with a particular resource, be it habitat, food, geographic
area, etc. (Cody and Diamond, 1975). Ecological biogeographers are primarily concerned with
temporally proximate factors that determine the composition of the species assemblage but not
the origins of these same factors. Little concern or attention is expressed or given to the
phylogenetic relationships of the constituent taxa and thus there is a large or even total
disregard for historical components because notions about the latter can only be developed in
terms of detailed knowledge of phylogeny. Less frequently, ecological determinants of
distributions of constituents of monophyletic groups, say a genus or family, have been studied,
although rarely if at all with specific reference to detailed phylogenetic relationships.
Much of ecological biogeography addresses questions of dynamics and proximal origins of
diversity, best applicable to islands or insular situations. Diversity patterns on continents are
much more complex and are not so easily studied. Unfortunately, knowledge of population
dynamics and detailed data about natural history attributes, permitting determination of niche
breadths, are required for much meaningful ecological biogeography to be conducted (Cody,
1975; Pianka, 1975). Because of this, the approach I will take herein is only one of examination
New World Cleonini
575
of natural history attributes and distributions, with specific reference to present environmental
parameters.
Historical biogeographers, on the other hand, recognize and stress the need for knowledge of
phylogeny, but generally at the expense of ignoring the role ecology might play in determining
distributions. Indeed, to many, knowledge of phylogeny is essential prior to attempts at any
meaningful historical biogeography. Wiley (1981) has noted three schools of historical
biogeographers: 1, evolutionary; 2, phylogenetic; 3, vicariance (or panbiogeographic, although
Croizat [1982] has disputed this). Detailed knowledge of phylogeny and strictly monophyletic
groups are the units of study for adherents of phylogenetic and vicariance biogeography and
stress vicariance and allopatric speciation. Evolutionary biogeography however, emphasizes the
role of dispersal with little or no regard for speciation mechanism or for details of phylogeny.
This latter is not employed by many contemporary biogeographers for it is the approach
resulting in the narratives and “scenarios” frowned upon for their ad hoc nature, and lack of
applicability and rigour. Heuristic advantages of the phylogenetic and vicariance schools of
biogeography are consistent with the advantages of a phylogenetic approach to phylogeny
reconstruction and are discussed in more detail by Wiley (1981), Patterson (1981), and
Platnick and Nelson (1978).
Ball (1975) recognizes three phases through which biogeographic studies pass: 1, descriptive
or empirical phase; 2, narrative phase; and 3, analytical phase. These phases, as noted
elsewhere herein and by Patterson (1981), apply equally well to systematic studies, especially
methods of phylogeny reconstruction. Descriptive or empirical biogeography is simply the data
base. What are the extant distributions and natural history attributes of the taxa in question?
What habitats do they occupy? What do they eat? When are they active?
In large part, narrative and analytical phases contrast inductive and hypothetico-deductive
approaches to biogeographic studies. Narrative explanations, or general evolutionary
“scenarios” as they have been called by Eldredge (1979), have frequently been criticized as
pseudo-science or bad science because of their unfalsifiable and ad hoc nature, and for their
explanatory value, which is, more often than not, concerned only with particulars as they apply
to the taxon in question (Goudge, 1961; Hull, 1974; Ball, 1975; Eldredge, 1979; Eldredge and
Cracraft, 1980; Brown and Gibson, 1983). General applicability is not usually a concern of
narrative explanations, although there is no reason why it should not be.
Consequently, Eldredge (1979) notes two ways of improving these “scenarios”. Firstly, base
them more explicitly on knowledge of phylogenetic relationships, and secondly, eliminate some
if not all of the more purely speculative elements. To these suggestions, I add a third; formulate
explanations as hypotheses in terms that are testable, predictive, and have potential for more
general applicability than just the taxon in question. With these improvements in mind,
“scenarios” are capable of producing ideas that can be subsequently rigorously tested in a
hypothetico-deductive manner for acceptance as general scientific facts, ideally of widespread
applicability and significance. In essence, this is the analytical phase of Ball (1975).
In my analysis of historical biogeography I employ a phylogenetic approach to place aspects
of the phylogeny of North American Cleoninae in a geographic perspective and thus come to
some degree of understanding of the geographic history of that specific taxon. Resultant
potential generalizations and predictions based on these results are enumerated, discussed and
formulated as hypotheses of general potential utility.
A vicariance approach, examining distribution patterns of other North American arid land
taxa and the plotting of generalized tracks will not be attempted herein but should be carried
Quaest. Ent., 1987, 23 (4)
576
Anderson
out as more studies of such taxa, that include a phylogenetic basis, become available.
Genus Apleurus Chevrolat
Diversity patterns
The genus Apleurus is represented in arid land habitats of southwestern North America and
Mexico by eight species placed in two subgenera. Maximum species diversity is in southern
California, northwestern Mexico (including Baja California Norte), southern Arizona and
southern New Mexico (Fig. 230). Diversity decreases beyond this area especially to the east
and the north. Species of Apleurus do not occur north of 45°N latitude, east of 95°W longitude,
or south of 1 5°N latitude (Fig. 230).
Five of the eight species are stenotopic, variously restricted in their distributions and of these
five, only two, A. jacobinus and A. hystrix are macrosympatric. The remaining three species,
A. lutulentus , A. angularis, and A. albovestitus, are eurytopic, widespread and broadly
sympatric with each other and various of the five geographically restricted species. Gross
geographic range overlap is relatively low with 12 of 28 possible species pairs (42.8%) having at
least partial sympatry. Nine of the 12 sympatric species pairs exhibit overlapping distributions;
three are nested (the area of distribution of one species occurring entirely within the area of
distribution of the other).
Habitat associations
Habitat associations of the species are varied, ranging from grasslands to desert-grassland
transitional habitats or deserts in eurytopic species (three of the eight); others are more
stenotopic, found only in grasslands, desert-grassland transitional habitats, or in deserts (five of
the eight) (Fig. 238).
Apleurus aztecus is a mesquite-grassland inhabitant in central Mexico. Apleurus
lutulentus, although also found in desert-grassland transitional habitats, is primarily a
mesquite-grassland and grassland inhabitant as evidenced by its more easterly distribution into
eastern Texas, Oklahoma and Kansas and disjunct southerly distribution into central Mexico,
but not in the Mexican portion of the Chihuahuan desert (Fig. 207). Apleurus angularis is also
found in grasslands, however, in contrast to A. lutulentus , does not extend as far east or south
as does that species. Rather, A. angularis extends further west and north into Great Basin
desert. Apleurus jacobinus and A. porosus are semi-desert and semi-desert to desert inhabiting
species respectively. Apleurus albovestitus is distributed even more to the west than is A.
angularis. It occurs in most western semi-desert and desert, and desert-grassland transitional
habitats. Apleurus saginatus is a desert-grassland inhabitant bordering the eastern Sonoran
Desert region (Fig. 206). Apleurus hystrix is a sand dune inhabiting species in the California
semi-desert habitat.
Relative measures of association of species with particular types of habitats reveals a
slightly higher level of association with desert-grassland transitional habitats than with others.
These measures were obtained by summing the total number of species of Apleurus occupying
a particular habitat type scoring one for a marked association (solid box in Fig. 238) and
one-half for a less marked association (half box in Fig. 238). This number is then divided by the
number of possible associations with that habitat type to give a relative measure of the degree
to which species of the genus occur in the various habitat types. Measures of association with
the four desert habitats is 0.156; with semi-deserts, 0.250; with desert-grassland transition
New World Cleonini
577
habitat, 0.375; and, with mesquite-grasslands and grasslands, 0.313.
Plant associations
Details of host plant associations are largely unknown for all species and so all plant
associations, many likely accidental, have been used as indicative of the probable taxonomic
range of host plants (Fig. 239). Adults of all species are found primarily or exclusively on
Compositae although in some species, especially A. albovestitus , adults are found sporadically
on a variety of other types of plants as well. Other than Compositae, Chenopodiaceae and
Leguminosae constitute the bulk of the remainder of the associations. Clear associations with
taxa lower than the hierarchical level of family are not evident.
Altitudinal zonation
Altitudinal ranges are shown as a Hubbs-Hubbs diagram (Fig. 232). Although species may
be represented over an extensive range of altitudes, most members of each species are found
over a narrower part of this range.
Apleurus hystrix, A. jacobinus and A. porosus, species restricted to the Pacific coastal
areas, are found over a much narrower range of altitudes than are more widespread species.
Altitudinal ranges of these three coastal species overlap extensively. On the other hand, among
the remaining species, only A. lutulentus and A. aztecus have altitudinal ranges which overlap
extensively. The three remaining species, A. angularis, A. albovestitus , and A. saginatus, have
altitudinal ranges which differ from each other, and from any other species in the genus. There
is, in general, little overlap in the altitudinal ranges of species of Apleurus (Fig. 232). There is
also a trend towards occupation of lower altitudes by increasingly derived species in the genus.
Seasonal occurrence
Patterns of adult seasonal occurrence are broad with A. angularis, A. jacobinus, A.
albovestitus and A. saginatus found throughout the year. Apleurus porosus adults are found
from January to November and are likely active throughout entire year. On the other hand, A.
lutulentus adults are known from March to October; A. aztecus from June to August; and A.
hystrix from February to July. Few records are available, however, for these last two species.
Detailed life histories are unknown for all species.
Co-existence
Consideration of these various niche dimensions as a group reveals that no two species of
Apleurus share the same or largely overlapping ranges in all niche dimensions. Statistical
measures aside, inspection shows species differences along the dimensions of altitude,
geographic distribution, and habitat association. All species share a plant association with
Compositae. Macrosympatric species differ in various aspects of altitudinal range and habitat
association. Species with overlapping altitude ranges are allopatric or differ in habitat
association. In general, niche overlap between species of Apleurus is low.
Wing length polymorphism
At least some adult members of most Apleurus species lack wings; 97.9% of A. lutulentus,
4.5% of A. angularis, 99.8% of A. albovestitus, and all A. saginatus are the only macropterous
individuals known. Adults examined of species of restricted distribution such as A. hystrix, A.
aztecus, A. jacobinus and A. porosus are all apterous. The most widespread species, A.
Quaest. Ent., 1987, 23 (4)
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Anderson
albovestitus, A. lutulentus and A. angularis have various percentages of macropterous
individuals. These individuals are scattered throughout the species ranges and are not restricted
in their distributions.
Genus Cleonidius Chevrolat
Diversity patterns
The genus Cleonidius is represented in North and Central America by 19 species; a single
species has so far been recognized from the Palearctic Region but is not considered further in
the biogeographic discussions following. Maximum species diversity is in the southwestern
United States, especially southern and central California, southern Nevada, Arizona, southern
Utah, New Mexico, Colorado and western Texas (Fig. 231). Diversity decreases slightly to the
north in northern Utah and Wyoming into eastern Washington and western Montana, but more
markedly into Oregon and western Washington to the west, into eastern Texas, Oklahoma and
Kansas to the east, and especially to the south into Mexico (Fig. 231) (although this may prove
an artifact of insufficient collecting). Species of the genus extend north into southern Canada
and south as far as Honduras; two species occur along the east coast of North America.
Species of Cleonidius tend to be more or less widespread in their geographic distributions. I
consider only five of the nineteen species restricted in their geographic distributions. Three of
these five, C. americanus , C. placidus, and C. eustictorrhinus are macrosympatric and found
only in California; C. collaris is found only in eastern Colorado and nearby localities in
adjacent states; and C. calandroides is found only along the northern Atlantic coast. Three
other species (C. puberulus, C. notolomus , and C. infrequens ) are slightly less restricted in
their distributions. Of the remaining 1 1 species, 10 are moderately to very widespread in the
western or southwestern United States and Mexico and are largely macrosympatric with each
other and with various of the species with restricted distributions. One species, C.
subcylindricus, is found along the Atlantic Coastal Plain. One of the six species in the C.
erysimi group, one of two in the C. poricollis group, one of four in the C. boucardi group, and
five of seven in the C. americanus group have more or less restricted distributions.
Gross geographic range overlap is slightly more extensive than in Apleurus. In the C.
erysimi group, eight of 15 possible species pairs have at least partial sympatry (three
overlapping, five nested); two of six comparisons in the C. boucardi group (two overlapping)
and 11 of 21 in the C. americanus group (seven overlapping, four nested) result in at least
partial sympatry. Species in the C. poricollis group are allopatric. For the genus Cleonidius as
a whole, 87 of 171 possible combinations result in at least some sympatry (62 overlapping, 25
nested). In other words, a single species is on average macrosympatric with 50.9% of the genus,
or 9.67 other species of Cleonidius.
Habitat associations
Habitat associations are varied in species of the genus, ranging, as in Apleurus species, from
grasslands to deserts in some, to principally deserts, to principally grasslands in others (Fig.
240). Measures of primary habitat associations (calculated as for Apleurus species), differ
slightly from those in Apleurus however, because in Cleonidius , association with desert
habitats is low (0.099), increasing in semi-desert habitats to 0.171, and desert-grassland
transitional habitats to 0.316. The highest assocation is with mesquite-grasslands and
grasslands (0.474). Association with western xerophytic evergreen forest is 0.132 whereas no
New World Cleonini
579
Apleurus were associated with this habitat type. Among the species groups, the C. erysimi
group is relatively well represented in deserts (0.125), also in semi-desert (0.250), but less so in
desert-grassland (0.167) and markedly less so in mesquite-grasslands and grasslands (0.083).
In contrast, the C. americanus group has values of 0.054 for deserts, 0.107 for semi-desert,
0.429 for desert-grassland, and 0.714 for mesquite-grasslands and grasslands. The C. boucardi
group has values of 0.000, 0.125, 0.250, and 0.500 respectively for these same habitats. The
only southwestern species in the C. poricollis group is found in a variety of habitats. With the
exception of this species, C. poricollis , other Cleonidius species are restricted to one or only a
few habitat types.
Plant associations
Unlike in Apleurus species, which are all associated primarily with Compositae, plant
associations in Cleonidius species are much more varied and span a much broader taxonomic
range of plant families (Fig. 241). A number of species are associated with Compositae,
Rosaceae, Cruciferae, and Leguminosae. There is slight association with Chenopodiaceae and
scattered records of associations with members of various other plant families. In the C. erysimi
group, associations are varied, with one species on each of Compositae, Rosaceae and
Rhamnaceae, and Cruciferae, and two on Leguminosae. Plant associations are not well known
for C. eustictorrhinus. In the C. boucardi group, two species are associated with Leguminosae,
one species with Rosaceae and Rhamnaceae, but also Leguminosae; plant associations are not
known for C. boucardi. Associations are primarily with Compositae and Cruciferae in C.
poricollis ; exclusively with Cruciferae in C. calandroides. Finally, plant associations are few
for most species in the C. americanus group with the exception of C. americanus which appears
to be associated with Compositae.
Altitudinal zonation
Altitudinal ranges are shown as a Hubbs-Hubbs diagram (Fig. 233). In general, each
species is found over a wide range of altitudes which overlap extensively with ranges of many
other species such that Cleonidius species are not as markedly altitudinally segregated as are
Apleurus species. This marked overlap also exists within each of the species groups as well.
Cleonidius poricollis , and three members of the C. boucardi group, C. boucardi , C. trivittatus
and C. quadrilineatus , have the largest altitudinal ranges of species in Cleonidius. Individuals
of these species have been recorded at maximum altitudes ranging from 3276 to 4004 m.
Seasonal occurrence
Patterns of adult seasonal occurrence are broad with most species found throughout the
greater part of the year from early spring to late winter months.
Co-existence
In contrast to species of Apleurus , examination by inspection of niche dimensions as a group
shows the majority of Cleonidius species differ most in terms of host plant (Fig. 241) and
habitat associations (Fig. 240). Species geographic range overlap is generally high (see Table
V), as is the degree of altitudinal range overlap and as is overlap in seasonal occurrence. Within
the C. erysimi group, extent of sympatry and altitudinal range overlap are high, but different
species in the group are associated with different habitat types and with different plant
families. In the C. americanus group, extent of sympatry and altitudinal range overlap are low
Quaest. Ent., 1987, 23 (4)
580
Anderson
New World Cleonini
581
and habitat associations overlap extensively; plant associations are mostly unknown. In the C.
boucardi group, sympatry is low, habitat associations differ, host plant associations differ
slightly, but altitudinal range overlap is high.
The genus Cleonidius is more diverse than Apleurus and is more extensive in its
distribution. Whereas Apleurus species appear to be more associated with deserts and less so
with grasslands, Cleonidius is very well-diversified in grasslands and desert-grassland
transitional habitats and is not associated with deserts to any significant degree (except for
species in the C. erysimi group). Only in Cleonidius are some species associated with the
western xerophytic evergreen forest. Patterns in host plant association differ markedly, for
species of Apleurus are almost exclusively associated with Compositae, whereas species of
Cleonidius are associated to various degrees with one of Compositae, Cruciferae, Leguminosae,
Rhamnaceae, Rosaceae and to a lesser extent Chenopodiaceae.
Wing length polymorphism. — All individuals examined of the C. boucardi group are
macropterous; in the C. poricollis group 99% of C. poricollis and all C. calandroides are
macropterous; and, in the C. erysimi group, only in C. erysimi are brachypterous individuals
known (91%). In contrast, in the C. americanus group, all C. canescens, C. puberulus and C.
collaris , and 79% of C. notolomus and 38% of C. infrequens , are brachypterous.
Apleurus and Cleonidius : Historical biogeography of New World species
Introductory Comments - A Warning
The evolutionary history of plant and animal taxa that presently inhabit the southwestern
United States and adjacent areas has been very complex. Not surprisingly, determination of
what this history has been is also a very difficult task and based on various assumptions.
Changing, locally variable climates, mainly a trend during the Tertiary of increasing aridity
(but here and there punctuated by pluvial periods of various duration) resulting in various and
ephemeral habitat types, continual but localized orogenic activity, and locally variable
topography, all result in a complex framework for speciation events to be understood and even
approximately dated. Adding to this is perhaps the most significant problem, that of the
Quaternary, for it is known that distributions of many organisms changed radically during this
time period (Coope, 1979). Owing to these presumptive changes, many of the distribution
patterns may reflect only Quaternary distributional shifts and not original speciation events,
these perhaps having occurred elsewhere and under very different ecological conditions. In
other words, the present distributions may be the result of secondary, not primary contact zones
(Endler, 1977). In the absence of fossils, the heuristic value of historical biogeography may
therefore be questioned because in order to overcome this problem, assumptions about habitat
associations, other ecological attributes, and especially distributions of the species at different
times, or of their ancestors, are required, based on these features in extant species. These can
only be made with reference to a reconstructed phylogeny and as such the claim of phylogeny
before biogeography is in large part valid for analyses to be maximally scientific. Whether or
not these assumptions should be accepted is a moot point.
Because biogeographic statements are so dependent on a reconstructed phylogeny, another
problem is that of confidence in the reconstructed phylogeny. A weak reconstructed phylogeny
results in weak biogeographic statements that may prove misleading should they be uncritically
accepted as facts.
Quaest. Ent., 1987, 23 (4)
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Anderson
In Apleurus, only the possibility of changed distributions is a problem, for confidence in the
reconstructed phylogeny is high. On the other hand, both are problems in Cleonidius.
Confidence in the fully resolved reconstructed phylogeny is low due to the large number of
secondary characters used in the analysis and concomitant homoplasy in these characters.
Some biogeographic statements about details of species origins in Cleonidius must therefore be
cautiously accepted, and alternatives, some of which are discussed, must also be considered.
A further complication in Cleonidius is recognition of a Palearctic species component which
is not herein considered in the reconstructed phylogeny. The single species examined from the
Palearctic region can, however, be placed either as sister to the Nearctic Cleonidius , or as a
member of the C. americanus group. Biogeographic significance of both placements will be
discussed.
Historical Explanations
In Apleurus , diversity gradients do not appear to be easily explainable in terms of latitude,
elevation, or any other climatic factor with the possible exception of regional degree of variation
in aridity. As noted elsewhere, evolution of Apleurus appears to parallel evolution of
increasingly arid habitats. This is supported by comparison not only of habitat associations with
the reconstructed phylogeny (Fig. 238), but also of altitudinal ranges with the reconstructed
phylogeny which trend toward occupation of lower, and thus likely more arid altitudes. Thus, if
any ecological factor might prove important in explaining why southern California, Arizona,
New Mexico and extreme northwestern Mehave the highest species diversity (Fig. 230), it is
likely the degree of habitat and associated floral complexity largely due to the occurrence of
desert and desert-grassland transitional habitats in these areas. Species diversity drops
markedly beyond the area where these habitats occur.
