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13
it
Quaestiones
Entomologicae
JVL
A periodical record of entomological investigations/
published at the Department of Entomology/
University of Alberta, Edmonton, Canada.
VOLUME 17
1981
CONTENTS
Lafontaine-Classification and phylogeny of the Euxoa detersa group (Lepidoptera:
Noctuidae) 1
Oliver-Redescription and systematic placement of Oreadomyia albertae Kevan and
Cutten-Ali-Khan (Diptera: Chironomidae 121
Corkum and Clifford-Function of caudal filaments and correlated structures in mayfly
nymphs, with special reference to Baetis (Ephemeroptera) 129
Scholefield, Pritchard and Enfield-The distribution of mosquito (Diptera, Culicidae) larvae in
Southern Alberta, 1976-1978 147
Note-Schaber-Recovery of Patasson luna (Hymenoptera: Mymaridae) a parasite of the alfalfa
weevil, Hypera postica (Coleoptera: Curculionidae), in Alberta 169
Book Review-Larsson, Sven Gisle. 1978. Baltic Amber - a Palaeobiological Study.
Entomograph, vol. 1 171
Book Review-Gillott, Cedric. 1980. Entomology 173
Hudson, J.E.-Studies on flight activity and control of mosquitoes in the Edmonton area,
1971-73 179
Fredeen, F.J.H.-Keys to the black flies (Simuliidae) of the Saskatchewan River in
Saskatchewan 189
Doane, J.F.-Seasonal captures and diversity of ground beetles (Carabidae: Coleoptera) in a
wheat field and its grassy borders in central Saskatchewan 211
Carter, A.-Aspects of the comparative ecology of populations of four Patrobus species
(Coleoptera: Carabidae: Patrobini) at George Lake, Alberta 235
Roughley and Pengelly-Classification, phylogeny, and zoogeography of Hydaticus leach
(Coleoptera: Dytiscidae) of North America 249
Book Review-Tschirnhaus, M von. 1981. Die Halm- und Minierfliegen im Grenzbereich
Land-Meer der Nordsee 3 1 5
Editor's Acknowledgements 317
Index 319
CORRIGENDA-Quaestiones Entomologicae, Volume 17(1,2)
Corkum and Clifford-Caudal filaments in Mayfly nymphs
page/line
131/18-19 Clifford (1970) and were ... therefore relatively small.”
should read, “... Clifford (1970) and were, therefore, relatively small.'
132/first entry in table “50°30’26” ” should read “50°30’47” ”
138/last line “(r = 0.019)” should read “(r = -0.019)”
141 /20 “ B . apinus” should read “/?. alpinus”
143/15 “/?. hageni” should read “/?. rhodani"
143/1 6 -1 7 “ Baetis , B. lapponicus, Baetis and B. rhodani ”
should read “5. alpinus, B. lapponicus, B. lutheri and B. rhodani ”
ANNOUNCEMENT
The Willi Hennig Society, whose purpose is the furtherance of scientific procedure in
systematics, will hold its third annual meeting at the University of Maryland at College Park
on November 19-22, 1982. The meeting is sponsored by the Maryland Center for Systematic
Entomology. There will be symposia as well as contributed papers. Further details may be
obtained from the organizers: C. Mitter, Entomolgy Department, University of Maryland,
College Park, MD 20742; V.A. Funk, Department of Botany, National Museum of Natural
History, Smithsonian Institution, Washington, D.C. 20560; and F.C. Thompson, Systematic
Entomology Lab, USDA, Smithsonian Institution, Washington, D.C. 20560.
Quaest. Ent. Vol. 17(3,4) mailed from Edmonton , Alberta 17.VI.19S2
CORRIGENDA-Quaestiones Entomologicae , Volume 17(3,4)
Roughley and Pengelly - Classification, Phylogeny, and Zoogeography of
Hydaticus Leach (Coleoptera: Dytiseidae) of North America.
page/line
249/35". . . (ancestral stock of H. modes tus) . " should read, "...(ancestral
stock of H. bimarginatus - H. rimosus) , and via Beringia
(H. modes tus) ."
251/26"BMNH British Museum (Natural History, London..." should read
"BMNH British Museum (Natural History), London..."
2 53/ 3 8" Roman numberals. . . " should read "Roman numerals..."
257/31"... Beauline, 1917:124..." should read "...Beaulne, 1917 :124. . .
259/21". ..(Fig. 12) l... H. modestus Sharp." should read "...(Fig 12)...
H. modestus Sharp * . "
262/4"... <?1 12.8 7.3 should read "... <f . . . 1 ... 12.8 ... -
... 7.3 ...
269/31,32"... aedoeagus long x * 3.3 mm..." should read "aedoeagus long
(x * 3.3 n*n..."