Similarly, in Cleonidius , maximum species diversity is again noted in these same areas, but
is also high throughout the rest of Arizona and New Mexico, western Texas, central California,
and southern Nevada, Utah, Colorado and Wyoming (Fig. 231). Species diversity thus appears
again to be influenced by degree of arid habitat complexity. However, high species diversity is
more widespread in Cleonidius because of the higher level of association of the species in the
genus with more widespread grassland and woodland habitats (Fig. 240). Although evolution in
this genus has likely also been influenced by increasing aridity in southwestern North America,
these influences are not as clearly indicated as in Apleurus. In Cleonidius , adaptation to the
most arid habitats is low, and to the least arid, relatively high. Furthermore, there is no clearly
evident evolutionary trend for species of the genus to be found in increasingly arid habitats
(Fig. 240), or at successively lower altitudes as are noted in Apleurus. High diversity in
Cleonidius thus extends outside the area of the highest habitat complexity and especially into
areas of grassland, woodland, and desert-grassland transitional habitats. High degree of
association with less arid habitats also explains the broader geographic and altitudinal range of
the species in the genus and perhaps also the occurrence of the genus in the Palearctic region as
well. In addition to the effects of habitat on the evolution of the group, unlike Apleurus , which
are all apparently associated with Compositae, species of Cleonidius are associated with a
much broader taxonomic array of plants (Fig. 241). Clearly, host plant shifts are an additional
factor to consider in the evolution of this latter genus.
In the present study I address the hypothesis that evolution and speciation within Apleurus
and Cleonidius closely parallel increasing aridity and associated changes in distribution and
composition of habitat types in the southwestern United States and Mexico. History of
New World Cleonini
583
distribution and composition of habitats in these areas has been dealt with extensively in
paleobotanical studies by Axelrod (1950, 1975, 1979, 1985), Axelrod and Raven (1985) and
Raven and Axelrod (1978) and thus interpretation of the geographical history of Apleurus and
Cleonidius draws largely upon the results of these publications.
Of three possible origins of arid land faunas in North America, only two are likely for
Cleonini. First, as discussed by Axelrod (1975), a continuous North American-western
Eurasian belt of Madrean-Tethyan sclerophyll woodland vegetation may have lived in the early
Paleogene, with subsequent vicariance of this habitat, as a result of Atlantic Ocean floor
spreading, proposed to account for any present disjunctions in distributions. Secondly, and
perhaps the generally more favourable hypothesis, Webb (1977) notes the marked similarity
between savanna-adapted mammal faunas in Asia and North America in the late Eocene
(postdating similarities between the faunas of Europe and North America), and invokes
subsequent vicariance of these habitats across Beringia as responsible for present distributions.
A third possibility, South American origins, as noted by Raven (1963) and Wells and Hunziker
(1976) for various plants, does not appear likely, because no Cleonini are known from that
continent (Wibmer and O’Brien, 1986) and the Lixus known from there do not appear related
to Cleonidius.
Apleurus seems to be North American in origin and likely evolved in situ , on Compositae, in
isolation from its sister genus, the Palearctic Chromoderus, as a result of vicariance of
sclerophyll woodland or savanna habitats in Holarctica, either between western Europe and
North America no later than early Eocene or between eastern Asia and North America no
earlier than the late Eocene (Fig. 242) respectively. Accepting that Apleurus evolved in North
America and on Compositae, investigation of the role of history proceeds by examination of
distributions and aspects of natural history with respect to the reconstructed phylogeny.
First, aside from A. hystrix, the next two most plesiotypic species, A. aztecus and A.
lutulentus, are exclusively and primarily mesquite-grassland and grassland inhabiting species,
respectively (Fig. 238). This, and the increased association of more derived species with
semi-desert and desert habitats, demonstrates a clear trend from mesquite-grassland and
grassland to desert habitation within species of the genus (Fig. 238), and indicates that
mesquite-grasslands, grassland, or a precursory habitat type not present today, or present but
lacking association with any Apleurus species, was likely the ancestral habitat for the genus.
This habitat was likely a grassland or savanna habitat, because no Apleurus are known from
woodland habitats, plesiotypic Apleurus are currently associated with this or a similar type of
habitat (Fig. 238), no eastern North American Apleurus are known, and Chromoderus species
judging from their distributions, are likely steppe or grassland inhabitants. In view of this, a
trans-Beringian savanna or grassland vicariance origin of Apleurus (Fig. 242) is to be preferred
over the Madrean-Tethyan vicariance of Axelrod (1975), which would predict southwestern
North America-Mediterreanean area relationships, presence of plesiotypic species in woodland
habitats, presence of plesiotypic species in southeastern North America, and presence of closely
related taxa in similar extralimital woodland habitats.
Grasses were present in North America in the early Tertiary as evidenced by first
appearance of grass pollen in the Paleocene, and first grass macrofossils and first appearance of
structural adaptations in mammals to open habitats (including especially, the initial albeit
slight development of high crowned teeth resistant to the abrasive properties of grasses) in the
Oligocene (Clayton, 1981; Webb, 1977).
Quaest. Ent., 1987, 23 (4)
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Anderson
Savanna or woodland precursors of mesquite-grassland and grassland and desert habitats
likely first emerged as distinct habitats in the mid- to late-Eocene as a result of a slowly
expanding dry climate and a marked increase in seasonal aridity fostering in situ adaptation by
already existing subhumid plants to these new and increasingly semi-arid conditions (Axelrod,
1950, 1975, 1979; Axelrod and Raven, 1985; Cronquist, 1978; Webb, 1977). Arboreal
vertebrates are less frequent in fossil deposits of this age, a number of North American groups
developed special adaptations to savanna habitation, and new, savanna-adapted groups
emigrated from Asia where they had already developed their adaptations for this type of life
(Webb, 1977). Adaptation and radiation in animals and plants (especially dicotyledonous
herbs) continued into the Oligocene as arid habitats proceeded to expand and diversify
(Axelrod, 1979; Cronquist, 1978; Webb, 1977). At this time semi-arid woodland or savanna
habitats were apparently relatively extensive in North America (Axelrod, 1975) and were likely
interspersed with localized grasslands. Both faunas of late Eocene (Webb, 1977), and floras of
the Oligocene (Axelrod, 1979), document relationships between taxa in southern California
and those in the Rocky Mountain region (Axelrod, 1975; Raven and Axelrod, 1978). Axelrod
(1979, Figure 6) and Raven and Axelrod (1978) further note that a number of congeneric
“closely related” woodland taxa are presently disjunct in insular and maritime southern
California and the highlands of central Mexico. A continuum may therefore have commented
these two areas and perhaps also the southern Rocky Mountain region from perhaps the late
Eocene, at the earliest, into the Miocene (Axelrod, 1950; Raven and Axelrod, 1978). The
present woodland in the uplands of central Mexico is the habitat thought to be most like the
ancestral Oligocene-Miocene woodland, hence the name Madrean for this vegetation type
(Axelrod, 1975, 1979). Development of increasingly arid habitats intermediate between these
areas throughout the later Tertiary, likely resulted in at least some of these present disjunctions
(Axelrod, 1975).
This geographic pattern applies well to Apleurus because the most plesiotypic species of
Apleurus, A. hystrix, is found in sand dunes in southern California, whereas the next two most
plesiotypic species, A. aztecus and A. lutulentus, live respectively, inland in central Mexico,
and in central Mexico, Arizona, New Mexico, Texas and adjacent states, (Fig. 242). This
pattern, although between a California sand dune species and Mexican or southern Rocky
Mountain mesquite-grassland and grassland taxa, is geographically similar to the relict
disjunctions noted above for semi-arid woodland plants and is suggestive that the same may
also apply to taxa initially adapted to savanna or grasslands. If isolation and the relict nature of
A. hystrix is a result of an early vicariance of California and inland semi-arid grassland or
savanna habitats, we might expect similar phylogenetic and geographic relationships to those in
Apleurus in other taxa in which a structurally markedly distinct species is restricted to one of
the southern California coastal dune habitats. Unfortunately, although endemics are known to
occur in many California dune fields (Andrews et al., 1979), phylogenetic and geographic
relationships of the endemics have not been studied in detail. Most extensive survey efforts have
also concentrated principally on inland dune fields (Andrews et al ., 1979; Hardy and Andrews,
1976). Some of these endemics are also not of a relict nature and are likely a product of the
later Tertiary climatic changes that fostered the rapid and numerous speciation events thought
to have contributed so extensively to the marked levels of endemism in the California flora
(Stebbins and Major, 1965; Raven and Axelrod, 1978). Perhaps relicts such as A. hystrix in
California, are managing to survive in restricted or marginal habitats having changed habitat
associations from the grassland or savanna habitat perhaps due to exclusion by newly evolved
New World Cleonini
585
competitively superior species, or due to the changing nature and ephemeral status of habitats
to which they were initially adapted. Raven and Axelrod (1978) note that among plants, a
number of Tertiary relicts have survived in California owing to its sheltered and equable
climate.
Grassland habitat only reached its present form and more extensive distribution during the
Miocene and early Pliocene (but especially at the Miocene-Pliocene transition) (Axelrod, 1950,
1985; Webb, 1977). Because of this, it is unlikely that divergence between A. aztecus and the
ancestor of the remainder of Apleurus took place before this time. Restriction of A. aztecus to
mesquite-grasslands, and primary association of A. lutulentus with mesquite-grasslands and
grasslands, suggests that the ancestor of the subgenus Apleurus , like the ancestor of the genus
Apleurus as noted, was also associated with grassland-like habitats. Present restriction of A.
aztecus to central Mexico is unlikely to have been a result of competitive exclusion by A.
lutulentus in the grasslands of the United States, for A. lutulentus coexists with A. aztecus in
central Mexico (Figs. 207, 210). Rather, a late Miocene or early Pliocene north-south
vicariance of Plains and Mexican Plateau grasslands as a result of increasing aridity and
development of more extensive semi-desert habitats (Webb, 1977) can account for the
distribution, with a subsequent dispersal of A. lutulentus south through desert-grassland
transitional habitat into central Mexican mesquite-grasslands and the Hidalgan Desert region
(term of Shreve, 1940; Axelrod, 1979) of a later Pleistocene origin. This later dispersal route
may have been through a desert-grassland transitional habitat, not grassland or
mesquite-grassland, because of the absence of concomitant dispersal north by A. aztecus.
Alternatively, lack of dispersal by the latter species may be due to individuals being flightless
whereas those of A. lutulentus are macropterous and presumably capable of flight. Webb
(1977) has drawn attention to the role of central Mexico, noting that Miocene faunules from
this area play a focal role in the history of the semi-arid biota of North America. Similarly,
Cronquist (1978) notes that the tribe Heliantheae of the Asteraceae (Compositae herein) has
its present center of diversity in the arid highlands of central Mexico, suggesting that this area
has been important in the evolutionary history of the group.
Increased aridity during the middle Pliocene resulted in increased extent of semi-desert
habitats and likely the origin of the ancestor of the remainder of Apleurus in western
desert-grassland transitional habitats, and A. lutulentus in more easterly, primarily grassland,
habitats (Fig. 243).
Divergence of the A. porosus-A. jacobinus-A. angularis and A. saginatus-A. albovestitus
lineages is more problematical, for extant species of the two groups, especially A. angularis and
A. albovestitus , do not differ much in their habitat associations or distributions.
In the A. saginatus-A. albovestitus lineage, an east-west vicariance of desert-grassland and
semi-desert habitats respectively (Axelrod 1979, figure 4) as a result of late Quaternary aridity
following the increased precipitation of the late Pliocene and early Pleistocene (Axelrod, 1979)
likely resulted in the origin of A. saginatus and A. albovestitus respectively (Fig. 245).
Although A. albovestitus is presently a widespread species, examination of intraspecific
variation shows the most plesiotypic form to be that from coastal Californian semi-desert
(possession of large glabrous abdominal patches, deeper and larger punctation; see “Geographic
variation” of A. albovestitus). Presence of other forms of A. albovestitus to the east in other
semi-desert and desert habitats is therefore suggestive of recent dispersal followed by
post-glacial local or regional selection pressures in various regional habitats which are
hypothesized to be largely of a recent origin in the late Quaternary (Spaulding et al., 1985).
Quaest. Ent., 1987,23 (4)
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Unlike A. albovestitus , A. saginatus has not expanded its range and is still found only in the
desert-grassland transition bordering the eastern portion of the Sonoran desert (Fig. 206).
In the A. porosus-A. angularis-A. jacobinus lineage, patterns of vicariance (Fig. 244)
among the three species are clearly evident for they are allopatric (Figs. 208, 210-211).
Because it has only been during the last four million years that Baja California reached its
present position and acquired a notably drier climate and coastal foggy desert (Axelrod, 1979),
origin of A. porosus does not likely predate that time. Increased aridity in the area likely
allowed entry of a desert-grassland-adapted ancestor from the north, as Baja California was
reaching its present position. This form is presumed to have evolved in situ into A. porosus. An
east-west vicariance of desert-grassland and semi-desert-adapted populations of the ancestor of
A. jacobinus-A. angularis, as in the A. albovestitus-A. saginatus lineage, likely accounts for
the origin and present distribution of A. jacobinus and A. angularis . Present distributions of A.
porosus , A. jacobinus, and A. angularis may be maintained as allopatric by competitive
exclusion or by selection pressures promoting reproductive isolation.
For Apleurus then, increasing aridity and concomitant changes in habitat structure and
availability appears to be the scenario most consistent with ideas and evidence about
paleoenvironments and their distribution in space and time in southwestern North America.
In Cleonidius, the pattern is somewhat different. Cleonidius species, in general, appear to
parallel some of the aspects of evolution of Apleurus species or lineages, but differ markedly in
various others.
As mentioned, at least one species of Cleonidius is known from the Palearctic Region.
However, its relationships are at present unclear (see Phylogeny section).
Unlike Apleurus , Cleonidius does not appear to be North American in origin because the
Palearctic species component of Cleonidius appears to be sister to the Nearctic species
component (Fig. 246). Thus, a Palearctic origin for the genus is to be preferred with subsequent
entry into North America of this ancestral stock. It is most likely that a trans-Beringian
grassland or savanna vicariance of this stock (Fig. 246), such as that proposed for the origin of
Apleurus, resulted in the evolution of Nearctic Cleonidius. The evidence for this is as follows:
1, Palearctic Cleonidius are eastern in their distribution and are apparently associated with
grassland or steppe habitat; 2, association with grasslands is widespread in Cleonidius
(although not restricted to primitive lineages or species) (Fig. 240); 3, association with
woodland, although widespread, is only in more derived species (Fig. 240) (suggestive of
secondary entry into that habitat type, a hypothesis supported by patterns of host plant
association); and 4, the two eastern North American species are unrelated and also of a
generally more derived nature. This trans-Beringian savanna or grassland event is likely to have
occurred only post-late Eocene.
Within Cleonidius there are five of what I consider to be main, perhaps interrelated,
differences from Apleurus: 1, species of Cleonidius are known from western xerophytic conifer
woodland (Fig. 240); 2, species of Cleonidius are known from eastern North America (Figs.
215, 229); 3, species exhibit primary host plant associations with one of a various number of
plant families (Fig. 241); 4, habitat association overlap (Fig. 240), altitudinal range overlap
(Fig. 233), and association with less arid habitats are high; and 5, comparison of habitat
associations or altitudinal ranges with the reconstructed phylogeny does not demonstrate a
clear correlation with increasingly arid habitats (Fig. 240) or with lower altitudes, respectively.
These differences are certainly suggestive of a different evolutionary history, one influenced
less so by shifts to increasingly arid habitats, but rather by host plant shifts, allopatric
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587
speciation within a habitat type, and habitat shifts to both more and less arid types at both
lower and higher elevations over a more extensive geographic range.
There are, however, some similarities. Distributions of three of the four species groups can
be geographically centered in certain areas. The C. erysimi group has five of six species
distributed in whole or in part in California (Fig. 257); the C. americanus group has six of
seven species distributed in whole or in part in the southern Rocky Mountain region (Fig. 258);
and, the C. boucardi group has three of four species in the central to southern Rocky Mountain
region or central Mexico (the most primitive being in central Mexico) (Fig. 259). With certain
restrictions, namely the question of the origin of the C. poricollis group, this pattern brings to
mind the California-central Mexico-southern Rocky Mountain pattern in primitive Apleurus
(Fig. 242) because the common ancestor of the C. boucardi group (Mexico) and the C.
poricollis group (questionable origin) is sister to the C. americanus group (southern Rocky
Mountains), the common ancestor of which is sister to the C. erysimi group (California).
This pattern is suggestive of similar historical processes accounting for diversification in the
basal lineages of the two genera, although it is possible that in Cleonidius the ancestral habitat
association is with sclerophyll woodland rather than grasslands or savanna for the two were
undoubtedly intermixed (Axelrod 1979). As such, an Eocene to Miocene vicariance of
California, southern Rocky Mountain region, and central Mexico was likely responsible for the
initial diversification within the genus (Fig. 246).
Parsimony, however, dictates preference for an association with grasslands or savanna
habitat because most C. americanus group members have a primary association with that
habitat type; C. poricollis and C. erysimi group members, although not primarily associated
with grasslands, are not primarily associated with woodlands; and primitive C. boucardi
members are associated with grasslands (Fig. 240). Furthermore, the two species associated
with western xerophytic conifer woodland and the single species associated with eastern North
American mesophytic evergreen forest, are more or less structurally derived and not relictual in
distribution. Similarly, the other eastern North American species is also structurally derived,
and although restricted in distribution, is the sister-species of a structurally similar western
North American species (Fig. 236).
The best evidence, however, for association with xerophytic woodland being secondarily
derived concerns host plant associations. Cleonidius texanus and C. quadrilineatus are both
associated primarily with Rosaceae and Rhamnaceae and each have their two closest relatives
associated exclusively with Leguminosae (Fig. 241). This, and presence of infrequent
association of C. texanus and C. quadrilineatus with Leguminosae (but not of relatives on
Rosaceae or Rhamnaceae), suggests Leguminosae was the plesiotypic host plant for these
lineages and that Rosaceae and concomitantly association with xerophytic woodland, derived.
Indeed, the widespread distribution and abundance of species associations with
Leguminosae suggests that association with this family, or an ancestral form of this family,
Rosaceae and Rhamnaceae (the families are phylogenetically closely related and placed in the
order Rosales [Cronquist, 1968]), may be ancestral for Cleonidius as a whole. Even more
parsimonious, however, is the hypothesis that association with Compositae is ancestral for the
genus. Association with this family is equally widespread and abundant in the genus, but unlike
Leguminosae, is found particularly in plesiotypic species in the C. erysimi , C. americanus , and
C. poricollis species groups, not in the more derived species, as is the association with
Leguminosae. Association with Compositae is also known in numerous other Cleoninae (e.g.,
Apleurus , Cleonis, Cyphocleonus, some Lixus ). Accordingly, preference must herein be given
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Anderson
to a plesiotypic association with Compositae, but final resolution of these competing hypotheses
rests largely on determination of host plant relationships in those Cleonidius species in which
they are as yet unknown, particularly, those of a primitive phylogenetic position and especially
those of the Palearctic species component.
Further supportive of the more recent trans-Beringian vicariance of savanna or grassland
are: 1, non-relictual geographic or structural status of any Cleonidius', 2, lack of extensive
structural divergence between species perhaps indicative of less available time for evolutionary
change to have occurred; and 3, likelihood that C. vibex and relatives are the sister group to
Nearctic Cleonidius (see Phylogeny section).
Axelrod’s (1975) Madrean-Tethyan hypothesis has not been given much support from other
studies. Floras of the eastern Gulf Coast and southeastern North America do not show evidence
of arid-adapted taxa (Graham, 1965). The hypothesis also does not receive any clear support
from the biogeographic analyses presented herein. Clearly, I believe we must accept an
ancestral association with grassland or savanna, as in Apleurus. Diversification in Cleonidius,
however, has apparently been much more complex than in Apleurus as evidenced by: lack of a
clear correlation between phylogeny and habitat association (Fig. 240); numerous different and
variable habitat associations of the species (Fig. 240); different host plant associations (Fig.
241); and present relatively extensive species sympatry and altitudinal range overlap (Fig.
233).
Within species groups, caution must be exercized because degree of confidence in various of
the hypothesized relationships is low (Fig. 237). Within the C. erysimi group, monophyly of C.
subcylindricus-C. texanus-C. longinasus is strongly supported by three primary characters.
Relationships of this lineage and the remaining three species in the species group are however
much less clear and are left as an unresolved trichotomy (Fig. 237). Assuming, as noted, that
the C. erysimi species group evolved in California (Fig. 246), it is likely that, as conditions
became more arid, there was an entry into semi-desert habitat in the C. eustictorrhinus-C.
erysimi lineage (Fig. 247). The hypothesized ancestral association with Compositae was
retained in this lineage and C. pleuralis, and the ancestor of the C. subcylindricus-C.
texanus-C. longinasus lineage likely shifted onto Leguminosae. C. eustictorrhinus and C.
erysimi may then have diverged in part as a result of an east-west vicariance between
California and the Great Basin Region with subsequent dispersal of C. erysimi back into
California (Fig. 248). This event likely also involved, or may have even been exclusively due to,
a host plant shift involving Compositae and the phylogenetically unrelated (Cronquist, 1968)
Cruciferae. Evidence for this is as follows. The sister-species C. americanus and C. frontalis (of
the C. americanus group) have similar geographic distributions to C. eustictorrhinus and C.
erysimi', C. americanus is associated with Compositae in California; host plant associations of
C. frontalis are not known. In C. eustictorrhinus and C. erysimi the opposite is true; the
widespread C. erysimi is found on Cruciferae whereas plant associations of the California C.
eustictorrhinus are not known. Furthermore, in the C. poricollis group, both C. poricollis and
C. calandroides are associated with Cruciferae, but C. poricollis is also markedly associated
with Compositae. Patterns of Cruciferae-Compositae association between closely related taxa
therefore (and, as noted, Leguminosae-Rosaceae/Rhamnaceae associations of a similar
nature), appear to repeat within Cleonidius (Fig. 241) and may thus represent more general
patterns of host plant relationships. In view of this, association with Compositae can be
hypothesized for C. eustictorrhinus, and association with Cruciferae for C. frontalis. If C.
pleuralis ultimately proves sister-species of the C. eustictorrhinus-C. erysimi lineage then
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589
evidence for C. eustictorrhinus being associated with Compositae is more marked, because then
only one shift from Compositae in the evolution of the three species is required. Within C.
pleuralis there has been a continued adaptation to more markedly arid lands (Fig. 240),
perhaps interrelated with association with Compositae (note relatively high level of association
with arid habitats in Apleurus, all of which are on Compositae; association of C. poricollis, the
only other Cleonidius that is associated with desert habitats, is also with Compositae).