271/8”ll"Dytiscus bimarginatus Say, 1831:5. — Say, 1834:442. Hydaticus
bimarginatus, LeConte, 1869:556. — Crotch, 1873:404 - Sharp,
1382:654. — Blatchley, 1910:233. — Leng and Mutchler, 1918:89 —
Blatchley, 1919:314. — Young, 1954:113. — Matta and Michael,
1977:48. Hydaticus fulvicollls Aub4, 1838:184. Hydaticus rimosus?
Young, 1954:113; nee Aub4, 1838:182. "should read
"Dytiseus bimarginatus Say, 1831:5. — Say 1834:442.
Hydaticus bimarginatus, LeConte, 1869:556. — Crotch, 1873:404
~ Sharp, 1882 ;654. - — Blatchley, 1910:233. ~ Leng and Mutchler,
1918:89. — Blatchley, 1919:314. — Young, 1954:113. — Matta
and Michael, 1977:48.
Hydaticus fulvicollis Aub4, 1838:184
Hydaticus rimosus? Young, 1954:113; nec Aub4, 1838:182.
271/13". . . (Le Conte, 1869a, p. VI)." should read "...(LeConte, 1869a,
p. VI)."
271/19". . .H. rimosus. . should read "... H. fulvicollis. .
273/22"min. - 8, N 1 150)." should read "min. - 8, N - 150).
274/34"sub-lateal. . . " should read "sub-lateral..."
281/ 36 "He continentalis J. Balf our-Brovne. . should read "H (H.) continentalis
J. Balfour-Browne. . . "
286/27". .. two taxa was probably been..." should read "...two taxa probably
has been. . ."
289/10"3especies. . should read "d ' especes. . . "
299/2 Add: Elytral coloration characterized as vittate (V, Fig. 9),
Fasciate (F, Fig. 8), and non-fasciate (F, Fig. 7); elytral sculpture
characterized as rugose (R, Figs. 9 and 17) and smooth (S, Figs. 7
and 8). Based on Table 2, p. 264.
292/ "Ross, H.H. 1874..." should read "Ross, H.H. 1974..."
Quaestionej-
Entomologicae
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada,
VOLUME 17
NUMBER 1-2
JANUARY-APR1L 1981
CLASSIFICATION AND PHYLOGENY OF THE EUXOA DETERSA GROUP
(LEPIDOPTERA: NOCTUIDAE)1
J. D. Lafontaine
Biosystematics Research Institute
Agriculture Canada
Ottawa K1A 0C6 Quaestiones Entomologicae
17:1-1201981
ABSTRACT
Thirty-one species and two subspecies are included in the detersa group of the genus Euxoa
(Lepidoptera: Noctuidae). Two species are described as new: Euxoa castanea Lafontaine, new
species, type locality - Golden, British Columbia ; Euxoa inscripta Lafontaine, new species,
type locality - Craig, Colorado. All 31 species in the E. detersa group are distributed in
western North America; the ranges of four of these extend into eastern North America.
In North America, Euxoa includes about 180 species in 65 species groups and 7 subgenera
( two unnamed). In the Palearctic region the genus includes about 100 species in 4 subgenera
and about 20 species groups. Few species are distributed south of northern Mexico, northern
Africa and northern India. A North American origin for the genus in mid-Tertiary time is
most probable, based on specializations of the genus for aridland living.
A phytogeny of the E. detersa group is preceded by a reconstructed phytogeny of species
groups and subgenera of Euxoa and an evaluation of structural characters.
Sommaire
Dans le group detersa du genre Euxoa nous incluons 31 especes et deux sous-especes. Nous decrivons deux nouvelles
especes: Euxoa castanea Lafontaine, n. esp., localite ty pique - Golden, Colombie britanique; Euxoa inscripta Lafontaine,
n. esp., localite typique Craig, Colorado. Les 31 especes se recontrent tous dans I ’ouest e I’Amerique du nord. Quatres de
ces especes s’ettendent dans Vest du continent.
En Amerique du nord on y rencontrent 180 especes d’ Euxoa reparties en 65 groupes d’especes et sept sous-genres
(deux ne sont pas decrits). En Eurasie il y a environs 100 espe reparties en 20 groupes d’especes et quatre sous-genres.
Peu d’especes atteignent le nord <Ju Mexique, de I’Afrique et de I’Inde. Nous postulons pour ce genre une origine nord
americaine lors du Tertiaire moyen. Cette hypothese est basee sur le mode de vie adapte aux region aride chez ces
insectes.
La phylogehse du group detersa suit celles des groupes et sous-genres des Euxoa a la suite de revaluation des
characteres structuraux.
'Based on a doctoral thesis submitted at the University of Alberta, Edmonton, Canada
2
Lafontaine
TABLE OF CONTENTS
INTRODUCTION
The genus Euxoa has a notorious reputation for the difficulty encountered in identifying
constituent species. This notoriety stems from three factors: the large number of species,
structural uniformity among species, and high degree of intraspecific variability.
In this paper, a revision of the Euxoa detersa group, the largest species group in the genus,
is presented. Thirty-one species and two subspecies are included in the group. Classification and
phylogeny of the E. detersa group are treated in detail. More generalized discussions on
phylogeny, distribution, origin, and bionomics of the genus are included.