In the C. subcylindricus-C. texanus-C. longinasus lineage there appears to be independent
entry into xerophytic and mesophytic evergreen woodlands in the central-southern Rocky
Mountain region and eastern North America by C. texanus and C. subcylindricus respectively
as evidenced by the pattern of host plant associations. An initial vicariance in this lineage
between C. subcylindricus in eastern North America and the C. texanus-C. longinasus lineage
in western North America, lineages both ancestrally on Leguminosae, was likely followed by a
host plant shift from Leguminosae to Rosaceae and Rhamnaceae and concurrent shifts to
xerophytic woodland habitat by C. texanus and to mesophytic woodland habitat by C.
subcylindricus (Fig. 249).
Within the C. americanus group, most of the species are primarily associated with
grasslands; C. americanus and C. notolomus are associated exclusively with Pacific semi-desert
and desert-grassland transition respectively (Fig. 240). Vicariance patterns are clearly evident
in the present distributions of species in this group. A California-Great Basin or southern Great
Plains vicariance can be proposed to account for the origin of the C. americanus-C. frontalis
lineage and the ancestor of the remainder of the C. americanus group (Fig. 250). Within the
former lineage, a California-inland vicariance again very likely explains the evolution of C.
americanus and C. frontalis (Fig. 251). This was likely concomitant with a shift to more arid
habitats, especially by C. americanus.
Origins of C. canescens and of the C. notolomus-C. collaris and C. puberulus-C. infrequens
lineages are not clear. However, a north-south vicariance of both C. collaris and C. notolomus
(Fig. 253), and of C. puberulus and C. infrequens (Fig. 252), accounts for the present
distribution of these species. Of the four, only C. notolomus has changed from a grassland
association to desert-grassland transitional habitat (Fig. 240). Few structural differences
between these sister-species, and the allopatric distributions favour a relatively recent
occurrence of these speciation events. Association with Compositae is hypothesized as
plesiotypic for the species group.
Origin of the C. poricollis group is unclear principally because of the widespread
distribution and eurytopic nature of C. poricollis (Fig. 228). It was undoubtedly concurrent
with the hypothesized initial radiation of species group lineages and while an east coast
isolation of C. calandroides likely accounts for the origin of that species, it is not clear where
the ancestor of C. poricollis and C. calandroides evolved. Association with Compositae is
proposed as ancestral for this species group as well.
In the C. boucardi group, there was likely a Mexican highlands isolation of C. boucardi in
mesquite-grassland and a southern Rocky Mountain isolation of the C. trivittatus-C.
quadrilineatus-C. placidus lineage in grassland habitat, an event brought about by increasing
aridity and the formation of intermediate desert-grassland transitional or semi-desert habitats
(Fig. 254). In the C. trivittatus-C. quadrilineatus-C. placidus lineage, in which association was
likely with Leguminosae, an east-west vicariance of C. trivittatus and C. quadrilineatus-C.
placidus in grassland and semi-desert habitats respectively, appears to account for the origin of
C. trivittatus (Fig. 255). Subsequent entry into xerophytic evergreen woodland habitat by the
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Anderson
semi-desert adapted ancestor, which, as in the C. erysimi group, continued to be associated with
Leguminosae, resulted in C. quadrilineatus in the eastern xerophytic woodlands and C.
placidus in the semi-desert habitat of California (Fig. 256). Concurrent with this habitat shift
by C. quadrilineatus was a host plant shift to Rosaceae and Rhamnaceae; C. placidus
continued to be associated with Leguminosae. Present occurrence of C. quadrilineatus in
California is likely a recent dispersal event. Ancestral host plant relationships of this group are
also likely with Compositae.
Summarizing, both Apleurus and Cleonidius are hypothesized to have entered North
America in savanna or grassland habitats (or their precursors) across Beringia during late
Eocene time. In both genera there appears to be an initial association with Compositae and
vicariance of lineages in grassland or mesquite-grassland between California, central Mexico,
and the southern Rocky Mountains. Further speciation event in Apleurus appear due primarily
to shifts to apotypic habitat types brought about by increased aridity during the later Tertiary;
there is little within habitat-type allopatric speciation. Host plant shifts are not a factor, for all
Apleurus species remain associated with Compositae. Host plant shifts, however, primarily
between Leguminosae-Rosaceae and Compositae-Cruciferae, and, to a lesser degree,
within-habitat-type allopatric speciation, appear to predominate as the causes of further
speciation in Cleonidius, most species of which remain associated with plesiotypic less-arid
habitat types such as grasslands and desert-grasslands.
Evolutionary trends and adaptations
Adaptive trends to arid conditions
Numerous apparent adaptations to arid habitats are known in Coleoptera, including
Curculionidae (Crowson, 1981). Some of these are exhibited in Cleoninae.
Wing length and flightlessness. — Adults of various species of North American Cleoninae
are apterous or brachypterous to varying degrees and in varying frequency. Some, such as
Stephanocleonus confusus or Apleurus aztecus, have the elytra more or less solidly fused
together along the median suture. The cuticle is also thick and hard in many of these same
individuals. Flightlessness appears more frequenty in traditional Cleonini than in traditional
Lixini.
In Apleurus no clear phylogenetic trend is evident in this character although adults of the
closely related A. angularis , A. jacobinus and A. porosus are all more or less apterous. Lack of
a trend in Apleurus is perhaps surprising in view of the correlation of phylogenetic position with
occupation of increasingly arid habitats (Fig. 238). Similarly, in Cleonidius, there is no clear
phylogenetic trend in this character. Adults of species of the C. americanus group are
predominantly brachypterous but they are mostly associated with grassland habitats and not
more arid habitats. Undoubtedly, degree of aridity is not the sole Undoubtedly, degree of
aridity is not the sole factor influencing the frequency of brachyptery and although the
aforementioned adaptations may serve to prevent desiccation in arid environments, the spatial
distribution of the habitat-type or of host plants, likely also influences the tendency towards or
away from brachyptery. Compromise between these differing selective forces likely results in
the wing-length polymorphisms noted in some species and the lack of a clear trend.
In species in the southwestern United States, populations on both sides of the Continental
Divide, in which individuals are flightless, are structurally more distinct in the north, where the
divide is high in elevation, as compared to the south, where the divide is low in elevation and
where gene flow seems likely. Species exhibiting similar distributions but which are composed
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591
of macropterous individuals do not show such patterns of variation.
Diel activity. — There is no evidence to indicate distinct peaks of diel activity in North
American Cleoninae which might indicate avoidance of diurnal stressful conditions in an arid
habitat. Such diel activity is known in other arid land Curculionidae (C.W. O’Brien, E.L.
Sleeper, pers. comms.).
Burrowing adaptations. — Modifications of legs for burrowing are not known in North
American Cleoninae. This is not surprising in view of 2, above. There are, however, adults of
some species that possess long erect surface vestiture which may prove correlated with life on
sand dunes and which may facilitate movement through sandy substrates. Such long vestiture is
known in various obligate sand dune Scarabaeidae and other Curculionidae.
Ventral tarsal pilose vestiture. — Within Apleurus (Figs. 234-235), and to a lesser extent
also in Cleonidius (Figs. 236-237), a clear trend is evident towards reduction in ventral tarsal
pilose vestiture. This reduction has been reasonably suggested to be due to living predominantly
on the ground rather than on plants (Kissinger, 1964; Crowson, 1981) and is widespread in
many arid-adapted Curculionidae. In Apleurus the trend is as expected, because those more
derived species in more arid environments, where plants are more widely spaced, and where it is
less desirable to spend time exposed on the plant, lack ventral tarsal pilose vestiture or have
pilose pads that are markedly reduced. In Cleonidius however, this is not clearly so, for
although more arid habitat species ( e.g ., C. erysimi ) possess reduced pilose pads, it is the
members of the C. americanus group which exhibit the greatest frequency of lack of ventral
tarsal pilose vestiture. This is perhaps surprising, for the species in this group are associated
principally with plesiotypic grassland habitats; predicted adaptive correlates are thus not
always evident in Cleonidius.
Other evolutionary trends
Host plant associations. — No trends in host plant relationships are evident in Apleurus,
because all species appear associated with Compositae (Fig. 239). In Cleonidius however, two
repeated patterns warrant mention (Fig. 241). Leguminosae-Rosaceae/Rhamnaceae
associations are found in the sister-species C. longinasus-C. texanus and C. placidus-C.
quadrilineatus, and Compositae-Cruciferae associations are found in C. poricollis-C.
calandroides and are hypothesized in C. americanus-C. frontalis and C. eustictorrhinus-C.
erysimi. It is unlikely that the Compositae-Cruciferae association is phylogenetically based, but
rather reflects the co-occurence (perhaps recent) of these two plant taxa in the same, more arid
habitats. The Leguminosae-Rosaceae/Rhamnaceae association, however, is likely
phylogenetically based for the taxa involved are closely related and the host plant shift is
concomitant with a shift in habitat association.
Groups in which sympatry is low, altitude range overlap low, and habitat association overlap
low, tend to have species which differ little in host plant relationships. Increasing levels of these
factors appear to be correlated with more varied host associations. Host plant associations,
largely unknown in the C. americanus group, can thus be predicted to vary little in terms of
their taxonomic range and thus largely be with Compositae or Cruciferae.
There is no trend from generalist to specialist feeding in either Apleurus or Cleonidius.
Habitat associations. — As noted in detail in the biogeography section, there is a clear trend
in Apleurus for phylogenetically more derived species to be associated with increasingly arid
habitats (Fig. 238) and lower elevations. This is not so in Cleonidius wherein most species are
associated with grassland or grassland-desert habitats (Fig. 240).
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Size. — Within Apleurus there is a tendency towards increased size (Figs. 234-235); such is
not clearly evident in Cleonidius. Increased size in an arid environment may serve to aid in
prevention of water loss for surface area to volume ratios would be lower in larger individuals
which are, notably, found in more arid environments. In Cleonidius however, the largest species
are those in the C. boucardi group. These are not found in notably arid habitats but rather
grasslands, semi-desert and xerophytic evergreen woodlands.
Genitalia. — Form of male genitalia including the internal sac is very conservative in species
of both Apleurus (Figs. 99-104) and Cleonidius (Figs. 177-196) and form of female genitalia
is very conservative in Cleonidius (Figs. 157-176). Such is not so in females of Apleurus
wherein there is a clear trend towards reduction in size of the stylus on gonocoxite II and an
increasingly subapical position of attachment of the same (Figs. 89-96). This trend is likely
related to the oviposition habits of the species but such are not known in detail for any North
American species of Cleoninae.
Form of the aedeagus (Figs. 31-36) and of the internal sac (Figs. 43-48) differs markedly
between species of Stephanocleonus, in contrast to the lack of differences in Cleonidius and
Apleurus. Reasons for this are unknown.
Elytral scale patterns. — Elytral scale patterns are generally uniform within and between
Apleurus species except for A. angularis (Figs. 72-74) and A. albovestitus (Figs. 75-79)
wherein various forms are recognizable. Correlation of particular scale patterns with
geographic distribution suggests a local adaptive significance, one I believe associated with
crypsis on the ground substrate or in debris under host plants. A thermoregulatory role
however, may apply to those A. albovestitus in the lower Colorado River drainage. In Apleurus
there are no species which possess a distinctly vittate scale pattern; most are more or less
mottled in general appearance.
In Cleonidius elytral patterns vary between species but not to a significant extent within a
species (with the exception of C. canescens ). Adaptive significance of the vittate pattern ( e.g .,
Fig. 131) found in most Cleonidius is likely one of concealment on host plants or in debris
under the host plant. Crypsis on the ground substrate is not likely, except perhaps in the C.
americanus group, for otherwise, most species have moderately developed ventral tarsal pilose
vestiture indicating life predominantly on plants. Only in the C. americanus group is there
predominance of species in which individuals have reduced ventral tarsal pilose vestiture. Also
in this group are found the only species with mottled elytral scale patterns {e.g., Figs. 124-125).
These patterns are similar to those of Apleurus species, and likely serve as an adaptation to life
on the ground substrate.
Vittate (and margined [Figs. 118-123]) patterns predominate in species of Cleonidius with
extensive ventral tarsal pilosity; non-vittate patterns predominate in species with reduced tarsal
vestiture. Wide ranging species with reduced ventral pilose tarsal vestiture in Apleurus exhibit
geographic variation in scale pattern, whereas restricted species or those wide-ranging species
with extensive ventral pilose vestiture do not show variation in elytral scale pattern. In
Cleonidius nearly all species exhibit little or no variation in elytral scale pattern (except for C.
canescens of the C. americanus group, which like A. angularis , has more or less vittate and
non-vittate forms occurring on the west and east sides of the Continental Divide). Within the C.
americanus group, which contains mostly allopatric species with reduced ventral vestiture,
there is the most extensive within-species-group variation in scale pattern. Other species groups
do not show within group variation in scale pattern. It thus appears that there is an inverse
correlation in both Apleurus and Cleonidius between degree of variation in elytral pattern and
New World Cleonini
593
extent of ventral tarsal pilose vestiture, but only seen in species or species groups distributed
over large geographic areas.
Patterns and predictions
From results of this study of Cleoninae, some tentative generalizations can be made as
regards arid adapted faunas in North America.
1 . Preference must go to a post-late Eocene vicariance of savanna or grassland habitat, or
precursory habitat, across Beringia as the original mode of isolation of Nearctic arid land taxa
which possess Palearctic relatives also inhabiting arid lands (Figs. 242, 246). There is no
evidence for direct Madrean-Tethyan connections as proposed by Axelrod (1975).
2. The most plesiotypic species or lineages in the Nearctic region will be found on
Compositae in various combinations of the mesquite-grasslands of the highlands of central
Mexico, the southern Rocky Mountain grasslands, and in one of various habitats in California
which may be relictual or predominantly grassland in nature (Figs. 242, 246).
3. Arid-adapted species in eastern North America are more recent in origin, not relictual,
and are related to a species or a lineage of few species in western North America.
4. California is the geographical area recognizable as possessing the most arid land
endemics. All have their closest relatives outside of California.
5. Evolution in arid-adapted taxa may involve a trend toward occupation of increasingly arid
habit by successively more derived species (Fig. 238). This is accompanied by few or no host
plant shifts (Fig. 239), and may be limited to non-speciose groups. Sympatry, altitudinal range
overlap, and habitat association overlap is low in these taxa.
6. Alternatively to 5 above, host plant shifts (Fig. 241) permit sympatry, altitudinal range
overlap and habitat association overlap and are perhaps associated only with more speciose
groups. No clear correlation between phylogenetic position of the species and degree of aridity
of habitat occupied is evident (Fig. 240).
7. Species inhabiting North American desert habitats are of recent origin (but not
necessarily of derived phylogenetic position), and are related to taxa associated with precursory
North American non-desert habitats although not in a generally predictable manner. That
regional deserts are composed of taxa from non-desert arid habitats and are of recent origin is
supported. No regional desert endemics are known in Apleurus or Cleonidius .
8. Contrary to statements of other authors (Axelrod, 1985), grassland endemics are known,
are primitive or derived in their phylogenetic position, and should thus be expected in other arid
land adapted taxa.
9. Compositae-Cruciferae and Leguminosae-Rosaceae/Rhamnaceae host plant shifts may
represent general patterns to be found in other phytophagous taxa that feed on vegetative plant
structures. Shifts from Leguminosae to Rosaceae and Rhamnaceae may prove generally
associated with entry into western upland xerophytic evergreen woodland habitat by some
phytophagous taxa. These shifts appear to have a phylogenetic basis as indicated by the fact
that the families are placed in the order Rosales (Cronquist, 1968).
In contrast, Compositae-Cruciferae host plant shifts are likely due to occurrence of both the
two plant and two weevil taxa in the same habitat types. No phylogenetic basis can be inferred
because the two families are not apparently phylogenetically closely related as indicated by
their placement in separate, unrelated subclasses (Cronquist, 1968).
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CONCLUSIONS AND FUTURE RESEARCH
Scientific studies should not only attempt to answer specific questions but should also strive
to pose questions which might prove the basis for further inquiry. More than anything else, I
believe this study draws attention to the need for examination of world faunas or at the least,
regional faunas placed in a world perspective. Traditionally, North American Cleonini have all
been treated as Cleonis , but as now should be clearly evident, the relationships of many taxa in
North America are actually quite phylogenetically distant.
Relationships of Apleurus are largely resolved. Unfortunately, this cannot be said with the
same confidence for Cleonidius. World study of the speciose and structurally diverse genus
Li xus should help resolve the still problematical relationships and even species constituency of
Cleonidius. What other Old World Lixus may require placement in Cleonidius and subsequent
study of large numbers of individuals of these species and determination of what their
relationships are to North American Cleonidius , should help to resolve the phylogenetic
placement of the Old World Cleonidius , further resolve the interrelationships of North
American species, and answer some still remaining questions of biogeographic importance. This
will be a large task, for within Lixus are placed a great number of species. More complete
knowledge of structural diversity of Lixus species and the phylogenetic relationships of
component species-groups or subgenera should ultimately lead to a clearer understanding of
suprageneric relationships in Cleoninae and a well substantiated tribal classification.
Not to be forgotten are the interrelationships among genera of traditional Cleonini. A start
at a study of these relationships leading, I hope, to a generic reclassification, has been made as
part of this present study. Results are not presented in detail here; only characters of relevance
to our understanding of the North American fauna have been included. Study of
representatives of generic-group names not yet examined is required, as is study of more species
in more speciose genera to assure proper placement and uniformity of features that appear to
prove phylogenetically valuable at the generic level.
Even within the North American fauna itself, work remains to be done. For Apleurus and
Cleonidius , I feel confident that no new species will be discovered; for Stephanocleonus ,
largely because the genus is northern and Holarctic, I suspect that one or more new species
remain to be discovered, perhaps in the Yukon Territory and Alaska. Whether any of the North
American species are Holarctic in distribution (and perhaps conspecific with nominal Old
World species) also needs to be determined for it has not been possible to examine types or
representatives of even a small number of the many Palearctic species. Only when this has been
done can phylogenetic relationships of the species in this genus be reconstructed.
For all Cleoninae, more host plant information needs to be compiled. Hosts, even plant
associations, are not known for Stephanocleonus species; the relationship with Compositae
needs to be further substantiated in Apleurus species; and, various of the species of Cleonidius
do not have hosts or even plant associations established, and those that are established need
variable degrees of further supportive data. With host plant shifts appearing to play an integral
role in Cleonidius , more such information can only increase understanding of the evolution of
species in that genus and determine whether predictions about specific host plant relationships
made herein, and whether predictions that these patterns are repeated and general are borne
out.
Immature stages of Cleoninae are largely unknown and unstudied. Collection and study of
eggs, larva and pupae, particularly in North America, may provide characters of phylogenetic
New World Cleonini
595
use in substantiating relationships as based on adult characters, or, in resolving problematical
relationships or those in which confidence is low.
ACKNOWLEDGMENTS
First and foremost I wish to thank George E. Ball, for his criticism, advice and support
during the course of this study. The academic environment for systematic studies at the
University of Alberta, created largely by his enthusiasm and dedication, was a pleasure to work
in. In addition, financial aid from Natural Sciences and Engineering Research Council grant
A- 1399, held by G.E. Ball, is also gratefully acknowledged.
For advice in selecting a thesis topic and for their assistance and guidance at various stages
and with various aspects of the study, I thank D.R. Whitehead, C.W. O’Brien, E.L. Sleeper,
H.R. Burke, W.E. Clark, and A.T. Howden; they were always available for consultation. R.T.
Thompson was especially helpful in finding specimens and literature during my visit to the
British Museum (Natural History). His help in solving nomenclatural problems and in
demonstrating his novel preparatory techniques is also very gratefully acknowledged.
I also thank members of my advisory committee (M.V.H. Wilson, D.A. Craig, R.A.
Stockey) and external thesis examiner (W.E. Clark) for their critical review of the penultimate
draft of the thesis on which this paper is based. For help in finding collecting localities in
Arizona and for companionship in the field, I thank Scott McCleve of Douglas, Arizona. For
permission to collect in Big Bend National Park, Texas, I thank C.M. Fleming of that same
facility.
Finally, I thank all my fellow students at the University of Alberta (D.C. Currie, J.M.
Cumming, G.A.P. Gibson, J.E. 0‘Hara, D.R. Maddison, J.S. Ashe, R.E. Roughley, J.-F.
Landry, J.H. Acorn) for their friendship at home, companionship in the field, for their constant
encouragement, but most of all, for expressing a keen interest in problems in weevil systematics
and biogeography when called upon for their help and comments. Special thanks are also due
A. Sen for preparation of the photographic plates and J.E. 0‘Hara for assistance in preparation
of the revised manuscript for publication.
Additional financial support during the course of this study came from The University of
Alberta, (Dissertation Fellowship, 1985-1986), and the Government of the Province of Alberta
(Province of Alberta Scholarships, 1983-1984, 1984-1985).
I offer a last word of fond appreciation to my parents Robert Hamilton and Daisy, and to
my wife Catherine; you were there when I needed you in ways you didn’t realize and I can’t
explain.
LITERATURE CITED
Anderson, D.M. 1956. Notes on Cleonus piger (Scop.) in the United States (Coleoptera;
Citrculionidae). The Coleopterists' Bulletin 10:81-85.
Anderson, E.M. 1914. Insects collected in the Okanagan Valley, 1913. Proceedings of the
Entomological Society of British Columbia 4:54-58.
Anderson, R.S. 1984a. Taxonomic studies of Cleonis piger (Scop.) and Cyphocleonus achates
(Fahrs.) (Coleoptera: Curculionidae;Cleoninae) from Centaurea as biological agents for
control of Centaurea diffusa (Compositae) in Canada. Unpublished Report for Agriculture
Canada, Regina, Saskatchewan. 49 pp + 8 figures.
Quaest. Ent., 1987, 23 (4)
596
Anderson
Anderson, R.S. 1984b. Connatichela artemisiae, a new genus and species of weevil from the
Yukon Territory (Coleoptera: Curculionidae: Leptopiinae): Taxonomy, paleontology, and
biogeography. The Canadian Entomologist 116:1571-1580.
Anderson, R.S. and S.B. Peck. 1986. Geographic patterns of colour variation in North
American Nicrophorus burying beetles (Coleoptera; Silphidae). Journal of Natural
History 20:283-297.
Andrews, F.G., A.R. Hardy, and D. Giuliani. 1979. The coleopterous fauna of selected
California sand dunes. California Department of Food and Agriculture, Insect Taxonomy
Laboratory, i + 142 pp. Sacramento, California.
Arnaud, P.H. 1978. A host-parasite catalogue of North American Tachinidae (Diptera).
United States Department of Agriculture, Miscellaneous Publications 1319. 860 pp.
Arnett, R.H., Jr. 1960-1962. The beetles of the United States (a manual for identification).
Catholic University Press, xi + 1 1 12 pp. Washington D.C..
Arnett, R.H., Jr., N.M. Downie, and H.E. Jaques. 1980. How to know the beetles (2nd
edition). The Pictured Key Nature Series. Wm. C. Brown Co., viii + 416 pp. Dubuque,
Iowa.
Aslam, N.A. 1961. An assessment of some internal characters in the higher classification of the
Curculionidae s.l. (Coleoptera). The Transactions of the Royal Entomological Society of
London 113:417-489.
Aslam, N.A. 1963. On the genera of Indo-Pakistan Cleoninae and Hylobiinae (Coleoptera:
Curculionidae). Bulletin of the British Museum ( Natural History ), Entomology. 13:47-66.
Axelrod, D.I. 1950. Evolution of desert vegetation in western North America, pp. 215-306. In:
Axelrod, D.I. Contributions to Paleontology. Studies in Late Tertiary Paleobotany.
Carnegie Institute of Washington Publication 590, Washington D.C. 1-323 pp.
Axelrod, D.I. 1975. Evolution and biogeography of Madrean-Tethyan sclerophyll vegetation.
Annals of the Missouri Botanical Garden 62:280-334.
Axelrod, D.I. 1979. Age and origin of Sonoran Desert vegetation. Occasional Papers of the
California Academy of Sciences 132:i-vi + 74 pp.
Axelrod, D.I. 1985. Rise of the grassland biome, Central North America. The Botanical
Review 51:1 63-20 1 .
Axelrod, D.I. and P.H. Raven. 1985. Origins of the Cordilleran flora. Journal of Biogeography
12:21-47.
Ball, I.R. 1975. Nature and formulation of biogeographical hypotheses. Systematic Zoology
24:407-430.
Bargagli, P. 1884. Rassegna biologica di Rincofori Europei. Bollettino della Societa
Entomologica Italiana 16:149-258.
Batra, S.W.T., J.R. Coulson, P.H. Dunn, and P.E. Boldt. 1981. Insects and fungi associated
with Carduus thistles (Compositae). United States Department of Agriculture. Technical
Bulletin 1616. 100 pp.
Bedel, L. 1882-1888. Faune des Colepteres du bassin de la Seine. Rhynchophora. Annales de
la Societe Entomologique de France 6: 1-442.
Berg, F.G.C. 1884. (Communication). Bulletin de la Societe Entomologique de France
1884:XCVIII-C.
Billberg, G.J. 1820. Enumeratio insectorum in museo Gust. Joh. Billberg. Stockholm. 138 pp.
Blackwelder, R.E. 1939. Fourth supplement 1933 to 1938 (inclusive) to the Leng Catalogue of
Coleoptera of America, north of Mexico. Mount Vernon, New York. 146 pp.
New World Cleonini
597
Blackwelder, R.E. 1947. Checklist of the Coleopterous insects of Mexico, Central America, the
West Indies and South America. Part 5. Bulletin. United States National Museum
(185)I-IV, 765-925.
Blanchard, C.E. 1851. Fauna Chilena. Insectos. Coleopteros. pp. 285-563. In: Gay, C. Historia
fisica y politica de Chile. Volume 5. Zoologia. Paris.
Blatchley, W.S. 1914. Notes on the winter and early spring Coleoptera of Florida with
descriptions of new species. The Canadian Entomologist 46:247-251.
Blatchley, W.S. 1930. Blatchleyana. A list of the published writings of W.S. Blatchley, A.B.,
A.M., LL.D. of Indianapolis, Indiana and Dunedin, Florida. Together with a chronology of
his life: the fixation of types of new genera and species described by him, etc. etc. 77 pp.
Indianapolis, Indiana.
Blatchley, W.S. and C.W. Leng. 1916. Rhynchophora or weevils of North Eastern America.
The Nature Publishing Company. Indianapolis, Indiana. 682 pp.
Bleasdell, G.G. 1937. The Rhynchophora of Iowa. Iowa State College Journal of Science
11:405-445.
Boheman, C.H. 1836. In: Schoenherr, C.J. Genera et species Curculionidum, cum synonymia
hujus familiae. Species novae aut hactenus minus cognitae, descriptionibus a Dom.
Leonardo Gyllenhal, C.H. Boheman, et entomologis aliis illustratae. Volume 3, Part 1, 505
pp. Paris.
Boving, A.G. 1927. Immature stages of Eumycterus (?) saccharides Barber, with comments on
the classification of the tribe Barini (Coleoptera: Curculionidae). Proceedings of the
Entomological Society of Washington 29:151-159.
Bradley, J.C. 1930. A manual of the genera of beetles of America, north of Mexico, x + 360
pp. Ithaca, New York. Mimeographed.
Brethes, J. 1910. Coleopteros argentinos y bolivianos. Anales de la Sociedad Cientifica
Argentina 69:205-227.
Brimley, C.S. 1938. The insects of North Carolina. Being a list of the insects of North Carolina
and their close relatives. North Carolina Department of Agriculture. 560 pp.
Brown, D.E. (ed.). 1982. Biotic Communities of the American Southwest-United States and
Mexico. Desert Plants 4:1-342.
Brown, J.H. and A.C. Gibson. 1983. Biogeography. The C.V. Mobby Company, St. Louis, i-xii
+ 643 pp.
Brown, W.J. 1940. Notes on the American distribution of some species of Coleoptera common
to the European and North American continents. The Canadian Entomologist 72:65-78.
Bruhn, A.F. 1947. The external male genitalia of some Rhynchophora. The Great Basin
Naturalist 8:1-35.
Buchner, P. 1933. Studien an Intracellularen Symbionten. VII. Die Symbiontischen
Einrichtungen der Russelkafer. Zeitschrift fur Morphologie und Okologie der Tiere.
26:709-777.
Buchner, P. 1953. Endosymbiosis of animals with plant microorganisms. Revised English
Version. Interscience Publishers, New York.
Burke, H.R. and D.M. Anderson. 1976. Systematics of larvae and pupae of American
Curculionoidea: Status report, historical review and bibliography. The Southwestern
Entomologist 1:56-73.
Capiomont, G. and M.C.-E. Leprieur. 1873. Monographic des Lixus. Deuxieme partie.
Annales de la Societe Entomologique de France 5:41-64.
Quaest. Ent., 1987, 23 (4)
598
Anderson
Casey, T.L. 1891. Coleopterological Notices. III. Annals of the New York Academy of
Sciences 6:9-214.
Casey, T.L. 1904. On some new Coleoptera, including five new genera. The Canadian
Entomologist 36:312-324.
Champion, G.C. 1902-1906. Biologia Centrali-Americana. Insecta. Coleoptera.
Rhynchophora. Curculionidae. Curculionidae (part). Volume 4, Part 4, viii + 750 pp.
Chevrolat, L.A.A. 1860. Description de Coleopteres nouveaux d‘Algerie. Revue et Magasin de
Zoologie, Series 2 12:75-82.
Chevrolat, L.A.A. 1873. Memoire sur les Cleonides. Memoires de la Societe Royale des
Sciences de Liege, Series 2 5(6):viii-l 18.
Chittenden, F.H. 1911. Some miscellaneous results of the work of the Bureau of Entomology -
IX. Insects injurious to the loco weeds. United States Department of Agriculture. Bureau of
Entomology. Bulletin. 64:33-42.
Clayton, W.D. 1981. Evolution and distribution of grasses. Annals of the Missouri Botanical
Garden. 68:5-14.
Cody, M.L. 1975. Towards a theory of continental species diversities, pp. 214-257. In: Cody,
M.L. and J.M. Diamond, Eds. Ecology and Evolution of Communities. Bellknap Press of
Harvard University, Cambridge, i-xiv + 545 pp.
Cody, M.L. and J.M. Diamond, Eds. 1975. Ecology and evolution of communities. Bellknap
Press of Harvard University, Cambridge, i-xiv + 545 pp.
Coope, G.R. 1979. Late Cenozoic fossil Coleoptera: evolution, biogeography, and ecology.
Annual Review of Ecology and Systematics. 10:247-267.
Correll, D.S. and M.C. Johnston. 1970. A manual of the vascular plants of Texas. Texas
Research Foundation. Renner, Texas, xvi +1881 pp.
Croizat, L. 1982. Vicariance/vicariism, panbiogeography, “vicariance biogeography,” etc. : a
clarification. Systematic Zoology 31:291-304.
Cronquist, A. 1968. The evolution and classification of flowering plants. Thomas Nelson and
Sons, London, xii + 396 pp.
Cronquist, A. 1978. The Biota of the intermountain region in geohistorical context. Great Basin
Naturalist Memoirs 2:3-15.
Crosby, C.R. and W.E. Blauvelt. 1930. A European beetle found in New York (Coleop.:
Curculionidae). Entomological News 41:164.
Crowson, R.A. 1955. The natural classification of the families of Coleoptera. Nathaniel Lloyd,
London. 187 pp.
Crowson, R.A. 1981. The biology of the Coleoptera. Academic Press. London, xii + 802 pp.
Csiki, E. 1934. Coleopterorum Catalogus, pars 134, Curculionidae: Subfam. Cleoninae, 152 pp.
Danks, H.V. 1981. Arctic arthropods. A review of systematics and ecology with particular
reference to the North American fauna. Entomological Society of Canada, viii + 608 pp.
Ottawa, Ontario.
Dejean, P.F.M.A. 1821. Catalogue de la collection de Coleopteres de M. le Baron Dejean, viii
+ 136 pp. Paris.
Eldredge, N. 1979. Cladism and Common Sense, pp. 165-198. In: Cracraft, J. and N. Eldredge,
Eds. Phylogenetic Analysis and Paleontology. Columbia University Press, New York. 1-233
pp.
Eldredge, N. and J. Cracraft. 1980. Phylogenetic patterns and the evolutionary process.
Columbia University Press, New York, ix + 349 pp.
New World Cleonini
599
Ely, C.R. 1913. The food plant of Cleonus calandroides Rand. Proceedings of the
Entomological Society of Washington 15:104-105.
Endler, J.A. 1977. Geographic variation, speciation, and clines. Monographs in Population
Biology 10. Princeton University Press, Princeton, i-ix + 1-246 pp.
Endler, J.A. 1982. Problems in distinguishing historical from ecological factors in
biogeography. American Zoologist 22:441-452.
Essig, E.O. 1958. Insects and mites of Western North America (revised edition). The
MacMillan Co., xiv + 1050 pp. New York, New York.
Everts, J.E. 1903. Coleoptera Neerlandica. De Schildvleugelige Insecten van Nederland en het
Aangrenzend Gebied. Tweede Deel Martinus Nijhoff. iv + 798 pp. + 8 pi. ‘Sgravenhage.
Fabricius, J.C. 1792. Entomologia systematica emendata et aucta. Secundum classes, ordines,
genera, species adjectis synonimis, locis, observationibus, descriptionibus. Volume 1. xx +
538 pp. Hafniae.
Fabricius, J.C. 1801. Systema eleutheratorum secundum ordines, genera, species: adiectis
synonymis, locis, observationibus, descriptionibus. Volume 2. 687 pp. Kiliae.
Fahraeus, O.I. von. 1842. In: Schoenherr, C.J. Genera et species Curculionidum, cum
synonymia hujus familiae. Species novae aut hactenus minus cognitae, descriptionibus a
Dom. Leonardo Gyllenhal, C.H. Boheman, et entomologis aliis illustratae. Volume 6, Part
2, 495 pp. Paris.
Fall, H.C. 1897. A list of the Coleoptera of the Southern California islands, with notes and
descriptions of new species. The Canadian Entomologist 29:233-244.
Fall, H.C. 1901. List of the Coleoptera of southern California, with notes on habits and
distribution and description of new species. Occasional Papers of the California Academy
of Sciences 8: 1-282.
Fall, H.C. 1913. A brief review of our species of Magdalis, with notes and descriptions of other
North American Rhynchophora. Transactions of the American Entomological Society
39:23-72.
Fall, H.C. and T.D.A. Cockerell. 1907. The Coleoptera of New Mexico. Transactions of the
American Entomological Society 33:145-272.
Faust, J. 1890. Notizen iiber Russelkafer. Stettiner Entomologische Zeitung 51:109-1 16.
Faust, J. 1904. Revision der Gruppe Cleonides vrais. Deutsche Entomologische Zeitschrift
1904:177-284.
Fletcher, J. 1906. The entomological record, 1905. 36th. Annual Report of the Entomological
Society of Ontario 1905:90-105.
Gaffney, E.S. 1979. An Introduction to the logic of phylogenyrreconstruction. pp. 79-111. In:
Cracraft, J. and N. Eldredge, Eds. Phylogenetic analysis and paleontology. Columbia
University Press, New York. 1-233 pp.
Gebler, F.A. von. 1830. Bemerkungen iiber die Insecten Sibiriens, vorziiglich des Altai.
Ledebours Reise 2(3): 1-228.
Gemminger, M. and E. von Harold. 1871. Catalogus Coleopterorum hucusque descriptorum
synonymicus et systematicus. Volume 8. Curculionidae. pp. 2181-2668. Monachii.
Gibson, A. 1914. The entomological record, 1913. 44th. Annual Report of the Entomological
Society of Ontario 1913:1 16-139.
Gmelin, J.F. 1790. Caroli a Linne Systema Naturae per regna tria naturae, secundum classes,
ordines, genera, species, cum characteribus. differentiis, synonymis, locis. Editio decima
tertia, aucta, reformata. Beer, Lipsiae. Volume 1, Part 4 (Coleoptera, Hemiptera). pp.
Quaest. Ent., 1987,23 (4)
600
Anderson
1517-2224.
Goeze, J.A.E. 1777. Entomologische Beitrage zu des Ritter Linne swelften Ausgabe des
Natursystems. Volume 1. 736 pp. Leipzig.
Goudge, T.A. 1961. The Ascent of Life. A philosophical study of the theory of evolution.
University of Toronto Press, Toronto. 236 pp.
Gould, S.J. 1985. The Flamingo‘s smile. Reflections in natural history. W.W. Norton and
Company. 476 pp. New York.
Graham, A. 1965. Origin and evolution of the biota of southeastern North America: evidence
from the fossil plant record. Evolution 18:571-585.
Gyllenhal, L. 1834. In: Schoenherr, C.J. Genera et species Curculionidum, cum synonymia
hujus familiae. Species novae aut hactenus minus cognitae, descriptionibus a Dom.
Leonardo Gyllenhal, C.H. Boheman, et entomologis aliis illustratae. Volume 2, Part 1, 326
pp., Part 2, 329-669 pp. Paris.
Hardy, A.R. and F.G. Andrews. 1976. A final report to the office of endangered species in
contract 14-16-0008-966. California Department of Food and Agriculture, Insect
Taxonomy Laboratory. 53 pp. Sacramento, California.
Hatch, M.H. 1971. The beetles of the Pacific Northwest. Part V: Rhipiceroidea, Sternoxi,
Phytophaga, Rhynchophora, and Lamellicornia. University of Washington Publications in
Biology 16. xiv + 662 pp.
Hecht, M.K. and J.L. Edwards. 1976. The determination of parallel or monophyletic
relationships: The proteid salamanders - a test case. American Naturalist 100:653-677.
Hecht, M.K. and J.L. Edwards. 1977. The methodology of phylogenetic inference above the
species level, pp. 3-51, In: Hecht, M.K., P.C. Goody and B.M. Hecht, Eds., Major patterns
in vertebrate evolution. Volume 14. NATO Advanced Study Institute Series A. Plenum
Press, New York. 908 pp.
Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana. 263 pp.
Henshaw, S. 1881-1882. Index to the Coleoptera described by J.L. LeConte, M.D.
Transactions of the American Entomological Society 9:197-272.
Henshaw, S. 1885. List of the Coleoptera of America, north of Mexico. American
Entomological Society. 161 pp. Philadelphia, Pennsylvania.
Herbst, J.F.W. 1795. Natursystem aller bekannten in= und auslandischen Insekten, als eine
Forsetzung der von Biiffonschen Naturgeschichte. Der Kafer. VI. Theil, xxiv + 520 pp.
Berlin.
Hicks, S.D. 1947. Additonal notes on Coleoptera taken in Essex County and southern Ontario.
The Canadian Entomologist 79:1 17-1 19.
Hicks, S.D. 1949. A note on the occurrence of Cleonus piger, a European weevil near Ottawa,
Ontario. The Coleopterists' Bulletin 3:7.
Horn, G.H. 1894. The Coleoptera of Baja California. Proceedings of the California Academy
of Sciences, Series 2 4(l):302-449.
Hughes, O.L., C.R. Harrington, J.A. Janssens, J.V. Matthews, Jr., R.E. Morlan, N.W. Rutter
and C.E. Schweger. 1981. Upper Pleistocene stratigraphy, paleoecology, and archaeology of
the northern Yukon interior, eastern Beringia 1. Bonnet Plume Basin. Arctic 34:329-365.
Hull, D.L. 1974. Philosophy of biological science. Prentice-Hall Inc., Englewood Cliffs, i-xii +
1-148 pp.
Kearney, T.H. and R.H. Peebles. 1960. Arizona Flora. University of California Press,
Berkeley, i-viii + 1085 pp.
New World Cleonini
601
Kingsolver, J.M. 1961. Evolutionary studies in the weevil genus Lixus (Coleoptera,
Curculionidae). Unpublished Ph.D. Thesis. University of Illinois, Urbana. 108 pp.
Kingsolver, J.M. 1972. A catalogue of the genus Lixus F. for America north of Mexico
(Coleoptera: Curculionidae). The Coleopterists‘ Bulletin 26:151-154.
Kirby, W. 1837. Part the fourth and last. The insects. In: Richardson, J. Fauna
Boreali-Americana; or the zoology of the northern parts of British America: containing
descriptions of the objects of natural history collected on the late Northern Land
Expeditions, under command of Captain Sir John Franklin, R.N. xxxix + 325 pp.
Kirkland, R.L. and R.D. Goeden. 1977. Descriptions of the immature stages of imported
puncturevine weevils, Microlarinus lareynii and M. lypriformis. Annals of the
Entomological Society of America 70:583-587.
Kirkland, R.L. and R.D. Goeden. 1978a. Biology of Microlarinus lareynii (Col.:
Curculionidae) on puncturevine in Southern California. Annals of the Entomological
Society of America 71:13-18.