A section on the biogeography of the Euxoa detersa group, originally included in this
revision, will be included in a paper on the biogeography of the genus Euxoa which is in
preparation.
Historical review
A review of taxonomic work on Euxoa in North America was given by Hardwick (1970a).
In the latter half of the nineteenth century, workers in both Europe and North America
described many new species of noctuids. Many new species were named on the basis of
peculiarities in color or maculation. The genus Euxoa was particularly susceptible to a
proliferation of names because of polymorphism and geographical variation in wing color in
many species.
In the twentieth century, use of male genitalia as a taxonomic tool was an important
advance that allowed many taxonomic problems to be resolved and correct synonymies to be
established in many genera. In Euxoa , however, structural uniformity, coupled with the large
number of species and available names kept the genus in a state of confusion. The fact that
Euxoa could not be subdivided in smaller, more manageable monophyletic units seriously
hampered revisionary work on the genus.
Revision of Euxoa of the Soviet Union by Kozhanshikov (1929, 1937) was a significant step
forward in that both male and female genitalia were used. In North America it was not until
1950 that female genitalia were used in revisionary work on Euxoa. At that time McDunnough
(1950) emphasized characters of the female genitalia in a revision of eastern North American
The Euxoa detersa group (Lepidoptera: Noctuidae)
3
species.
A breakthrough in Euxoa systematics came when Hardwick (1970a) demonstrated the
value of the male vesica showing that characters of this structure could be used to arrange
species into subgenera and species groups as well as to resolve taxonomic problems among
closely related species. Subdivision of Euxoa into subgenera and species groups initiated a
series of studies on individual species groups (see papers by Hardwick and Lafontaine).
Distribution of the genus
In general, Euxoa is distributed in dry regions of the northern hemisphere. The genus is
most diverse in North America, particularly in the west; it includes about 180 species in 65
species groups and 7 subgenera (see Lafontaine, 1982 and Phylogeny below).
In the Old World, about 70 species occur in the Soviet Union, Mongolia and China. To this
total, additional species can be added from Kashmir, India, and Nepal (eight species), the
Middle East (four species), Europe (10 species) and northern Africa (10 species) (Boursin,
1940a, 1940b, 1944, 1952, 1964; Hartig and Heinicke, 1973; Kovaks and Varga, 1973;
Kozhantschikov, 1929, 1937).
The Palearctic fauna, consisting of approximately 100 species, can be arranged in about 20
species groups (most of which also occur in North America) and three subgenera (all of which
also occur in North America). Few species of Euxoa occur south of northern Africa, northern
India and northern Mexico.
Although about 24 species of Euxoa were formerly thought to occur in South America (e.g.
Kohler, 1945; Kozhantshikov, 1937), all but two of these have now been transferred to other
genera (see Kohler, 1955, 1967). The status of the two South American Euxoa species remains
unclear.
Origin of the genus Euxoa
In the absence of a fossil record for Noctuidae (see Wilson, 1978a; Skalski, 1976) other
evidence must be considered to establish a possible time of origin of Euxoa.
The Lepidoptera are generally believed to have arisen some time between the Triassic and
the Early Cretaceous (Shields, 1976; Whalley, 1978). The oldest known Lepidoptera fossils are
Lower Cretaceous and consist of a species of Micropterygidae and a probable species of
Incurvariidae (Whalley, 1978). Since most families of Lepidoptera were probably present by
the Oligocene (Shields, 1976; Common, 1975; Wilson, 1978a, 1978b), radiation of the order
must have occurred during Late Mesozoic or Early Tertiary times, even though most known
Late Cretaceous fossil Lepidoptera specimens belong to primitive groups (Wilson, 1978b;
Whalley, 1978). Speculation, based on distribution patterns, indicates an earlier radiation of
families (see Shields, 1976). Later arguments by Shields and Dvorak (1979), whish expanded
on this theme, are weak because of a lack of a substantiating phylogenetic analysis.
The “higher noctuids”, particularly cutworm genera adapted to desert and grassland
conditions, probably originated, or at least radiated, during the period when such habitats were
developing in the world. Desert and grassland habitats began to develop in late Oligocene and
Miocene times (Cracraft, 1973; Axelrod, 1976). Desert flora and aridland avifauna are
believed to have evolved during the Miocene (Hubbard, 1973) as did such grassland
mammalian families as Bovidae (cattle, antelope, goats) and Antilocapridae (pronghorn)
(Romer, 1966). The Miocene is also the period of origin of grazing horses (Simpson, 1951) and
aridland katydids (Cohn, 1965). Although grass pollen is first recorded from Upper Cretaceous
Quaest. Ent., 1981, 17 (1,2)
4
Lafontaine
(Maastrichtian) deposits in West Africa “grasses do not become frequent in the fossil record
until the Lower Eocene, probably correlated with the rise of grazing mammals and the origin of
non-bambusoid tribes of grasses” (Raven and Axelrod, 1974, p. 395).