Kirkland, R.L. and R.D. Goeden. 1978b. Biology of Microlarinus lypriformis (Col.:
Curculionidae) on puncturevine in Southern California. Annals of the Entomological
Society of America 7 1 :65-69.
Kissinger, D.G. 1964. Curculionidae of America, north of Mexico. A key to the genera.
Taxonomic Publications, v + 143 pp. South Lancaster, Massachusetts.
Kissinger, D.G. 1970. Curculionidae tribe Ophyrastini of North America (Coleoptera).
Taxonomic Publications, vi + 238 pp. South Lancaster, Massachusetts.
Knoch, A.W. 1781. Beitrage zur Insectengeschichte. Leipzig, Schwickert. Stuck 1. 10, 98u 3p.
6 col. Taf.
Krombein, K.V. 1979. Superfamily Sphecoidea. pp. 1573-1740. In: Krombein, K.V. et al.
Catalog of Hymenoptera in America North of Mexico. Volume 2: Apocrita (Aculeata).
Smithsonian Institution Press, xvi + 2209 pp. Washington D.C.
Krombein, K.V. et al.. 1979. Catalog of Hymenoptera in America North of Mexico. Volumes
1, 2, 3. Smithsonian Institution Press. Washington D.C.
Kumar, R., R.J. Lavigne, J.E. Lloyd and R.E. Pfadt. 1976. Insects of the Central Plains
Experiment Range, Pawnee National Grassland. Agricultural Experiment Station,
University of Wyoming, Science Monograph No. 32. iv + 74 pp.
Kuschel, G. 1949. Los “Curculionidae” del extremo norte de Chile (Coleoptera, Curcul. ap.6°).
Acta Zoologica Lilloana 8:5-54.
Kuschel, G. 1950. I. Nuevos Curculionidae de Bolivia y Peru. II. Notas a algunas especies de
Brethes. Revista del Museo de La Plata 6(Zool. 38):69— 1 16.
Lacordaire, J.T. 1863. Histoire naturelle des insectes. Genera des Coleopteres ou expose
methodique et critique de tous les genres proposes jusqu‘ici dans cet ordre d‘insectes.
Volume 6. pp. 1-608, 615-637. Paris.
La Ferla, A. 1939. Contributo alia conosczenza del Cleono del Carciofo ( Cleonus piger Scop.).
Bollettino del Laboratorio di Entomologia della Facolta agraria in Portici 3:3-33.
Latreille, P.A. 1810. Considerations genenerale sur 1‘ordre natural des animaux composant les
classes des crustaces, des arachnides, et des insectes; avec un tableau methodique de leurs
genres, disposes en families. Schoell, Paris. 444 pp.
LeConte, J.L. 1850. General remarks upon the Coleoptera of Lake Superior. Article IV, pp.
201-242 + 8 pi. In: Agassiz, J.L.R. and J.E. Cabot. Lake Superior: Its physical character,
vegetation and animals. 461 pp. Boston, Massachusetts.
Quaest. Ent., 1987, 23 (4)
602
Anderson
LeConte, J.L. 1858. Descriptions of new species of Coleoptera, chiefly collected by the United
States and Mexican Boundary Commission, under Major W.H. Emory, U.S.A. Proceedings
of the Academy of Natural Sciences of Philadelphia. 10:59-89.
LeConte, J.L. 1859a. The Coleoptera of Kansas and eastern New Mexico. Smithsonian
Contributions to Knowledge 2(6) :i-vi + 58 pp.
LeConte, J.L. 1859b. The complete writings of Thomas Say on the entomology of North
America. Volume 1. xxiv + 412 pp.
LeConte, J.L. 1876a. Tribe IV. Cleonini. pp. 144-160. In: LeConte, J.L. and G.H. Horn. The
Rhynchophora of America, north of Mexico. Proceedings of the American Philosophical
Society 15(96):i-xvi + 1-455.
LeConte, J.L. 1876b. Appendix, pp. 412-442. In: LeConte, J.L. and G.H. Horn. The
Rhynchophora of America, north of Mexico. Proceedings of the American Philosophical
Society 15(96):i-xvi + 1-455.
LeConte, J.L. 1878. The Coleoptera of the Alpine Regions of the Rocky Mountains. Bulletin of
the United States Geological and Geographical Survey of the Territories 4(2)
1878:447-480.
LeConte, J.L. and G.H. Horn. 1883. Classification of the Coleoptera of North America.
Smithsonian Institution. 567 pp. Washington D.C.
Leng, C.W. 1919. The Rhynchophora (except Ipidae). pp. 19E-21E In: Report of the Canadian
Arctic Expedition. Volume 3: Insects, part E: Coleoptera. pp. 3E-27E.
Leng, C.W. 1920. Catalog of the Coleoptera of America, north of Mexico, x + 470 pp. The
Cosmos Press. Cambridge, Massachusetts.
Leng, C.W. and A.J. Mutchler. 1933. Second and third supplements 1925 to 1932 (inclusive)
to Catalogue of the Coleoptera of America, North of Mexico. 1 12 pp. The Cambridge Press.
Cambridge, Massachusetts.
Leonard, M.D. 1926. A list of the insects of New York with a list of the spiders and certain
other allied groups. Cornell University, Agricultural Experiment Station. Memoir # 101.
1121 pp.
Linnaeus, C. 1758. Systema naturae, per regna tria naturae, secumdum classes, ordines,
genera, species cum characteribus, differentiis, synonymis, locis, ed. 10, reformata, Volume
1, 823 pp. Holmiae.
Linnaeus, C. 1767. Ibid., Vol. 1, pars 2, editio duodecimo reformata , pp. 533-1327. Holmiae.
Livingston, B.E. and F. Shreve. 1921. The distribution of vegetation in the United States, as
related to climatic conditions, xiv + 590 pp. Carnegie Institute of Washington, Washington
D.C..
Lohse, G.A. 1983. Unterfamilie: Cleoninae. pp. 7-29. In\ Freude, H., K.W. Harde, and G.A.
Lohse. Die Kafer Mitteleuropas. Band 1 1. Goecke and Evers, Krefeld. 342 pp.
Lukjanovitsh, T. 1926. Bemerkungen iiber Systematik und geographische Verbreitung der
Curculioniden (Coleoptera). Entomologicheskoe Obozrenie 20: 140-143.
Maddison, W.P., M.J. Donoghue, and D.R. Maddison. 1984. Out-group analysis and
parsimony. Systematic Zoology 33:83-103.
Maddox, D.M. 1976. History of weevils on puncturevine in and near the United States. Weed
Science 24:4 1 4-4 1 9.
Mann, J.S. and R.A. Crowson. 1983. On the occurrence of mid-gut caeca, and organs of
symbiont transmission, in leaf beetles. The Coleopterists Bulletin 37:1-15.
Mannerheim, G.C.G. 1843. Beitrag zur Kafer-Fauna der Aleutischen Inseln der Insel Sitkha
New World Cleonini
603
und Neu-Californiens. Bulletin de la Societe Imperiale des Naturalistes de Moscou
1 6(2) : 1 7 5—3 1 4.
Marcovitch, S. 1923. A root weevil ( Stephanocleonus plumbeus Lee.) reared from strawberry.
The Canadian Entomologist 55:218.
Marshall, G.A.K. 1932. Notes on the Hylobiinae (Col., Cure.). Annals and Magazine of
Natural History, Series 10 9:341-355.
Matthews, J.V. Jr. 1974. Quaternary environments at Cape Deceit (Seward Penninsula,
Alaska): evolution of a tundra ecosystem. Geological Society of America, Bulletin
85:1353-1384.
Matthews, J.V. Jr. 1975. Insects and plant macrofossils from two Quaternary exposures in the
Old Crow-Porcupine region, Yukon Territory, Canada. Arctic and Alpine Research
7:249-259.
Matthews, J.V. Jr. 1977. Tertiary Coleoptera fossils from the North American Arctic. The
Coleopterists Bulletin 31:297-308.
Mayr, E. 1963. Animal species and evolution. Belknap Press, Harvard University. Cambridge,
Massachusetts. 797 pp.
Melsheimer, F.E. 1853. Catalogue of the described Coleoptera of the United States, xvi + 174
pp.
Morgan, A.V., A. Morgan, A.C. Ashworth and J.V. Matthews, Jr. 1984. Late Wisconsin fossil
beetles in North America, pp. 354-363. In: Porter, S.C., Ed. Late-Quaternary
Environments of the United States. Volume 1. The Late Pleistocene. University of
Minnesota Press, Minneapolis.
Morimoto, K. 1962. Comparative morphology and phylogeny of the superfamily
Curculionoidea of Japan. Journal of the Faculty of Agriculture, Kyushu University
11:331-373.
Morlan, R.E. and J.V. Matthews, Jr. 1983. Taphonomy and paleoecology of fossil insect
assemblages from Old Crow River (CRH- 15) Northern Yukon Territory, Canada.
Geographic physique et Quaternaire 37:147-157.
Motschulsky, T.V. von. 1845. Observations sur la musee entomologique de 1‘Universite
Imperiale de Moscou. Bulletin de la Societe Imperiale des Naturalistes de Moscou
18:332-388.
Motschulsky, T.V. von. 1860. Coleopteres rapportes par M. Severtsef des Steppes meridionales
des Kirghises. Bulletin de TAcademie Imperiale des Sciences de St.-Petersbourg
2:513-544.
Mott, R.J., T.W. Anderson and J.V. Matthews, Jr. 1981. Late-glacial paleoenvironments of
sites bordering the Champlain Sea based on pollen and macrofossil evidence, pp. 129-171.
In: W.C. Mahaney, Ed. Quaternary Paleoclimate. Geoabstracts, Norwich, England.
Muller, O.F. 1776. Zoologiae Danicae prodromus, seu animalium Daniae et Norwegiae
indigenarum characteres, nomina, et synonyma imprimis popularium. xxxii + 282 pp.
Hafniae.
Munz, P.A. and D.D. Keck. 1968. A California Flora. University of California Press. Berkeley,
California. 1681 pp.
O’Brien, C.W. and G.J. Wibmer. 1978. Numbers of genera and species of Curculionidae
(Coleoptera). Entomological News 89:89-92.
O’Brien, C.W. and G.J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae
sensu lato) of North America, Central America, and the West Indies (Coleoptera:
Quaest. Ent., 1987, 23 (4)
604
Anderson
Curculionoidea). Memoirs of the American Entomological Society 34:x + 382 pp.
O’Brien, C.W. and G.J. Wibmer. 1984. Annotated checklist of the weevils (Curculionidae
sensu lato) of North America, Central America, and the West Indies. Supplement 1. The
Southwestern Entomologist 9:286-307.
Olivier, A.G. 1807. Entomologie, ou histoire naturelle des insectes, avec leurs caracteres
generiques et specifiques, leur description, leur synonymie, et leur figure enluminee.
Coleopteres. Volume 5. 612 pp. Paris.
Pallas, P.S. 1781. leones insectorum praesertim Rossiae Sibiriaeque peculiarium quae collegit
et descriptionibus illustravit. Walther, Erlangae. 56 pp.
Patterson, C. 1981. Methods of paleobiogeography. pp. 446-489. In : Nelson, G. and D.E.
Rosen, Eds. Vicariance biogeography. A Critique. Columbia University Press, New York,
i-xviii + 1-593 pp.
Peschken, D.P. 1984. Host range of Lema cyanella (Coleoptera: Chrysomelidae), a candidate
for biocontrol of Canada Thistle, and of four stenophagous, foreign thistle insects in North
America. The Canadian Entomologist 116:1377-1384.
Petri, K. 1905. Bestimmungs-Tabelle der mir bekannt gewordenen Arten der Gattung Lixus
Fab. aus Europa und der angrenzenden Gebieten. Wiener Entomologische Zeitung
24:33-48, 101-117, 155-167.
Petri, K. 1912. Ein neuer Lixus aus Turkestan und Bemerkungen zu meiner
Bestimmungstabelle des Genus Lixus Fabr.. Annales Historico-naturales Musei Nationalis
Hungarici, Budapest 10:277-280.
Pianka, E.R. 1975. Niche relations of desert lizards, pp. 292-314. In: Cody, M.L. and J.M.
Diamond, Eds. Ecology and evolution of communities. Belknap Press of Harvard University,
Cambridge, i-xiv + 545 pp.
Pierce, W.D. 1907. On the biologies of the Rhynchophora of North America. Studies from the
Zoological Laboratory, The University of Nebraska 78:249-319.
Platnick, N.I. and G. Nelson. 1978. A method of analysis for historical biogeography.
Systematic Zoology 27:1-16.
Randall, J. 1838. Descriptions of new species of coleopterous insects inhabiting the state of
Massachusetts. Boston Journal of Natural History 2:34-52.
Raven, P.H. 1963. Amphitropical relationships in the floras of North and South America.
Quarterly Review of Biology 38:151-177.
Raven, P.H. and D.I. Axelrod. 1978. Origin and relationships of the California flora.
University of California Publications in Botany 72:i-viii + 1-134.
Reitter, E. 1916. Fauna Germanica. Die Kafer des Deutschen Reiches. Nach der analytischen
Methode bearbeitet. Band V. K.G. Lutz‘ Verlag. Stuttgart. 343 pp.
Rzedowski, J. 1978. Vegetacion de Mexico. Editorial Limusa. 432 pp. Mexico City.
Sanders, H.O. 1960. The female genitalia and spermathecae of some of the Rhynchophora. The
Great Basin Naturalist 20:1-22.
Say, T. 1831. Descriptions of new species of Curculionites of North America, with observations
of some of the species already known. 30 pp. New Harmony, Indiana.
Scheinert, W. 1933. Symbiose und Embryonalentwicklung bei Riisselkafern. Zeitschrift fur
Morphologie und Okologie der Tiere 27:76-128.
Schenkling, S. and G.A.K. Marshall. 1931. Coleopterorum Catalogus. Pars 114.
Curculionidae: Cylindrorrhininae. pp. 1-23.
Schoenherr, C.J. 1823. Tabula synoptica familiae curculionidum. Isis Oken, heft X, columns
New World Cleonini
605
1132-1146.
Schoenherr, C.J. 1826. Curculionidum disposito methodica cum generum characterihus,
descriptionibus atque observationibus variis, seu prodromus ad synonymiae insectorum,
partena 4, x + 338 pp. Lipsiae.
Schoenherr, C.J. 1834. Genera et species Curculionidum, cum synonymia hujus familiae.
Species novae aut hactenus minus cognitae, descriptionibus a Dom. Leonardo Gyllenhal,
C.H. Boheman, et entomologis aliis illustratae. Volume 2, Part 1. 1-326 pp. Part 2.
329-669 pp. Paris.
Scopoli, J.A. 1763. Entomologia Carniolica exhibens insecta Carnioliae indigena distributa in
ordines, genera, species, varietates. Methodo Linnaeana. xxxiv + 420 pp. Vindobonae.
Scudder, S.H. 1893. Tertiary rhynchophorous Coleoptera of the United States. Monographs of
the United States Geological Survey 21:1-206.
Shreve, F. 1942. The desert vegetation of North America. Botanical Review 195-246.
Silfverberg, H. (Ed.) 1979. Enumeratio Coleopterorum Fennoscandiae et Daniae. vi + 79 pp.
Helsinki.
Sleeper, E.L. 1969. Two new Cleoninae from the Rita Blanca Lake deposits (Coleoptera:
Curculionidae). Geological Society of America, Memoirs 1 13:131-133.
Smith, P.W. 1957. An analysis of post-Wisconsin biogeography of the Prairie Peninsula region
based on distributional phenomena among terrestrial vertebrate populations. Ecology
38:205-218.
Solari, F. 1941. Revisione dei Neoplinthus Italiani ed alcune note di sistematica generate dei
Curculionidi (Coleoptera). Memoire Societa Entomologica Italiana 20:43-90.
Spaulding, W.G., E.B. Leopold, and T.R. Van Devender. 1985. Late Wisconsin paleoecology of
the American Southwest, pp 259-293. In: Porter, S.C., Ed. Late-Quaternary Environments
of the United States. Volume 1. The Late Pleistocene. University of Minnesota Press,
Minneapolis.
Sprague, P.S. and E.P. Austin. 1875. On the species of Coleoptera described by Mr. J.W.
Randall. Proceedings of the Boston Society of Natural History 17:373-385.
Stebbins, G.L. and J. Major. 1965. Endemism and speciation in the California flora.
Ecological Monographs 35:1-35.
Tanner, V.M. 1934. Coleoptera of Zion National Park No. II. Annals of the Entomological
Society of America 27:43-49.
Tanner, V.M. 1966. Rhynchophora beetles of the Nevada Test Site. Brigham Young University
Science Bulletin, Biological Series 8(2): 1-35.
Ter-Minasyan, M.E. 1963. Phytogeny of beetles of the subfamily Cleoninae (Coleoptera,
Curculionidae). Entomological Review 42:420-423.
Ter-Minasyan, M.E. 1978. Weevils of the subfamily Cleoninae in the fauna of the USSR.
Tribe Lixini. Zoological Institute, Academy of Sciences of the USSR, vii + 166 pp.
Amerind Publishing Co., New Delhi. (Translated from Russian; originally published in
1967).
Ter-Minasyan, M.E. 1979. Review of the weevil genus Stephanocleonus Motsch. (Coleoptera;
Curculionidae). Insects of Mongolia. Number 6. pp. 184-342.
Ting, P.C. 1936. The mouthparts of the coleopterous group Rhynchophora. Microentomology
1:93-114.
Van Dyke, E.C. 1953. New Coleoptera from western North America (Carabidae, Throscidae,
Curculionidae). The Pan-Pacific Entomologist 29:98-101.
Quaest. Ent., 1987, 23 (4)
606
Anderson
Villers, C.J. 1789. Caroli Linnaei entomologia. Band 1. Lugduni, Piestre et Delamolliere. 24u
765p. 3 Taf.
Vuilleumier, F. and D. Simberloff. 1980. Ecology versus history as determinants of patchy and
insular distributions in high Andean birds. Evolutionary Biology 12:235-379.
Watrous, L.E. and Q.D. Wheeler. 1981. The out-group comparison method of character
analysis. Systematic Zoology 30: 1-1 1 .
Webb, S.D. 1977. A history of savanna vertebrates in the New World. Part I: North America.
Annual Review of Ecology and Systematics 8:355-380.
Wells, P.V. and J.H. Hunziker. 1976. Origin of the creosote bush ( Larrea ) deserts of
southwestern North America. Annals of the Missouri Botanical Garden 63:843-861.
Wheeler, A.G., Jr. and D.R. Whitehead. 1985. Larinus planus (F.) in North America
(Coleoptera: Curculionidae: Cleoninae) and comments on biological control of Canada
thistle. Proceedings of the Entomological Society of Washington 87:751-758.
Whitehead, D.R. 1972. Classification, phylogeny, and zoogeography of Schizogenius Putzeys
(Coleoptera: Carabidae: Scaratini). Quaestiones Entomologicae 8:131-348.
Wibmer, G.J. and C.W. O’Brien. 1986. Annotated checklist of the weevils (Curculionidae
sensu lato) of South America (Coleoptera: Curculionidae). Memoirs of the American
Entomological Institute 39:i-xvi + 1-563 pp.
Wickham, H.F. 1889. Collecting notes. Entomologica Americana 5:77-78.
Wickham, H.F. 1896. A list of some Coleoptera from the southern portions of New Mexico and
Arizona. Bulletin from the Laboratories of Natural History, State University of Iowa
3:153-171.
Wickham, H.F. 1902. A catalog of the Coleoptera of Colorado. Bulletin of the State of Illinois
Laboratory of Natural History 5:217-310.
Wickham, H.F. 1909. A list of the Coleoptera of Iowa. Bulletin from the Laboratories of
Natural History of the State University of Iowa 6:1-40.
Wickham, H.F. 1911. Fossil Coleoptera from Florissant, with descriptions of several new
species. Bulletin of the American Museum of Natural History 30:53-69.
Wickham, H.F. 1912. On some fossil Rhynchophorous Coleoptera from Florissant, Colorado.
Bulletin of the American Museum of Natural History 31:41 —55 + 4 pi.
Wilcox, J., D.C. Mote, and L. Childs. 1934. The root weevils injurious to Strawberries in
Oregon. Oregon State Agricultural Experiment Station, Station Bulletin 330. 109 pp.
Wiley, E.O. 1981. Phylogenetics. The Theory and Practice of Phylogenetic Systematics.
Wiley-Interscience, John Wiley and Sons, New York, i-xvi + 1-439 pp.
Yothers, W. 1916. Bud weevils and other bud-eating insects of Washington. State College of
Washington Agricultural Experiment Station Bulletin 124. 43 pp. + 6 plates.
Zoubkoff, B. 1829. Notice sur un nouveau genre et quelques nouvelles especes de Coleopteres.
Bulletin de la Societe Imperiale des Naturalistes de Moscou 1:147-168, tab. 2.
Zoubkoff, B. 1833. Nouveaux Coleopteres recueillis en Turcmenie. Ibid., 6:310-340.
Zwolfer, H. and P. Harris. 1984. Biology and host specificity of Rhinocyllus conicus (Froel.)