Based on this information, aridland cutworm genera, including Euxoa are assumed to have
arisen during the middle Tertiary.
Statements about place of origin of Euxoa are tentative until a thorough phylogenetic
analysis of Palearctic species has been undertaken. A preliminary analysis of the phylogenetic
relationships of the Nearctic and Palearctic species leads me to suggest that the genus
originated in North America. Most species groups that occur in the Old World belong to the
highly derived lineages of the subgenus Euxoa (i.e. lineages 11 and 13, Fig. 186) and to the
subgenus Orosagrotis. The subgenus Chorizagrotis also occurs in the Palearctic region. This,
however, is not surprising since the subgenus contains migratory species.. One species of the
subgenus in North America occurs as far north as Greenland and Alaska (Hardwick, 1970a).
Origin of the Euxoa detersa group
Other than general arguments outlined above for a mid-Tertiary time of origin for Euxoa ,
little can be said regarding time of origin of the Euxoa detersa group, other than that it was
probably in Neogene time.
The sister group of the E. detersa group (see figure 186) is not helpful for determining a
place of origin of the E. detersa group (other than that it was somewhere in the Holarctic
region), since it consists of species groups that occur in both the Nearctic and the Palearctic
regions.
The E. detersa group, as well as about 50 species groups and three subgenera of Euxoa are
apparently restricted to the Nearctic region. The diversity of the Palearctic Euxoa fauna, as
compared to that of the Nearctic region, suggests that either relatively few New World species
groups reached the Old World, or many species groups have gone extinct there. The former
hypothesis is considered more likely because the Palearctic Euxoa fauna is primarily a
subtraction fauna of Nearctic groups consisting mainly of more northerly Holarctic species
groups.
Based on these data, a North American origin of the E. detersa group is considered to be the
most probable hypothesis.
Euxoa Survey
Long series of specimens from a large number of localities are required for revisionary work
on a taxonomically difficult genus. Such material is also invaluable for clarifying distribution
patterns and in determining habitat preferences for species. To these ends, a survey of Euxoa
species in western North America was initiated in 1960 by D.F. Hardwick of the
Biosystematics Research Institute, Ottawa. Approximately 50,000 specimens have been
collected to date. At each collecting site, detailed habitat notes and photographs were taken;
these were used to establish habitat preferences and tolerances of Euxoa species. These
specimens when added to the holdings of the Canadian National Collection (CNC) form a
sample or approximately 80,000 specimens of Euxoa. About 12,000 of these specimens are
species in the Euxoa detersa group. Most collections were made during August and September,
the period in which the majority of Euxoa species are flying. Distribution of spring flying
species remain inadequately known.
The Euxoa detersa group (Lepidoptera: Noctuidae)
5
Cutworm bionomics
Larvae of Euxoa species, as well as those of many other noctuid genera, are commonly
called cutworms. Cutworm ecology is a field of study that has received little attention. This is
surprising considering both total number of species involved and economic importance of many
species. Problems in identification is probably the main factor that has hampered ecological
work on cutworms. Only one book (i.e. Crumb, 1956) attempts to treat all North American
noctuid larvae; this work includes about 700 species, approximately one-fifth of the North
American fauna. Unfortunately, Crumb could find no characters for identifying to species
larvae of Euxoa.
Most papers on cutworm bionomics treat only a single, usually economically important
species, and contain only descriptions of larvae and life history information. A notable
exception to this is a series of papers about bionomics of the pale western cutworm (Agrotis
orthogonia Morr.) by H.L. Seamans, W.C. Cook, and L.A. Jacobson (see Jacobson, 1971, for
review). Unfortunately, comparable information for other cutworm species is lacking, although
there is a fair amount of piecemeal ecological information about some pest species.
Most cutworms are subterranean, surface-feeding, or climbing. Larvae of most species of
Euxoa are surface-feeding cutworms, although those of a number of species readily adopt a
climbing habit (Hardwick, 1970a).
Many factors of the physical environment are important to cutworm bionomics. Probably
the three most important factors are temperature, soil, moisture, and soil texture. Temperature
is important in controlling rates of metabolism and development, limits of distribution both
northward and southward, and number of generations per year. Many cutworms, including
species of Euxoa , aestivate through summer heat in earthen cells in the soil. Since there is a
limit to the amount of heat that they can tolerate, their southern limit is probably controlled by
summer temperature (Hardwick and Lefkovitch, 1971). In general, larvae of species that occur
in open aridlands have a longer prepupal aestivation than do those of species, that occur in
forested areas (J.R. Byers, Pers. comm.). Soil moisture and soil texture tolerances and
preferences of cutworm larvae are known for few species but these factors are undoubtedly
important in determining distribution patterns and in resource partitioning. The importance of
soil conditions to plant distributions has recently been recognized (e.g. see Kruckeberg, 1969;
Zamora and Tueller, 1973).
Ranges of many Euxoa species are similar to those of plant species and communities (e.g.
species associated with sagebrush areas or conifer forests). Since larvae do not eat these plants,
it is assumed that the association is due to similarities in habitat requirements rather than to a
direct dependence of the noctuid on the plant. A possible exception to this is that many aridland
Euxoa species occur in areas where rabbitbrush (Chrysothamnus spp.) is abundant. Although
this association could be attributed to a preference for arid habitats in both cutworms and
rabbitbrush, it is probably also related to feeding requirements of adults. In many arid regions,
rabbitbrush is one of few plants in bloom in the fall when adults of Euxoa species are flying; its
flowers are an essential food source for the moths. Flight of fall noctuids in many arid regions is
markedly correlated with flowering time of rabbitbrush (Cook, 1930b).
The most interesting aspect of cutworm bionomics, and the one that has received the least
attention, is resource partitioning by larvae. Although larvae of most species are general
feeders, they show preference for either broad-leaved herbs or grasses (e.g. see Strickland,
1948). Except for a few basic factors in niche segregation (e.g. subterranean vs.
surface-feeding; broad-leaf eating vs. grass-eating; and moisture loving vs. dry soil species)
Quaest. Ent., 1981, 17 (1,2)
6
Lafontaine
knowledge of cutworm bionomics is in a very crude state. Walkden (1943) found that fields
that differ in plant composition also differ in cutworm composition. Unfortunately, soil
conditions were not considered as a possible factor affecting species composition.
Experiments on effects of such factors as temperature, moisture and soil conditions on
cutworm bionomics are needed. Understanding of effects of these variables will greatly enhance
knowledge of resource partitioning and geographical distribution of cutworm species.
MATERIALS AND METHODS
Materials
During this study, approximately 16,000 specimens were examined. Most specimens (about
12,000) are in the Canadian National Collection (CNC), Ottawa. This material was
supplemented by specimens from the private collections of A. Blanchard, Houston, Texas, J.G.
Franclemont, Ithaca, New York, J.R. Heitzman, Independence, Missouri, M.C. Nielson,
Lansing, Michigan, R.W. Poole, United States Department of Agriculture, Washington, D.C.
and from collections in the following institutions:
The American Museum of Natural History (AMNH), New York, New York (F.H.
Rindge).
The California Academy of Sciences (CASC), San Francisco, California (D.H.
Kavanaugh).
California Department of Agriculture (CDAE), Sacramento, California (T.N. Seeno).
Cornell University (CU), Ithaca, New York (J.G. Franclemont).
North Dakota State University (NDSU), Fargo, North Dakota (E.U. Balsbaugh Jr.).
Los Angeles County Museum of Natural History (LACM), Los Angeles, California (C.L.
Hogue).
San Diego Natural History Museum (SDMC), San Diego,- California.
The United States National Museum of Natural History (USNM), Washington, District
of Columbia (E.L. Todd, R.W. Poole).
Peabody Museum of Natural History (PMNH), Yale University, New Haven,
Connecticut (C.L. Remington).
Methods
Collecting and preparation of material. - Adult moths were collected by light trapping with
a 125-watt, Osram, mercury vapor bulb as light source. Design of the light trap used was
described by Hardwick (1968).
Specimens were pinned in the field and later spread following the technique described by
Martin (1977).
Genitalia preparations were made following the technique described by Hardwick (1950)
with one modification. The male vesica was everted by squirting isopropyl alcohol through it
while immersed in a dish of isopropyl alcohol rather than while immersed in a 10 per cent
alcohol solution.
Ovipositor valves of some females were separated from the remainder of the genitalia and
mounted flat. This was done by cutting the membrane anterior to the valves and between the
The Euxoa detersa group (Lepidoptera: Noctuidae)
7
valves ventrally. The spread out valves were then mounted. This allowed valve shape and setae
to be studied more readily.
Structural characters. - There is a frustrating paucity of structural characters that can be
used to characterize adults of species of Euxoa other than those of the genitalia.
Males of most species have biserrate antennae, although those of a few species groups can be
characterized by antenna structure. Males of three species groups have plumose antennae;
those of two species groups have filiform antennae.
A frontal tubercle is present in adults of all but five species of Euxoa. Species that are
without a frontal tubercle live in sandy areas; adults of these species presumably do not have
difficulty in escaping from the subterranean pupal cell after eclosion.
Adults of most species have rounded, globular eyes. Those of some species that live in arctic
or alpine areas have reduced, elliptoid eyes.
Of the above three external structural characters, only one is of use in distinguishing species
in the Euxoa detersa group; a frontal tubercle is absent from most specimens of Euxoa detersa
(Walker).
Male genitalia possess many structural characters of taxonomic and phylogenetic
importance. Within Euxoa , size and shape of sacculus, sacculus extensions, cucullus, harpe,
juxta and uncus are all important characters. Of these structures, sacculus extensions and
harpes are most useful for distinguishing species of the E. detersa group, although other
structures may be important in specific instances. The most useful structure of the male
genitalia, both within the E. detersa group and Euxoa in general, is the vesica. Overall vesica
shape, as well as shape and position of accessory pouches, provide excellent characters for
taxonomic and phylogenetic studies.