(Col. Curculionidae), a successful agent for biocontrol of the thistle, Carduus nutans L.
Zeitschrift fur Angewandte Entomologie 97:36-62.
Zwolfer, H. and M. Preiss. 1983. Host selection and oviposition behaviour in West-European
ecotypes of Rhinocyllus conicus Froel. (Col. Curculionidae). Ibid., 95:1 13-122.
New World Cleonini
607
Figures 1-4. Lateral view of head of various Cleoninae. 1, Microlarinus species (Scale bar = 0.8 mm); 2, Rhinocyllus
conicus (Scale bar = 1.6 mm); 3, Larinus planus (Scale bar = 1.6 mm); 4, Lixus scrobicollis (Scale bar = 1.6 mm).
Figures 5-6. Apex of tibia of various Cleoninae (Scale bar = 0.5 mm). 5, Cleonidius infrequens; 6, Apleurus lutulentus.
Figures 7-8. Ventral view of prosternum of various Cleoninae (Scale bar = 1.6 mm). 7, Apleurus species; 8, Cleonidius
species.
Quaest. Ent., 1987, 23 (4)
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Figures 9- 18. Cleonis pigra. 9, lateral view of head (Scale bar =1.1 mm); 10, dorsal view of head (Scale bar =1.1 mm);
1 1, lateral view of internal sac of male (Scale bar = 0.7 mm); 12, dorsal view of internal sac of male (Scale bar = 0.7
mm); 13, lateral view of aedeagus of male (Scale bar = 1.3 mm); 14, ventral view of apical sclerite complex of internal sac
of male (Scale bar = 0,6 mm); 15, lateral view of gonocoxite II and stylus of female (Scale bar = 1.3 mm); 16,
spermatheca of female (Scale bar = 0.7 mm); 17, ventral view of sternum VIII of female (Scale bar = 1.3 mm); 18,
ventral view of metatarsus (Scale bar = 1.3 mm).
New World Cleonini
609
Figures 19-24. Dorsal habitus of: 19, Cleonis pigra (Scale bar = 5.0 mm); 20, Stephanocleonus confusus (Scale bar
= 5.0 mm); 21, S. immaculatus (Scale bar = 5.0 mm); 22, S. plumbeus (Scale bar = 5.0 mm); 23, S. stenothorax
(Scale bar = 5.0 mm); 24, Apleurus (Gibbostethus) hystrix (Scale bar = 2.5 mm).
Quaest. Ent., 1987, 23 (4)
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Anderson
Figures 25-30. Head of Stephanocleonus species (Scale bar = 2.4 mm) - a, lateral view; b, dorsal view. 25, S. confusus\
26, S. cristaticollis', 27, S. immaculatus\ 28, S. parshus ; 29, S. plumbeus ; 30, S', stenothorax.
-Q
New World Cleonini
611
Figures 31-36. Male genitalia of Stephanocleonus species (Scale bar = 1.3 mm) - a and b, lateral and ventral views of
aedeagus, respectively; c, ventral view of sternum VII. 31, S', confusus ; 32, S. cristaticollis-, 33, S. immaculatus ; 34, S.
parshus-, 35, S. plumbeus-, 36, S. stenothorax. Figures 37-42. Ventral view of sternum VIII of female Stephanocleonus
species (Scale bar = 1.3 mm). 37, S. confusus-, 38, S. cristaticollis-, 39, S. immaculatus-, 40, S. parshus-, 41, S. plumbeus-,
42, S. stenothorax.
Quaest. Ent., 1987, 23 (4)
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Figures 43-48. Internal sac of male Stephanocleonus species (Scale bar = 0.7 mm) - a and b, lateral and dorsal view,
respectively. 43, S. confusus ; 44, S. cristaticollis\ 45. S. immaculatus\ 46, S. parshus\ 47, S. plumbeus. 48, S.
stenothorax.
New World Cleonini
613
Figure 49. Hubbs-Hubbs diagram illustrating variation among specimens of Stephanocleonus species: length of elytra
(LEI) - a, males; b, females. The range of each species is indicated by a horizontal line and the mean by a long vertical
line. A solid rectangle represents two standard errors on either side of the mean and a hollow rectangle represents 1.5
standard deviations on either side of the mean. Only mean and range are shown for samples with less than five specimens.
Quaest. Ent., 1987, 23 (4)
614
Anderson
50 51
Figures 50-51. Hubbs-Hubbs diagrams illustrating variation among specimens of Stephanocleonus species - a, males; b,
females. See caption for Fig. 49 for explanation. 50, form of elytra (WE1M/LE1); 51, relative length of pronotum
compared to elytra (LP/LE1).
New World Cleonini
615
Figure 52. Hubbs-Hubbs diagram illustrating variation among specimens of Stephanocleonus species: form of rostrum
(WRA/WF) - a, males; b, females. See caption for Fig. 49 for explanation.
Quaest. Ent., 1987, 23 (4)
616
Anderson
S.
S.
S.
S.
S.
S.
confusus
cr i st at icol 1
immacul at us
par shu s
p 1 umbeus
stenothorax
53
Figure 53. Hubbs-Hubbs diagram illustrating variation among specimens of Stephanocleonus species: relative width of
frons compared to rostrum (WRA/WF) - a, males; b, females. See caption for Fig. 49 for explanation.
New World Cleonini
617
Figures 54-67. 54, lateral view of gonocoxite II and stylus of female Stephanocleonus (Scale bar = 0.7 mm). 55,
spermatheca of female Stephanocleonus (Scale bar = 0.7 mm). 56, genitalia of Chromoderus fasciatus; a, lateral view of
aedeagus of male (Scale bar =1.3 mm); b, lateral view of internal sac of male (Scale bar = 0.7 mm); c, dorsal view of
internal sac of male (Scale bar = 0.7 mm); d, ventral view of apical sclerite complex of internal sac of male (Scale bar =
0.6 mm); e, ventral view of sternum VIII of female (Scale bar = 1.3 mm). 57, genitalia of Cnemodontus limpidus\ a,
lateral view of aedeagus of male (Scale bar = 1.3 mm); b, lateral view of internal sac of male (Scale bar = 0.7 mm); c,
dorsal view of internal sac of male (Scale bar = 0.7 mm); d, ventral view of apical sclerite complex of internal sac of male
(Scale bar = 0.6 mm); e, ventral view of sternum VIII of female (Scale bar = 1.3 mm). 58, lateral view of mesosternum
of Apleurus ( Gibbostethus ) hystrix (Scale bar = 1.3 mm). 59, lateral view of mesosternum of Apleurus (Apleurus)
lutulentus (Scale bar = 1.3 mm). 60, spermatheca of female Apleurus (Scale bar = 0.7 mm). 61, ventral view of ligula of
Apleurus (Gibbostethus) hystrix (Scale bar = 0.2 mm). 62, ventral view of ligula of Apleurus ( Apleurus ) lutulentus
(Scale bar = 0.4 mm). 63, ventral view of ligula of Apleurus ( Apleurus ) saginatus (Scale bar = 0.4 mm). 64, ventral view
of maxilla of Apleurus ( Apleurus ) saginatus (Scale bar = 0.4 mm). 65, ventral view of maxilla of Apleurus
( Gibbostethus ) hystrix (Scale bar = 0.4 mm). 66, tarsal claws of Apleurus angularis (Scale bar = 0.7 mm). 67, tarsal
claws of Apleurus jacobinus (Scale bar = 0.7 mm).
Quaest. Ent., 1987, 23 (4)
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Figures 68-74. Dorsal habitus of species of Apleurus. 68, A. aztecus (Scale bar = 2.0 mm); 69, A. lutulentus (Scale bar
= 4.0 mm); 70, A. porosus (Scale bar = 4.0 mm); 71, A. jacobinus (Scale bar = 4.0 mm); 72, A. angularis (3 mi. N
Baker, Nevada) (Scale bar = 4.0 mm); 73, A. angularis (Colorado Springs, Colorado) (Scale bar = 4.0 mm); 74, A.
angularis (Baboquivari Mountains, Arizona) (Scale bar = 3.0 mm).
New World Cleonini
619
Figures 75-80. Dorsal habitus of species of Apleurus (Scale bar = 5.0 mm). 75, A. albovestitus (18 mi. SW Mendota,
California); 76, A. albovestitus (16 mi. S. Vidal, California); 77, A. albovestitus (Virgin, Utah); 78, A. albovestitus
(Phoenix, Arizona); 79, A. albovestitus (Yuma, Arizona); 80, A. saginatus.
Quaest. Ent., 1987,23 (4)
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Figures 81-88. Head of Apleurus species (Scale bar = 2.4 mm) - a and b, lateral and dorsal views, respectively. 81, A.
hystrix ; 82, A. azlecus ; 83, A. lutulentus\ 84, A. porosus; 85, A. angularis\ 86, A. jacobinus\ 87, A. saginatus\ 88, A.
albovestitus.
Figures 89-96. Female genitalia of Apleurus species. 89, A. hystrix (Scale bar = 1.0 mm); a, ventral view of sternum
VIII; b, lateral view of gonocoxite II and stylus. 90, A. aztecus\ a, ventral view of sternum VIII (Scale bar = 2.0 mm); b,
lateral view of gonocoxite II and stylus (Scale bar = 1.0 mm). 91, A. lutulentus\ a, ventral view of sternum VIII (Scale
bar = 2.0 mm); b, lateral view of gonocoxite II and stylus (Scale bar = 1.0 mm). 92, A. porosus; a, ventral view of
sternum VIII (Scale bar = 2.0 mm); b, lateral view of gonocoxite II and stylus (Scale bar = 1.0 mm). 93, A. angularis\ a,
ventral view of sternum VIII (Scale bar = 2.0 mm); b, lateral view of gonocoxite II and stylus (Scale bar = 1.0 mm). 94,
A. jacobinus\ a, ventral view of sternum VIII (Scale bar = 2.0 mm); b, lateral view of gonocoxite II and stylus (Scale bar
= 1.0 mm). 95, A. albovestitus ; a, ventral view of sternum VIII (Scale bar = 2.0 mm); b, lateral view of gonocoxite II
and stylus (Scale bar = 1.0 mm). 96, A. saginatus\ a, ventral view of sternum VIII (Scale bar = 2.0 mm); b, lateral view
of gonocoxite II and stylus (Scale bar = 1.0 mm).
Quaest. Ent., 1987, 23 (4)
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Figures 97-99. Male genitalia of Apleurus species. 97, A. hystrix; a, lateral view of internal sac (Scale bar = 0.4 mm); b,
dorsal view of internal sac (Scale bar = 0.4 mm); c, ventral view of apical sclerite complex of internal sac (Scale bar =
0.3 mm); d, lateral view of aedeagus (Scale bar = 1.3 mm); e, ventral view of aedeagus (Scale bar = 1.3 mm); f, ventral
view of stemite of sternum VII (Scale bar = 1.3 mm). 98, A. aztecus; a, lateral view of internal sac (Scale bar = 0.7
mm); b, dorsal view of internal sac (Scale bar = 0.7 mm); c, lateral view of aedeagus (Scale bar = 1.3 mm); d, ventral
view of apical sclerite complex of internal sac (Scale bar = 0.6 mm). 99, A. lutulentus (Scale bars as in Fig. 98); a, lateral
view of internal sac; b, dorsal view of internal sac; c, lateral view of aedeagus; d, ventral view of apical sclerite complex of
internal sac.
New World Cleonini
623
Figures 100-102. Male genitalia of Apleurus species - a, lateral view of internal sac (Scale bar = 0.7 mm); b, dorsal view
of internal sac (Scale bar = 0.7 mm); c, lateral view of aedeagus (Scale bar = 1.3 mm); d, ventral view of apical sclerite
complex of internal sac (Scale bar = 0.6 mm); e, ventral view of sternite of sternum VII. 100, A. angularis ; 101, A.
jacobinus ; 102, A. porosus.
Quaest. Ent., 1987, 23 (4)
624
Anderson
Figures 103-104. Male genitalia of Apleurus species - a, lateral view of internal sac (Scale bar = 0.7 mm); b, dorsal view
of internal sac (Scale bar = 0.7 mm); c, lateral view of aedeagus (Scale bar = 1.3 mm); d, ventral view of apical sclerite
complex of internal sac (Scale bar = 0.6 mm); e, ventral view of sternite of sternum VII (Scale bar = 1.3 mm). 103, A.
albovestitus\ 104, A. saginatus.
New World Cleonini
105
Figure 105. Hubbs-Hubbs diagram illustrating variation among specimens of Apleurus species: length of elytra (LEI)
males; b, females. See caption for Fig. 49 for explanation.
Quaest. Ent., 1987, 23 (4)
626
Anderson
106
8
13
33
7
95
28
99
40
8
6
37
8
'• 9 1
• 3 0
= 103
: 4 1
Figure 106. Hubbs-Hubbs diagram illustrating variation among specimens of Apleurus species: form of rostrum
(WRA/LR) - a, males; b, females. See caption for Fig. 49 for explanation.
New World Cleonini
627
Figures 107-108. Hubbs-Hubbs diagrams illustrating variation among specimens of Apleurus species - a, males; b,
females. See caption for Fig. 49 for explanation. 107, form of elytra (WE1M/LE1); 108, width of frons compared to width
at apex of rostrum (WRA/WF).
i
Quaest. Ent., 1987, 23 (4)
628
Anderson
0.80 0.90 1.00 1.10 1-20
109 110
Figures 109-110: Hubbs-Hubbs diagrams illustrating variation among specimens of Apleurus species - a, males; b,
females. See caption for Fig. 49 for explanation. 109, form of pronotum (WPT/WPB); 1 10, length of pronotum compared
to length elytra (LP/LE1).
New World Cleonini
629
Figures 1 1 1-1 17. Ill, Male genitalia of Cylindropterus luxeri (Scale bar = 0.7 mm); a, lateral view of internal sac; b,
dorsal view of internal sac. 1 12, Male genitalia of Lixus (Lixoglyptus) spartii (Scale bar = 0.7 mm); a, lateral view of
internal sac; b, dorsal view of internal sac. 1 13, Ventral view of sternum VIII of female Cylindropterus luxeri (Scale bar
= 1.3 mm). 1 14, Ventral view of sternum VIII of female Lixus ( Lixoglyptus ) spartii (Scale bar = 1.3 mm). 1 15, Internal
sac of male Cleonidius Lobes and pockets of internal sac labelled as noted (Scale bar = 0.7 mm); a, lateral view; b, dorsal
view. 116, Spermatheca of female Cleonidius (Scale bar = 0.3 mm). 117, Lateral view of gonocoxite II and stylus of
Cleonidius (Scale bar = 0.7 mm).
Quaest. Ent., 1987, 23 (4)
630
Anderson
Figures 1 18-123. Dorsal habitus of: 1 18, Cleonidius erysimi (Scale bar = 4.0 mm); 1 19, C. eustictorrhinus (Scale bar =
4.0 mm); 120, C. pleuralis (Scale bar = 4.0 mm); 121, C. subcylindricus (Scale bar = 4.0 mm); 122, C. longinasus
(Scale bar = 4.0 mm); 1 23, C. texanus (Scale bar = 4.0 mm).
New World Cleonini
631
Figures 124-130. Dorsal habitus of: 124, Cleonidius americanus 125, C. frontalis (Scale bar = 4.0 mm); 126, C.
canescens (Scale bar = 4.0 mm); 127, C. infrequens (Scale bar = 4.0 mm); 128, C. puberulus (Scale bar = 4.0 mm);
129, C. collaris (Scale bar = 4.0 mm); 130, C. notolomus. (Scale bar = 4.0 mm).
Quaest. Ent., 1987, 23 (4)
632
Anderson
Figures 131-136. Dorsal habitus of: 131, Cleonidius poricollis (Scale bar = 4.0 mm); 132, C. calandroides (Scale bar =
4.0 mm); 133, C. boucardi (Scale bar = 5.0 mm); 134, C. trivittatus (Scale bar = 5.0 mm); 135, C. placidus (Scale bar
= 5.0 mm); 136, C. quadrilineatus (Scale bar = 4.0 mm).
New World Cleonini
633
Figures 137-142. Head of Cleonidius species (Scale bar = 2.3 mm) - a and b, lateral and dorsal view, respectively. 137,
Cleonidius erysimr, 138, C. eustictorrhinus\ 139, C. pleuralis\ 140, C. subcylindricus ; 141, C. longinasus\ a, Lateral view
of heads of male and female; b. Dorsal view of head of female; 142, Cleonidius texanus.
Quaest. Ent., 1987, 23 (4)
634
Anderson
Figures 143-149. Head and femur of Cleonidius species - (Scale bar = 2.3 mm) a and b, lateral and dorsal view of head,
respectively; c, lateral view of pro-tibia of female. 143, C. americanus\ 144, C. frontalis ; 145, C. canescens\ 146, C.
infrequens', 147, C. puberulus\ 148, C. collaris ; 149, C. notolomus.
New World Cleonini
635
Figures 150-156. Head of Cleonidius species (Scale bar = 2.3 mm) - a and b, lateral and dorsal vies, respectively. 1
poricollis\ 151, C. calartdroides; 152, C. boucardi\ 153, C. trivittatus; 154, C. placidus; 155, C. quadrilineatus\ 1:
vibex. (Scale bar = 2.3 mm).
Quaest. Ent., 1987, 23 (4)
Os Q
636
Anderson
Figures 157-176. Ventral view of sternum VIII of female Cleonidius species (Scale bar = 0.8 mm). 157, C. erysimi’, 158,
C. eustictorrhinus ; 159, C. pleuralis', 160, C. subcylindricus; 161, C. longinasus ; 162, C. texanus\ 163, C. americanus;
164, C. frontalis\ 165, C. canescens\ 166, C. infrequens\ 167, C. puberulus\ 168, C. notolomus ; 169, C. collaris; 170, C.
poricollis’, 171, C. calandroides\ 172, C. boucardi; 173, C. trivittatus\ 174, C. placidus\ 175, C. quadrilineatus’, 176, C.
v/7?ex. (Scale bar = 0.8 mm).
New World Cleonini
637
Figures 177-182. Male genitalia of Cleonidius species - a and b, internal sac, lateral and dorsal view, respectively; c,
lateral view of aedeagus (Scale bars, 0.7 mm, 0.7 mm and 1.3 mm respectively). 177, C. erysimi\ 178, C. eustictorrhinus\
179, C. pleuralis; 180, C. subcylindricus\ 181, C. longinasus ; 182, C. texanus.
Quaest. Ent., 1987,23 (4)
638
Anderson
Figures 183-189. Male genitalia of Cleonidius species - a and b, internal sac, lateral and dorsal view, respectively; c,
lateral view of aedeagus (Scale bars, 0.7 mm, 0.7 mm and 1.3 mm respectively). 183, C. americanus-, 184. C. frontalis-,
185, C. canescens\ 186, C. infrequens\ 187, C. puberulus-, 188, C. notolomus\ 189, C. collaris.
New World Cleonini
639
Figures 190-196. Male genitalia of Cleonidius species - a and b, internal sac, lateral and dorsal views, respectively; c,
lateral view of adeagus (Scale bars, 0.7 mm, 0.7 mm and 1.3 mm, respectively): 190, C. poricollis\ 191, C. calandroides\
192, C. boucardi\ 193, C. trivittatus\ 194, C. placidus ; 195, C. quadrilineatus ; 196, C. vibex.
Quaest. Ent., 1987,23 (4)
640
Anderson
cn^rt^r^oco to *— o oocn to ir>
-r-^^r-r-cO-CNtD^lOyCOCOCO’-OO’-^
CD
Figure 197. Hubbs-Hubbs diagram illustrating variation among specimens of Cleonidius species; length of elytra (LEI) - a, males; b, females. See caption for Fig. 49 for explanation.
New World Cleonini
641
00
CD
0000000000000<J0<J00<J0
Quaest. Ent., 1987, 23 (4)
Figure 198. Hubbs-Hubbs diagram illustrating variation among specimens of Cleonidius species; form of rostrum (WRA/LR) - a, males; b, females. See caption for Fig. 49 for explanation.
642
Anderson
Figure 199. Hubbs-Hubbs diagram illustrating variation among specimens of Cleonidius species; width frons compared to width at apex of rostrum (WRA/WF) - a, males; b, females. See
caption for Fig. 49 for explanation.
New World Cleonini
643
Quaest. Ent., 1987, 23 (4)
compared to length of elytra (LP/LE1); 201, form of elytra (WE1M/LE1).
644
Anderson
Figures 202-203. Maps illustrating position of collecting localities for: 202, Stephanocleonus immaculatus\ 203, S.
parshus.
New World Cleonini
645
Figures 204-205. Maps illustrating position of collecting localities for: 204, Stephanocleonus confusus (dots) and S.
stenothorax (stars); 205, S. plumbeus (dots) and S. cristaticollis (stars).
Quaest. Ent., 1987, 23 (4)
646
Anderson
Figures 206-209. Maps illustrating position of collecting localities for: 206, Apleurus saginatus\ 207, A. lutulenlus\ 208,
A. jacobinus\ 209, A. albovestitus.
New World Cleonini
647
Figures 210-213, Maps illustrating position of collecting localities for: 210, Apleurus aztecus (dots) and A. porosus
(stars); 21 1, A. angularis\ 212, A. hystrix ; 213, Cleonis pigra.