Female genitalia have several very useful structural characters. Many closely related species
that are indistinguishable by structures of male genitalia can be distinguished by those of
female genitalia and vice versa. The most useful character for phylogenetic studies and for
distinguishing species groups is shape of the bursa copulatrix. Shape of ovipositor valves, their
vestiture, and presence or absence of sclerotized projections, are useful characters for
distinguishing species. In females of some species groups sclerotized plates in the dorsal and
ventral wall of ductus bursae, and anterior apophyses, may be used to distinguish species.
Taxonomic value of structural characters of Euxoa were reviewed by McDunnough (1950)
and Hardwick (1970a).
Measurements. - Genitalia measurements were made with a Leitz stereoscopic microscope
at 12.5X magnification using a micrometer eyepiece with a scale interval of 0.025 mm.
Length of four structures of male genitalia were measured: right sacculus extension, left
sacculus extension, right harpe, and right sacculus. From these data, mean and standard
deviation were calculated. A diagrammatic presentation of how these measurements are taken
was given by Hardwick and Lefkovitch (1973).
In addition to genitalia measurements, two wing measurements were taken. “Expanse” is
the distance between apices of the forewings of spread specimens. This distance varies slightly,
depending on how a specimen is spread, so not all specimens were included. “Length of
forewing” is length of right forewing from base to apex.
Maculation. - Maculation of Noctuidae was described in detail by Forbes (1954); only a
brief review is included here.
Specimens with maculation complete (fig. 47) have six transverse lines crossing the
forewing; basal line, transverse anterior line, median line, transverse posterior line, subterminal
Quaest. Ent., 1981, 17 (1,2)
8
Lafontaine
line, and terminal line. The basal line, near the wing base, is extended only half way across the
wing in most species. Transverse anterior and transverse posterior lines are double lines; they
are present and conspicuous in most species. The median line consists of a dark shade that
crosses the forewing between the reniform spot and orbicular spot close to the transverse
posterior line. In most species, the subterminal line is a pale line made evident by dark shading
in the terminal area and by dark sagittate spots proximal to it in the subterminal area. The
terminal line is along the outer edge of the forewing.
The area of the forewing is divided into four smaller areas. The basal area is between the
wing base and transverse anterior line; the median area is between transverse anterior and
transverse posterior lines; the subterminal area is between the transverse posterior line and
subterminal line; and the terminal area is between subterminal and terminal lines. The basal
area has a black, basal dash in many species (fig. 9). The median area has three spots in most
specimens (fig. 47), a rounded “orbicular spot”, a kidney-shaped “reniform spot”, and an
elongate “claviform spot” distal to the transverse anterior line. The terminal area is dark in
most species.
The cubital vein (Cu) of the forewing is just below the orbicular and reniform spots; it
branches into four veins, M2, M3, Cu2 and Cu2. In many species, veins Cu, Cux and M3 are
pale-lined and contrast with the ground color of the forewing (fig. 51).
The hind wing has a dark, crescentic, discal spot in many species. In some species, a trace of
a median line is present on the dorsal surface; this line is parallel to the outer edge of the wing
about 2/3 from wing base (e.g. see fig. 13). A median line on the ventral surface of the hind
wing is present in most species.
Description format. - The description format used follows that of Hardwick (1970a), and
includes: synonymy and selected bibliography; description of adult; type material; distribution
and period of flight; habitat; and remarks. A list of material examined is not included but
collection data are summarized in the discussion on distribution and period of flight.
Descriptions of adults are presented in the following sequence: vestiture of head and thorax;
ground color of forewing; transverse lines; areas of forewing; spots, dashes and streaks on
forewing; expanse; male genitalia; female genitalia.
Structures of male genitalia referred to most frequently in descriptions are shown in figure
65.
Habitat descriptions. - Habitat descriptions were prepared by using both field notes and
photographs from each collecting site. At any one locality, such material is of little use since it
can not be determined which portion of the collecting area a given specimen came from. Over a
large number of localities, however, a picture begins to emerge as to what conditions must be
present for each species to occur in an area. Some species for example, which were frequently
collected in pine forests in montane areas were collected in the Great Plains only at localities
where a pine forest was nearby. These species are assumed to occur only in pine forests and not
in treeless plains. Other species occur only where coarse, loose, soil is present and are found in
the Great Plains only in areas of heavy erosion such as badlands and river terraces.
Although habitat diagnoses obtained in this way are crude, they have proven to be useful in
interpreting distribution patterns.
A more refined knowledge of habitat preferences for each species must await field studies of
larval habitats.
Sorting of specimens. - Specimens were initially sorted into more or less homogeneous
populations by using wing color and maculation, geographical distribution, and period of flight.
The Euxoa deter sa group (Lepidoptera: Noctuidae)
9
These population samples were then studied more critically by using structural characters and
were arranged into more inclusive units. It was then determined whether these groupings
represented species, subspecies, or geographical populations.
Male and female specimens of each species are similar in wing pattern so no difficulty was
encountered in associating both sexes of each taxon.
After limits of variation of each species had been established, specimens of each were
arranged geographically to study geographical variation.
To facilitate taxonomic treatment and discussion, species were arranged into species groups.
Species groups are monophyletic groups of structurally similar species that share a derived
character state that distinguishes its members from those of other species groups. The rank of
species group is informal. Taxa belonging to the Euxoa detersa group are revised in this paper;
other species groups are discussed briefly in relation to the phylogeny of species groups of
Euxoa.
Criteria for species and subspecies. - “Species are groups of actually or potentially
interbreeding natural populations, which are reproductively isolated from other such groups”
(Simpson, 1961, p. 150).
In practice, presence or absence of reproductive isolation can rarely be demonstrated,
especially when preserved material is all that is available. Also, such a rigid definition is
frequently inadequate for species in which reproductive isolation occasionally breaks down
(Byers and Hinks, 1978; Teal et al., 1978). In these instances, Simpson’s evolutionary
definition of a species is more appropriate. “An evolutionary species is a linege ... evolving
separately from others and with its unitary evolutionary role and tendencies44 (Simpson, 1961,
p. 153).
“A subspecies is an aggregate of phenotypically similar populations of a species, inhabiting
a geographic subdivision of the range of a species and differing taxonomically from other
populations of the species” (Mayr, 1969, p. 41). There has been much discussion among
taxonomists as to what constitutes a subspecies (see discussion by Mayr, 1969). Some
taxnomists recognise subspecies only for allopatric pouplations of uncertain status (e.g.
Whitehead, 1972); others, at the opposite end of the spectrum, consider every population that
differs in any way from neighboring populations to be a subspecies. While a proliferation of
names for local populations is undesirable for many reasons, the subspecies taxon is useful if
applied judiciously.
I use two criteria for deciding whether or not a subspecific name is warranted. The degree of
phenotypic difference between subspecies should approach that found between species of the
same or related groups; the phenotypic differences are the result of isolation and differentiation,
not simply the effect of local conditions on phenotypic expression of the gene pool.
For example, specimens of Euxoa detersa detersa and Euxoa detersa personata appear to
be as different from each other as are those of Euxoa detersa from those of Euxoa cicatricosa.
The differences between these two aggregates of populations are found over a variety of
comparable habitats. The two subspecies were probably isolated in different geographical
regions during the last glacial maximum. Since Euxoa detersa personata occurs in the Great
Lakes region, the Great Plains, and the boreal forest, a vernacular name such as the “Great
Plains” populations is riot convenient; further, the subspecific name draws attention to the
distinctness of the two aggregates of pouplations.
On the other hand, specimens of Euxoa cicatricosa look different in various parts of the
range of the species. The differences, however, are in wing color which matches the color of the
Quaest. Ent., 1981, 17 (1,2)
10
Lafontaine
substrate where the moths occur. Subspecific names for such populations cannot be justified.
Inbreeding of some color forms of Euxoa species has produced forms typical of specimens from
other regions (J.R. Byers, pers. comm). These experiments indicate that habitat associated
color forms are the effect of local conditions on phenotypic expression of the gene pool; such
ecotypic forms could change rapidly in different habitats.
Geographical variation. - An excellent review of this topic was given by Hubbell (1956). In
adults of Euxoa , geographical variation in wing color is a common phenomenon in many
species. This has two components in most species: color or hue and shade or degree of darkness.
Wing color varies with substrate color in many species: brown forms occur in the east and
north; buff, or pale gray forms occur in the Great Plains and northern Great Basin; and reddish
orange forms occur in red sandstone areas of the southwest. In shade, specimens tend to be pale
in hot, arid regions and dark in cool, mesic regions.
Some geographical variation cannot be explained in terms of local conditions but is the
result of isolation and differentiation of populations. In many species geographical variation of
this type consists of differences in wing pattern rather than color.
Geographical variation in structural characters is rare in Euxoa species. In some species
length of sacculus extension in male genitalia varies in different parts of the species range (e.g.
Euxoa punctigera (Wlk.), see Lafontaine, 1974c). Within the Euxoa detersa group,
geographical variation in structural characters has only been found in shape of female
ovipositor valves of three species (E. castanea Laf., n. sp., E. idahoensis (Grt.), E.
quadridentata (Grt. & Rob.) it is discussed under those species.
Immature stages. - Immature stages were not used in taxonomic or phylogenetic analyses.
The reason for this is that immature stages are known for too few species, and those that are
known lack characters suitable for such analyses.
Eggs of two species in the Euxoa detersa group (E. servita (Sm.), E. detersa (Wlk.)) have
been described (Salkeld, 1975).