Quaest. Ent., 1987, 23 (4)
648
Anderson
Figures 214-219. Maps illustrating position of collecting localities for: 214, Cleonidius longinasus\ 215, C. subcylindricus\
216, C. eustictorrhinus\ 217, C. pleuralis\ 218, C. texanus\ 219, C. erysimi.
New World Cleonini
649
Figures 220-225. Maps illustrating position of collecting localities for: 220, Cleonidius trivittatus\ 221, C. boucardi\ 222,
C. placidus (dots) and C. infrequens (stars); 223, C. americanus (dots) and C. collaris (stars); 224, C. quadrilineatus ;
225, C. notolomus (dots) and C. puberulus (stars).
Quaest. Ent., 1987,23 (4)
650
Anderson
Figures 226-229. Maps illustrating position of collecting localities for: 226, Cleonidius frontalis-, 227, C. canescens\ 228,
C. poricollis ; 229, C. calandroides.
New World Cleonini
651
Figure 230. Numbers of species of Apleurus in 5° intervals.
Quaest. Ent., 1987, 23 (4)
652
Anderson
Figure 231. Numbers of species of Cleonidius in 5° intervals.
New World Cleonini
653
N = 26
N = 8
N = 40
N = 7
N= 1 4
N = 56
N=7
N = 38
N = 56
N = 7
N=4
N = 3
N = 1 4
N= 1 32
N= 1 0
N = 29
N=42
N= 1 0
N = 63
Figures 232-233. Hubbs-Hubbs diagrams illustrating variation among specimens of: 232, Apleurus species; elevation; 233,
Cleonidius species; elevation. See caption for Fig. 49 for explanation.
Quaest. Ent., 1987, 23 (4)
654
Anderson
o
A-
t
t
3(3)
5,8(3)
6 — 6 — 0 6 — 0
4 2, 3(2,3) ,4(3) ,5(2,3) ,6
MONOPHYLY CHARACTERS
234
Figure 234. Reconstructed phylogeny based on primary characters illustrating hypothesized phylogenetic relationships
among Apleurus species and related out-groups. Dots denote apotypic character states, open circles plesiotypic states. See
“Phylogeny” section and Tables I and II for discussion of characters and states (numbers in parentheses).
New World Cleonini
655
Figure 235. Reconstructed phylogeny based on primary and secondary characters illustrating hypothesized phylogenetic
relationships among Apleurus species and related out-groups. Dots denote apotypic character states, open circles
plesiotypic states. See “Phylogeny” section and Tables I and II for discussion of characters and states (numbers in
parentheses).
Quaest. Ent., 1987, 23 (4)
Cyl Indropterus
656
Anderson
<J<J<J<J<J<J <J<J000<J0 00 0000
Figure 236. Reconstructed phylogeny based on primary characters illustrating hypothesized phylogenetic relationships
among Cleonidius species and related out-groups. Dots denote apotypic character states, open circles plesiotypic states.
See “Phylogeny” section and Tables III and IV for discussion of characters and states (numbers in parentheses).
quadr i 1 i neat us
New World Cleonini
657
Figure 237. Reconstructed phylogeny based on primary and secondary characters illustrating hypothesized phylogenetic
relationships among Cleonidius species and related out-groups. Dots denote apotypic character states, open circles
plesiotypic states. See “Phylogeny” section and Tables III and IV for discussion of characters and states (numbers in
parentheses).
Quaest. Ent., 1987,23 (4)
658
Anderson
Mojave Desert
Sonoran Desert
Chihuahuan Desert
Great Basin Desert
Texas Semi-desert
Pacific Semi-desert
Deser t -Grass 1 and
Grass 1 and
Grass 1 and - Dec . For
Mesquite Grassland
Figure 238. Habitat associations of Apleurus species Solid box denotes marked association, half-box slight association.
Phylogeny following 235.
New World Cleonini
659
Asclepiadaceae
Chenopod i aceae
Compos i tae
Cruci ferae
Ephedraceae
Grami nae
Leguminosae
Malvaceae
Polygonaceae
Rosaceae
Sol anaceae
Umbel 1 i ferae
Zygophyl 1 aceae
239
Figure 239. Plant associations of Apleurus species Solid box denotes marked association, half-box slight association, check
mark few collection records. Plant taxa are listed alphabetically. Phytogeny following 235.
Quaest. Ent., 1987, 23 (4)
660
Anderson
Mojave Desert
Sonoran Desert
Chihuahuan Desert
Great Basin Desert
Texas Semi-desert
Pacific Semi-desert
Desert -Grass 1 and
Grassland
Grass 1 and- Dec . For .
S . E . Meso . Ever . For .
Wes t . Xero . Ever . For .
N . W. Meso. Ever . For .
Mesquite Grassland
Eastern Beaches
240
Figure 240. Habitat associations of Cleonidius species Solid box denotes marked association, half-box slight association.
Phylogeny following 237.
New World Cleonini
661
Asclepiadaceae
Capri fol iaceae
Chenopodi aceae
Compos i tae
Cruci ferae
Ephedraceae
Ericaceae
Legum i nosae
Loasaceae
Polygonaceae
Rhamnaceae
Rosaceae
Umbel 1 i ferae
Figure 241. Plant associations of Cleonidius species. Solid box denotes marked association, half-box slight association,
check mark few collection records. Plant associations are unknown for some species. Plant taxa are listed alphabetically.
Phylogeny following 237.
Quaest. Ent., 1987, 23 (4)
662
Anderson
Figure 242. Origin and initial diversification of Apleurus in North America: A. hystrix , A. azlecus , and A. lutulentus-A.
porosus-A. angularis-A. jacobinus-A. albovestitus-A. saginatus lineage. See text for detailed explanation.
New World Cleonini
663
Figures 243-245. Diversification of Apleurus in North America (see text for detailed explanation; dashed lines indicate
hypothesized ancestral distribution, solid lines indicate present distributions. 243, A. lutulentus and A. porosus-A.
angularis-A. jacobinus-A. albovestitus-A. saginatus lineage; 244, A. angularis , A. jacobinus and A. porosus\ 245, A.
albovestitus and A. saginatus.
Quaest. Ent., 1987, 23 (4)
664
Anderson
Figure 246. Origin and initial diversification of Cleonidius in North America: Cleonidius erysimi group, C. americanus
group, C. poricollis group, and C. boucardi group. See text, and caption for 242 for detailed explanation.
New World Cleonini
665
Figures 247-249. Diversification of Cleonidius in North America. See text, and captions for 243-245 for detailed
explanation. 247, C. erysimi-C. eustictorrhinus, C. pleuralis, and C. subcylindricus-C. longinasus-C. texanus lineages;
248, C. erysimi and C. eustictorrhinus-, 249, C. subcylindricus, C. longinasus and C. texanus.
Quaest. Ent., 1987, 23 (4)
666
Anderson
Figures 250-253. Diversification of Cleonidius in North America. See text, and captions for 243-245 for detailed
explanation. 250, Cleonidius americanus-C. frontalis , C. canescens-C. infrequens-C. puberulus-C. collaris-C. notolomus
lineages; 251, C. americanus and C. frontalis-, 252, C. puberulus and C. infrequens ; 253, C. collaris and C. notolomus.
New World Cleonini
667
Figures 254-256. Diversification of Cleonidius in North America. See text, and captions for 243-245 for detailed
explanation. 254, Cleonidius boucardi and C. trivittatus-C. placidus-C. quadrilineatus lineage; 255, C. trivittatus and C.
placidus-C. quadrilineatus lineage; 256, C. placidus and C. quadrilineatus.
Quaest. Ent., 1987, 23 (4)
668
Anderson
Figures 257-258. Geographic ranges of species in: 257, Cleonidius erysimi species group; 258, C. americanus species
group.
New World Cleonini
669
259. Geographic ranges of species in the Cleonidius boucardi species group
Quaest. Ent., 1987, 23 (4)
670
Anderson
Table 1. Descriptive statistics for S. confusus based on eight males and eight females from
Medicine Hat, Alberta
Males
A. Measurements, in mm.
New World Cleonini
671
Table 2. Descriptive statistics for S. cristaticollis based on one male and one female from
Medicine Hat, Alberta
Female
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
672
Anderson
Table 3. Descriptive statistics for S. immaculatus based on eight males and eight females from
throughout the species range
Females
A. Measurements, in mm.
New World Cleonini
673
Table 4. Descriptive statistics for S. parshus based on eight males and eight females from
southern Manitoba and Ontario
Females
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
674
Anderson
Table 5. Descriptive statistics for S. plumbeus based on eight males and eight females from
Riviere - au - Tonnerre, Quebec
Females
A. Measurements, in mm.
New World Cleonini
675
Table 6. Descriptive statistics for S. stenothorax based on eight males and eight females from
Bluefish Caves Archaeological Site, Yukon Territory
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
676
Anderson
Table 7. Descriptive statistics for A. hystrix based on eight males and eight females from the El
Segundo Sand Dunes, Los Angeles County, California
Females
A. Measurements, in mm.
New World Cleonini
677
Table 8. Descriptive statistics for A. aztecus based on 13 males and six females from
throughout the species range
Males
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
678
Anderson
Table 9. Descriptive statistics for A. lutulentus based on eight males and eight females from 2
mi. NE. Portal, Arizona
Females
A. Measurements, in mm.
New World Cleonini
679
Table 10. Descriptive statistics for A. porosus based on seven males and eight females from
68.0 km. W. Ejido Viscaino, Baja California Sur, Mexico
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
680
Anderson
Table 11. Descriptive statistics for A. angularis based on 15 males and 10 females from
Mercury, Nevada
New World Cleonini
681
Table 12. Descriptive statistics for A. jacobinus based on eight males and eight females from
Turlock, California
Females
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
682
Anderson
Table 13. Descriptive statistics for A. albovestitus based on eight males and eight females from
Portal, Arizona
Females
A. Measurements, in mm.
New World Cleonini
683
Table 14. Descriptive statistics for A. saginatus based on eight males and eight females from 5
mi. SW. Patagonia, Arizona
Males
A. Measurements, in mm.
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
684
Anderson
Table 15. Descriptive statistics for C. erysimi based on eight males and eight females from
Redondo, California
New World Cleonini
685
Table 16. Descriptive statistics for C. eustictorrhinus based on 13 males and 14 females from
throughout the species range
Females
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
686
Anderson
Table 17. Descriptive statistics for C. pleuralis based on eight males and eight females from
Palm Springs, California
Males
A. Measurements, in mm.
Females
A. Measurements, in mm.
New World Cleonini
687
Table 18. Descriptive statistics for C. subcylindricus based on seven males and 17 females from
throughout the species range
Males
A. Measurements, in mm.
Females
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
688
Anderson
Table 19. Descriptive statistics for C. longinasus based on 10 males and 10 females from
Hemet Reservoir, Herkey Creek, and Idyllwild, California
Females
A. Measurements, in mm.
New World Cleonini
689
Table 20. Descriptive statistics for C. texanus based on eight males and eight females from
Mile 19, Hitchcock Highway, Santa Catalina Mountains, Arizona
Quaest. Ent., 1987,23 (4)
690
Anderson
Table 21. Descriptive statistics for C. americanus based on eight males and eight females from
Mercy Hot Springs, California
Females
A. Measurements, in mm.
New World Cleonini
691
Table 22. Descriptive statistics for C. frontalis based on eight males and eight females from
Medicine Hat, Alberta
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
692
Anderson
Table 23. Descriptive statistics for C. canescens based on eight males and eight females from
Denver, Colorado
Females
A. Measurements, in mm.
New World Cleonini
693
Table 24. Descriptive statistics for C. infrequens based on two males and six females from
throughout the species range
Females
A. Measurements, in mm.
Quaest. Ent., 1987, 23 (4)
694
Anderson
Table 25. Descriptive statistics for C. puberulus based on six males and 10 females from
Colorado Springs, Colorado
Females
A. Measurements, in mm.
New World Cleonini
695
Table 26. Descriptive statistics for C. collaris based on seven males and six females from
throughout the species range
Males
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
696
Anderson
Table 27. Descriptive statistics for C. notolomus based on 11 males and 13 females from
throughout the species range
New World Cleonini
697
Table 28. Descriptive statistics for C. poricollis based on eight males and eight females from
lmi. N. Little Lake, California
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
698
Anderson
Table 29. Descriptive statistics for C. calandroides based on eight males and eight females
from Hampton, New Hampshire
Females
A. Measurements, in mm.
New World Cleonini
699
Table 30. Descriptive statistics for C. boucardi based on eight males and eight females from
north of Tepatitlan, Jalisco, Mexico
Males
A. Measurements, in mm.
Females
A. Measurements, in mm.
Quaest. Ent., 1987,23 (4)
700
Anderson
Table 31. Descriptive statistics for C. trivittatus based on eight males eight females from
Albuquerque, New Mexico
Females
A. Measurements, in mm.
New World Cleonini
701
Table 32. Descriptive statistics for C. placidus based on 10 males and eight females from
Panoche, California
Quaest. Ent., 1987, 23 (4)
702
Anderson
Table 33. Descriptive statistics for C. quadrilineatus based on eight males and eight females
from southern Arizona localities
APPENDIX 2
New World Cleonini
703
Appendix II: Species of genera of Cleoninae examined for purposes of determining relationships
of New World taxa. Genus-group name is listed first, followed by species examined on loan
from BMNH. An asterisk indicates that the species examined is the type-species of that
generic-group name. No asterisk indicates that the type species was not examined because, no
type-species has yet been designated for that genus-group name; or, type species was not
available for study in the BMNH.
Adosomus Faust
Amblysomus Faust
Ammocleonus Bedel
Atactogaster Faust
Bothynoderes Schoenherr
Brachycleonus Faust
Calodemas Faust
Centrocleonus Chevrolat
Chromoderus Motschulsky
Chromonotus Motschulsky
Chromosomus Motschulsky
Cnemodontus Chevrolat
Coniocleonus Motschulsky
Conorhynchus Motschulsky
Cosmogaster Faust
Cylindropterus Chevrolat
Cyphocleonus Motschulsky
Epirhynchus Schoenherr
Eumecops Hochhuth
Eurycleonus Bedel
Gonocleonus Chevrolat
Isomerus Motschulsky
Koenigius Heyd.
Leucochromus Motschulsky
Leucomigus Motschulsky
Leucosomus Motschulsky
Liocleonus Motschulsky
Lixocleonus Marshall
Lixomorphus Faust
Mecaspis Schoenherr
Menocleonus Faust
Microcleonus Faust
Nemoxenus Faust
Neocleonus Chevrolat
A. granulosus (Mannerheim)
A. brevis (Fahraeus) *
A. hieroglyphicus (Olivier) *
A. orientalis (Chevrolat) *
B. punctiventris (Germar) *
B. fronto (Fischer von Waldheim) *
C. vetustus Faust
C.fallax (Fahraeus) *
C. albidus (Fabricius) *
[ = C. fasciatus (Vill.)]
C. vittatus (Hochhuth) *
C.fischeri (Fahraeus) *
C. limpidus (Gyllenhal)
C. excoriatus (Gyllenhal) [type species,
C. carinirostris (Gyllenhal)]
C. conirostris (Gebler) *
C. lateralis (Gyllenhal) *
C. luxeri Chevrolat *
C. cenchrus (Pallas) *
E. humerosus Faust [type species,
E. argus (Sparrm.)]
Eumecops sp. [type species,
E. kittaryi Hochhuth]
E. baluchicus Marshall [type species,
E. gigas Mars.]
G. helferi Chevrolat *
I. granosus (Zoubkoff) *
K. palaestinus Heyd. *
L. imperialis (Zoubkoff) *
L. candidatus (Pallas) *
L. opthalmicus (Rossi) *
L. clathratus (Olivier) *
L. incanus Marshall *
L. ocularis (Fabricius) *
M. palmatus (Olivier)
M. implicatus Faust
M. panderi (Fischer von Waldheim) *
TV. zebra (Chevrolat)
TV. sannio (Herbst) *
704
Anderson
457
Nomimonyx Faust
Pachycerus Schoenherr
Pentatropis Faust
Phaulosomus Faust
Pleurocleonus Motchulsky
Porocleonus Motschulsky
Prionorhinus Chevrolat
Pseudocleonus Chevrolat
Rhabdorhynchus Motschulsky
Stephanocleonus Motschulsky
Temnorhinus Chevrolat
Tetragonothorax Chevrolat
Trichocleonus Motschulsky
Xanthochelus Chevrolat
Xenomacrus Faust
N. perturbans Faust *
P. cordiger (Germar) [type species,
P. scabrosus Brulle]
P. sparsus (Fahraeus) [type species,
P. formosus (Fahraeus)]
P. kilimanus Faust
P. quadrivittatus (Zoubkoff) *
P. candidus (Olivier) *
P. stillatus (Gyllenhal)
P. cinereus (Schrank)
R. anchusae Chevrolat [type species,
R. menetriesi (Gyllenhal)]
S. flaviceps (Pallas) *
T. brevirostris (Gyllenhal) *
T. retusus (Fabricius) *
T. leucophyllus (Fischer von Waldheim) *
X. longus Chevrolat *
X. glacialis (Herbst)
Quaest. Ent., 1987, 23 (4)
New World Cleonini
705
INDEX TO NAMES OF TAXA
(Synonyms in italics)
FAMILY GROUP TAXA
Amphibia, 446
Anobiidae, 549
Asclepiadaceae, 478, 506, 530
Capparaceae, 478, 502
Caprifoliaceae, 520
Carabidae, 466
Cerambycidae, 549
Chenopodiaceae, 445-446, 478, 484, 490,
506,517,520,527,531,579, 581
Chrysomelidae, 549
Cleoninae, 435, 437, 445-447, 450, 452,
467, 471, 546-550, 552, 591, 594
Cleonini, 435, 437, 445-446, 550, 594
Coleoptera, 435
Compositae, 445-446, 452, 470, 478, 480,
484, 486, 490-492, 502, 506, 515, 531,
579,581,585,588,590-591
Cruciferae, 445-446, 490, 502, 506, 523,
525,530-531,533,579,581,588
Curculionidae, 434, 446, 549, 591
Curculionoidea, 435
Cylydrorhinini, 546
Diptera, 446
Entiminae, 546
Ephedraceae, 490, 506
Ericaceae, 504
Euphorbiaceae, 470
Geomorinae, 446
Graminae, 490
Hylobiinae, 546, 549
Hymenoptera, 446
Labiatae, 531
Lagriidae, 549
Leguminosae, 445, 470, 478, 486, 490,
504, 506, 508, 511-512, 517, 520, 531,
541, 543, 574, 579, 581, 587-588, 590,
593
Leptopiinae, 546
Lepyrini, 550
Lixini, 445-446, 550
Loasaceae, 520
Malvaceae, 478, 486, 490
Molytinae, 546, 549-550, 552
Polygonaceae, 445, 490, 504
Rhamnaceae, 511-512, 543, 574, 581,
587, 590, 593
Rhinocyllini, 550
Rosaceae, 445, 461, 490, 502, 511-512,
531,543,579, 581,587, 590, 593
Roseaceae, 574
Solanaceae, 490
Tachinidae, 446
Umbelliferae, 486, 541
Vitaceae, 490
Zygophyllaceae, 446, 490
GENERA AND SUBGENERA
Adosomus Faust, 450-451
Apleurus Chevrolat, 435, 445, 449, 466,
468-472, 547, 550-551, 553, 576, 582,
586, 591,593-594
Bangasternus Gozis, 447, 550
Bothynoderes Schoenherr, 455
Centrocleonus Chevrolat, 471
Centrocleonus LeConte, 470-471
Chromonotus Motschulsky, 455
Chromosomus Motschulsky, 455
Cleonaspis LeConte, 470
Cleonidius Casey, 435, 440, 445, 447-449,
471, 492-493, 547, 550, 564, 572, 578,
582-583, 586-587, 591, 593-594
Cleonis Dejean, 445, 447, 449-451, 471
Cleonopsis LeConte, 470-471
Cleonus Schoenherr, 449
Coniocleonus Motschulsky, 453
Conorhynchus Motschulsky, 455
Cyphocleonus Motschulsky, 447, 450-451
Dinocleus Casey, 471-472, 476
Epimeces Billberg, 450
Geomorus Schoenherr, 449-450
Gibbostethus Anderson, new subgenus,
445, 447, 468, 472
Ileomus Schoenherr, 449
Larinus Schoenherr, 448
Lixestus Reitter, 493
Lixus Fabricius, 446, 448, 450, 471, 493,
Quaest. Ent., 1987, 23 (4)
706
Anderson
552-553, 583
Menecleonus Faust, 455
Microlarinus Hochhuth, 447, 551
Plagiographus Chevrolat, 453
Pleurocleonus Motschulsky, 455
Rhinocyllus Germar, 447, 550
Stephanocleonus Motschulsky, 435, 440,
445, 449, 453, 455, 458-460, 463, 558,
594
Temnorhinus Chevrolat, 455
Xanthochelus Chevrolat, 455
SPECIES AND SUBSPECIES
acradenius (Greene) Blake, Aplopappus,
490, 531
affinis J.G. Agardh., Lupinus, 512
albifrons Benth., Lupinus, 470
albovestitus (Casey), Apleurus, 468, 470,
474, 476, 478, 484, 486-487, 489, 491,
576-577, 585, 592
albovestitus Casey, Dinocleus , 486-487
altissimum L., Sisymbrium, 531
americanus Csiki, Cleonidius, 495, 498,
513, 573, 578-579, 588-589
americanus Csiki, Cleonus, 513
amygdalus Batsch., Prunus, 512
angularis (LeConte), Apleurus, 468, 470,
473, 476, 479, 481, 484-486, 492,
576-577, 586, 590, 592
angularis LeConte, Cleonus, 470-471, 481
arcadenius (Greene) Blake, Aplopappus,
484
arvense (L.), Cirsium, 452
aztecus (Champion), Apleurus, 468, 472,
474, 478, 576-577, 584-585, 590
aztecus Champion, Cleonus , 472, 474
basalis Fall, Cleonus , 513
bicarinatus Casey, Cleonus, '536-537
bisulcatus (Hook.) Gray, Astragalus, 538
boucardi (Chevrolat), Cleonidius, 496,
498, 500, 534, 537, 573, 579, 589
boucardi Chevrolat, Apleurus , 471, 534
breyeri Brethes, Argentinorhynchus, 546
bryanti Van Dyke, Dinocleus , 472, 476
calandroides (Randall), Cleonidius,
496-498, 505, 508, 531, 533, 572,
578-579, 581, 588-589
calandroides Randall, Lixus , 531
californica (DC.) Torr. and Gray,
Gutierrezia, 531
californica Harv., Dilthyrea, 502
californica Wats., Ephedra, 490, 506
californicus (Aves), Geococcyx, 446, 484
californicus Motschulsky, Lixus, 505, 545
californicus Muell.-Arg., Croton, 470
campestris L., Brassica, 531
canescens (LeConte), Cleonidius, 495,
499, 517, 533, 544, 572, 581, 589, 592
canescens (Pursh) Nutt., Atriplex, 517,
520, 527
canescens LeConte, Cleonus, 483, 517
capillosus Csiki, Cleonus , 484
capitatum (Dougl.) Greene, Erysimum,
502
carinirostris Gyllenhal, Cleonus , 453
carinirostris Gyllenhal, Coniocleonus, 453
carinocollis LeConte, Cleonus , 534
cernua DC., Flourensia, 478, 490
chilensis Blanchard, Cleonis, 546
cineritius Gyllenhal, Coniocleonus, 453
circumductus Casey, Cleonus , 510
collaris (LeConte), Cleonidius , 495, 523,
525,533,573,581
collaris LeConte, Cleonus , 496, 523
coloradensis Csiki, Cleonus , 518
communis L., Pyrus, 512
confertiflora (DC.) Rydb., Franseria, 490,
531
confusus Anderson, Stephanocleonus,
455-457, 459-460, 463-464, 590
conicus Froelich, Rhinocyllus, 445-446,
452
cooperi (Gray) Greene, Psilostrope, 484
cristaticollis Csiki, Cleonus , 459
cristaticollis Csiki, Stephanocleonus,
455-456, 459, 463
cumulicola Small, Lupinus, 508
cuneatus (Hook.) Nutt., Ceanothus, 453,
512
densus Casey, Dinocleus , 486-487
dentatus Champion, Dinnocleus, 481, 491
New World Cleonini
707
denticollis Casey, Dinocleus, 481, 483
denudatus Zoubkoff, Lixus, 543
diffusa Lam, Centaurea, 452
diffusus Nutt., Lupinus, 508
dumosa Gray, Franseria, 490
earlei Greene ex. Rydb., Astragalus, 538
edentula (Bigel.) Hook., Cakile, 533
enceloides (Cav.) Benth. and Hook.,
Verbesina, 478, 490
ericoides (Less.) H. & A., Aplopappus,
470, 502
erysimi (Fall), Cleonidius, 495, 497, 501,
503-504, 572, 581, 588, 591
erysimi Fall, Cleonus, 500, 502
eustictorrhinus Anderson, Cleonidius, 495,
498, 501-503, 508, 572, 578-579, 588
excoriatus Gyllenhal, Coniocleonus, 453
farctus Casey, Dinocleus , 479
fasciaturs Villers, Curculio, 451
fasciatus Gmelin, Curculio , 451
fasciculatum Benth., Eriogonum, 490
fasciculatum H. & A., Adenostoma, 502
ferrugineus Fahraeus, Coniocleonus, 453
filiformis Fabricius, Curculio, 450
flaviceps Palliser, Stephanocleonus, 453
flavus Nutt., Astragalus, 538
floridum Benth., Cercidium, 490
fossus Chevrolat, Apleurus, 466, 470-471,
475-476
frontalis (LeConte), Cleonidius, 495, 499,
515, 525, 573, 588-589
frontalis LeConte, Cleonus , 515
glariuscula DC, Chaenactis, 470
glaucus Fabricius, Coniocleonus, 453
glutinosa Pers., Baccharis, 478, 490, 492
grandirostris Casey, Cleonus , 5 1 8
graniferus Casey, Cleonus , 506
hirsutum L., Gossypium, 478, 486
hystrix (Fall), Apleurus, 468-469, 495,
576-577, 584
hystrix Fall, Dinocleus, 468-469
immaculatus Anderson, new species,
Stephanocleonus, 455, 459-460,
463-465
indicus Fahraeus, Cleonus , 451
infrequens Anderson, Cleonidius, 495,
499, 520, 525, 573, 578, 581, 589
inornatus LeConte, Cleonus , 536-537,
539
integerrimus Hook, and Arn., Ceanothus,
512
interruptus Casey, Dinocleus , 487
jacobinus (Casey), Apleurus, 468, 470,
473, 479, 484, 486, 491, 576-577, 586,
590
japonicus (Faust), Cleonus, 450-451
juliflora (Swartz) DC., Prosopis, 490
kali L., Salsola, 490, 530
karelini Boheman, Lixus , 543
kirbyi Casey, Cleonus , 528
laciniata Vasey and Rose, Encelia, 480
lambertii Pursh, Oxytropis, 538
lanata (Pursh), Eurotia, 530
lecontei Casey, Cleonus , 534
lecontellus Csiki, Cleonus , 534
ledifolius Nutt., Cercocarpus, 512
lentiformis (Torr.) Wats., Atriplex, 484,
490
leucodermis Greene, Ceanothus, 512
linearifolius DC, Aplopappus, 484
linearis (Cav.) Lag., Palafoxia, 490
lineaticollis Kirby, Lepidophorus, 466
lobigerinus Casey, Cleonus , 528
longinasus Anderson, new name,
Cleonidius, 495, 498, 508, 572
lucida Greene, Gutierrezia, 484
ludoviciana (Nutt.) Wats., Lesquerella,
523
lupinus Blatchley, Lixus, 506
lutulentus (LeConte), Apleurus, 468, 471,
473-478, 484, 491-492, 576-577,
584-585
lutulentus LeConte, Cleonus, 466, 470,
475-476
luxeri Chevrolat, Cylindropterus, 495
maritima (L.) Desv., Lobularia, 502
meles Boheman, Lixus, 495, 544
mexicana D. Don., Cowania, 512
mexicanus Casey, Dinocleus , 487
microcephala (DC.) Gray, Gutierrezia,
484, 490,531
mixtus LeConte, Lixus , 508
Quaest. Ent., 1987, 23 (4)
708
Anderson
modestus Mannerheim, Lixus, 505, 545
molitar LeConte, Centrocleonus, 487
molitor LeConte, Cleonus, 486
monogyra T. and G., Hymenoclea, 492,
531
montanus Raf., Cercocarpus, 512, 543
multiradiata Harv. and Gray, Baileya, 490
nauseosus (Pall.) Britton, Chrysothamnus,
478, 484, 490-492, 506, 515, 531
nebulosus Knoch, Curculio, 451
notolomus Anderson, Cleonidius, 495, 499,
525.533.573.578.581.589
nutalli Wats., Atriplex, 530
oleracea L., Brassica, 502, 531
oreophila Woot. and Standi., Verbesina,
478
pacificus Fall, Cleonus , 539
pallasi Faust, Lixus, 495, 544
paniculatus (Gral) H.M. Hall,
Chrysothamnus, 506
paraplecticus Linnaeus, Curculio, 450
parshus Anderson, new species,
Stephanocleonus, 455-456, 459,
461-465
persica (L.) Batsch., Prunus, 490, 502, 512
piger Scopoli, Curculio , 451
pigra (Scopoli), Cleonis, 435, 442,
450-453
pilosus LeConte, Centrocleonus, 484
pinnata (Pursh.) Britton, Stanleya, 502
pinnata (Walt.) Britton, Descurainia, 531
placidus Casey, Cleonus , 539
placidus Csiki, Cleonidius, 496, 499, 537,
539, 541,543,573,578,590
pleniradiata Harv. and Gray, Baileya, 478,
490
pleuralis (LeConte), Cleonidius, 495-496,
499. 504. 589
pleuralis LeConte, Lixus , 504
plumbeus LeConte, Stephanocleonus, 455,
457-458,461-465, 546
polycarpa (Torr.) Watts., Atriplex, 506
porcatus Casey, Dinocleus , 48 1 , 483
poricollis (Mannerheim), Cleonidius,
496-497, 499, 505, 525, 527-528,
533-534, 572, 579, 581, 588-589
poricollis (Mannerheim), Cleonus , 528
poricollis Mannerheim, Lixus , 493, 500,
527-528, 545
porosus (LeConte), Apleurus, 468, 473,
479, 484-486, 491, 576-577, 586, 590
porosus LeConte, Centrocleonus , 478
praepotens Say, Rhynchophorus, 536-537
praepoteus (Say), Lixus , 537
psilostachya DC., Ambrosia, 490
puberulus (LeConte), Cleonidius , 495,
521, 525, 573, 578, 581, 589
puberulus LeConte, Cleonus , 521
pulvereus LeConte, Cleonus, 470, 475-476
pungens (H. and A.) T. and G.,
Hemizonia, 486
quadrilineatus (Chevrolat), Cleonidius,
496, 500, 512, 534, 537, 539, 541,
572-573, 579, 587, 590
quadrilineatus Chevrolat, Apleurus , 471,
541
rosea L., Atriplex, 530
saginatus (Casey), Apleurus, 468,
473-474, 477, 484, 487, 491-492,
576-577
saginatus Casey, Dinocleus , 491
salsola T. and G., Hymenoclea, 490, 506
sardous (Chevrolat), Cleonus, 450-451
sativa L., Medicago, 490, 506, 520, 531
sativa L., Pastinaca, 541
sativus L., Raphanus, 490, 502, 531
sericea (Nutt.) Coville, Pluchea, 490
sericia Nutt., Oxytropis, 538
sophia (L.) Webb., Descurainia, 531
sparsus LeConte, Cleonus , 515
spinosa Zucc., Koeberlinia, 478
stenothorax Anderson, new species,
Stephanocleonus, 455-456, 465
stevioides Hook, and Arn.„ Chaenactis,
484
stratus Csiki, Cleonus , 515
subcylindricus Casey, Cleonidius, 495,
497, 503, 506, 508, 533, 578, 589
subcylindricus Casey, Cleonus , 506
subverticillata (Gray) Vail, Asclepias, 478
suffrutescens (Abrams) G. Rossb.,
Erysimum, 502
New World Cleonini
709
sulcirostris Linnaeus, Curculio , 449-451
tenuisectus (Greene) Blake, Aplopappus,
484
tesota Gray, Olneya, 490
texanus (LeConte), Cleonidius, 495, 498,
501, 503, 508, 510, 512, 543, 572, 587,
589
texanus LeConte, Lixus, 510
thulis Kissinger, Viatavitus, 466
transversofasciatus Goeze, Curculio , 45 1
tridentata (DC.) Coville, Larrea, 490
tridentata (Pursh.) DC., Purshia, 512, 531
tridentata Nutt., Artemesia, 490, 515
trisulcatus (Herbst), Cyphocleonus, 451
trivittatus (Say), Cleonidius, 471, 496,
500, 525, 534, 536-537, 539, 541, 573,
579, 589
trivittatus Chevrolat, Apleurus, 471
trivittatus Say, Cleonus, 536, 539
utahensis Torr. and Gray, Astragalus, 538
venetus (Hbk.) Blake, Aplopappus, 531
vermiculatus (Hook.) Torr., Sarcobatus,
490
vibex (Pallas), Cleonidius, 442, 496, 543,
572
vibex Pallas, Curculio, 493, 543
vibex Pallas, Lixus, 493, 495
vibex scutellaris Petri, Lixus , 544
virgatus LeConte, Cleonus , 527-528
viscidiflorus (Hook.) Nutt.,
Chrysothamnus, 484, 531
vittatus Kirby, Cleonis, 493, 527-528
vittatus Kirby, Cleonus , 528
vulgaris L., Beta, 530
vutgare (Savi), Cirsium, 452
wickhami Casey, Dinocleus, 487
wislizeni Gray, Hymenothrix, 490
wootoni Sheldon, Astragalus, 538, 543
Quaest. Ent., 1987, 23 (4)
711
BOOK NOTICES
ENTOMOFAUNAL PUBLICATIONS FROM THE AUSTRALIAN REGION
In recent years, Australian and New Zealand government and museum entomologists have
undertaken preparation of major series of faunistic treatments which are of substantial value
not only locally but also more distantly, for those extra-territorial biologists who want to gain
some level of familiarity with the notogaean arthropod fauna. Previously, Quaestiones
Entomologicae has published extensive reviews or notices of two series: Fauna of New Zealand,
and A Guide to the Genera of Beetles of South Australia. Series previously unnoticed in the
pages of Quaest. Ent. are the Zoological Catalogue of Australia, and the Fauna of Australia.
We have not yet seen any parts of the last named series. Below, comments are offered about
recently received parts of the first three series noted.
HARRIS, A. C. 1987. Pompilidae (Insecta: Hymenoptera). Fauna of New Zealand [no.]
12, 154 pp. DSIR Science Information Publishing Centre, P.O. Box 9741, Wellington, New
Zealand. Price $39.95 (New Zealand), $34.25 (Canadian).
The 11 species of spider wasps (10 endemic, one introduced from Australia) representing
four genera included in two subfamilies, are treated in fulsome manner, including for each
species data about: names; structural features of adults and larvae; and way of life, including
hunting of spiders by adult females, “nidification formula”, nest site and structure, life history,
and emergence and copulation. Variation in color and color pattern are clearly indicated by
stylized diagrams, and various structural features and nests are illustrated by line drawings and
photographs. The frontspiece is a superb photograph of a paralyzed ctenizid spider with a larval
pompilid attached.
To aid in identification of taxa, illustrated keys are provided to adults (males and females),
larvae, and type of nesting behavior.
An introductory part of the publication treats an appropriately wide variety of biological
information about New Zealand pompilids: structural features of adults and larvae,
geographical variation, mimicry, nesting behavior, biogeographic relationships, and
evolutionary relationships. I found especially interesting the author’s observation about
mimicry. Three complexes are recognized, based on color pattern of body and wings, and
pattern of movement of adults. Each complex has Mullerian and Bastesian components,
including collectively, elaterid beetles, asilid and calliphorid flies, ichneumonid, proctotrupid,
and sphecid wasps, and bees.
The substantial amount of attention devoted to prey capture and nesting behavior is
supplemented by an appendix that records, in association with the name of the predatory wasp
species, family, genus and species of the spider prey. Information of this kind will be of interest
to general biologists and behaviorists.
Overall, the author has provided a study that easily meets the very high standards associated
with publications about aculeate Hymenoptera, in the tradition of such masters as Howard E.
Evans, Karl V. Krombein, and Charles D. Michner.
MATTHEWS, E. G. 1987. A guide to the beetles of South Australia. Part 5 Polyphaga:
Tenebrionoidea, 67 pp. Special Educational Bulletin Series (No. 8), South Australian Museum,
Adelaide, Australia.
Quaest. Ent., 1987, 23 (4)
712
This volume, the primary purpose of which is to provide a simplified means of identifying
adult beetles to genus, consists principally of illustrated keys and 135 habitus illustrations-
some excellent line drawings, and some photographs- one such illustration for each genus. In a
chapter preceding the keys, each of the 17 families of tenebrionoids of South Australia is
briefly characterized by comparison or by reference to diagnostic structural features, and by
reference to way of life. Under each family, genera are listed, and number of species of each
genus in South Australia is reported. For the markedly speciose Tenebrionidae (including
Alleculidae, Cossyphodidae, Lagriidae, and Nilionidae), the genera are grouped by tribe and
subfamily.
The volume is attractively bound with a soft cover, on which is a color illustation of a
brownish “pie-dish” tenebrionid of the genus Helea on brownish pebbly soil. This useful
publication continues the high standard set in the previous volumes in this series.
LAWRENCE, J. F„ B. P. MOORE, J. E. PYKE, and T. P. WEIR. 1987. Volume 4
Coleoptera Archostemata, Myxophaga and Adephaga. viii + 444 pp. In , Zoological catalogue
of Australia (D. W. Walker, Executive Editor). Australian Government Publications Services,
Mail Order Sales, G. P. O. Box 84, Canberra A. C. T. 2601. Price $34.95 (Australian), $33.54
(Canadian).
This volume catalogues the taxa of three suborders: Archostemata, including families
Cupedidae and Ommatidae, and Myxophaga, family Microsoridae (Lawrence, Weir, and
Pyke); Adephaga- Geodephaga, including families Rhysodidae and Carabidae (Moore, Weir,
and Pyke); and Adephaga- Hydradephaga, including families Haliplidae, Hygrobiidae,
Noteridae, Dytiscidae, and Gyrinidae (Lawrence, Weir, and Pyke).
For taxa of supraspecific rank, arrangement is taxonomic according to the latest revision.
For species, arrangement within genera is alphabetical by specific epithet, and within species,
subspecies are arranged alphabetically by subspecific epithet.
Information presented for taxa is abundant. For each family, a brief account is given about
number of included species, composition, world distribution, fossil record, habits and life cycles,
as well as taxonomic history and present state of knowledge of the group within Australia.
Generic and specific synonymies are detailed for the Australian components. Data about
geographical distribution were compiled from published sources and from personal knowledge
of the authors, as appropriate. Habitat and life history data are also included.
Names of taxa in the text are set in boldface, and they stand out clearly. The text is
complemented by an extensive index of taxonomic names. The durable volume has hard covers,
with a covering of blue cloth.
This contribution is excellent, and is a credit to the authors, editors, and printers, and to the
Division of Entomology, CSIRO, under whose aegis the required bibliographical research was
carried out. Coleopterists will be looking forward to publication of the remaining parts of this
remarkable catalogue that deal with beetles.
George E. Ball
713
EDITOR’S ACKNOWLEDGEMENTS AND COMMENTS
Reviews of manuscripts were provided by the individuals whose names appear below. The
appearance of their names does not imply that the reviewers endorsed the papers published, but
only that they helped the Editor, as requested. Their assistance is very much appreciated.
J. S. Ashe
Department of Zoology, Field Museum of Natural History
S. A. Braimah
Edmonton, Alberta
Sir Cyril Clarke
Merseyside, England
Gregory R. Courtney
Department of Entomology, University of Alberta
R. W. Crosskey
Department of Entomology, British Museum (Nat. Hist.)
D. C. Currie
Department of Entomology, University of Alberta
H. Goulet
Biosystematics Research Institute, Agriculture Canada, Ottawa
S. A. Marshall
Department of Environmental Biology, University of Guelph
A. F. Newton
Department of Zoology, Field Museum of Natural History
J. R. Spence
Department of Entomology, University of Alberta
M. K. Thayer
Department of Zoology, Field Museum of Natural History
D. R. Whitehead
Systematic Entomology Laboratory, USDA, United States National
Museum
French translations of abstracts were provided by J. O. Lacoursiere, of my Department. J.
S. Scott and D. Shpeley read proof, as required, and assisted in other ways as well.
The Publications Manager, Mrs. S. Subbarao, provided excellent service both in production
and management, and thanks to her efforts, some important improvements to our operation will
be implemented in the near future.
The staff of Printing Services, University of Alberta, provided generally highly satisfactory
issues. This year, a special feature was the color plates in F. A. H. Sperling’s treatment of
Papilio machaon species group. It is a pleasure to acknowledge the careful work of E. Anaka
and J. Smith of Printing Services, in printing these plates.
New printing equipment, recently acquired by Printing Services, will provide higher
resolution of text and consequent improvement in readability. Eventually, we plan to take full
advantage of this improved facility to transfer sophisticated art work. The immediate
advantage to authors, of this new equipment, is that they may submit manuscripts on
Macintosh discs, using Microsoftword software. Authors who wish to submit manuscripts
Quaest. Ent., 1987, 23 (4)
714
generated on other microcomputers should consult the Publications Manager for information.
In an effort to improve the quality of paper, yet control the cost of production, page size will
be changed from 6.75 in X 9.75 in. to 6 in. X 9 in. beginning with Volume 24.
George E. Ball
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