Larvae of only one species in the Euxoa detersa group (E. detersa (Wlk.)) have been
described, although those of four other species (E. castanea Laf., n. sp., E. auripennis Laf., E.
niveilinea (Grt.), E. cicatricosa (Grt. & Rob.)) have been reared (Hinks and Byers, 1976).
At present, no characters have been found with which larvae of Euxoa species can be
differentiated (Crumb, 1956).
CLASSIFICATION
The Euxoa detersa group
Description. — Antenna of male varied from slightly to moderately biserrate and bifasciculate. Frontal tubercle
present except in most specimens of E. detersa.
Maculation of forewing complete in species that lack longitudinal streaking; transverse lines obscure or absent in
species with longitudinal streaks on forewing.
Male genitalia. Valves bilaterally symmetrical or slightly asymmetrical with respect to lengths of sacculus extensions;
left sacculus extension slightly shorter than right in many specimens. Right sacculus extension slightly to markedly longer
than right harpe in most species; markedly shorter in one species (E. melura). Sacculus extensions about as stout as harpes
to slightly stouter. Vesica bent to right or dorsally above apex of aedeagus. Sub-basal diverticulum foot-shaped in most
species; somewhat triangular in some species (e.g. E. costata), small and horn-like in males of E. citricolor. An additional
nipple-like diverticulum on vesica sub-basally in some species. Median diverticulum horn-shaped, basal bulge directed
toward sub-basal bend present in some species; median diverticulum near sub-basal bend in most species but positioned
about half way to apex in some species. Apical diverticulum in most species. Angle formed by apical half of aedeagus with
The Euxoa detersa group (Lepidoptera: Noctuidae)
11
respect to apical half of vesica 20-85°; 40-50° in most species.
Female genitalia. Bursa copulatrix unisaccate; corpus bursae oval or somewhat rectangular, slightly invaginated on
left side in females of some species; ductus bursae connected to corpus bursae 0.25 to 0.50 way along right side.
Sclerotized plates in dorsal and ventral walls of ductus bursae extended anteriorly slightly farther than anterior
apophyses, except in females of E. quadridentata. Ovipositor valve clothed with fine setae; long stout setae, sclerotized
apical projections, or short conical setae in some species. Sub-basal row of long, thin setae in most species. Ovipositor
valves rounded or triangular in shape; dorsal margins not fused.
Remarks. - The E. detersa group includes 31 species, all confined to North America:
Euxoa cost at a (Grote), E. castanea Lafontaine, new species , E. foeminalis (Smith), E.
idahoensis (Grote), , E. clausa McDunnough, E. brevipennis (Smith) , E. servita (Smith) , E.
redimicula (Morrison) , E. auripennis Lafontaine, E. arizonensis Lafontaine, , E. olivalis
(Grote) , E. agema (Strecker), E. oblongistigma (Smith) , E. citricolor (Grote), E. tronella
(Smith), E. teleboa (Smith) E. moerens (Grote), E: latro (Barnes and Benjamin) , E.
murdocki (Smith), E. dodi McDunnough , E. infracta (Morrison), E. laetificans (Smith), E.
quadridentata (Grote and Robinson), E. inscripta Lafontaine, new species , E. unica
McDunnough, E. niveilinea (Grote), E. dargo (Strecker), E. melura McDunnough, E. detersa
(Walker), E. cicatricosa (Grote and Robinson), E. recula (Harvey). Two species, E. detersa
and E. quadridentata include two subspecies: E. d. detersa (Walker), E. d. personata
(Morrison) ; E. q. quadridentata (Grote and Robinson), E. q.flutea Smith.
Adults of the E. detersa group are distinguished from those of other species groups by a
combination of male and female genitalic characters. These are: elongate sacculus extensions
and vesica shape in males; unisaccate, oval corpus bursae with ductus bursae on right side, and
ovipositor valves not fused together in females.
All members of the E. detersa group live in western North America; ranges of four species
extend into eastern North America as well.
Key to males of Euxoa detersa group
1 Right harpe at least 1.25 times length of right sacculus extension (Fig. 92) . . .
E. melura McD. (p. 42).
1 ' Right harpe less than 1.25 times length of right sacculus extension . . . 2
2 (L) Apical half of harpe microscopically pubescent, in addition to much larger
scattered setae .3
2 ' Apical half of harpe without pubescence, only larger scattered setae present,
these about half as long as diameter of harpe. . 8
3 (2) Orbicular spot elongate, bar-like, two or three times as long as wide; right harpe
less thant 1.2 mm long ... 4
y Orbicular spot rounded or oval; in some specimens elongate and fused with pale
costa, and right harpe more than 1.3 mm long 6
4 (3) Right sacculus extension about 1.50 times length of right harpe (Fig. 75)
E. olivalis (Grt.) (p. 27).
4 ' Right sacculus extension about 1.25 times length of right harpe (Fig. 76) . . . 5
5 (4') Forewing with costa pale, evenly colored, pale shading extended uninterrupted
to cubital vein on basal quarter of wing (Figs. 31. 32)
E. oblongistigma (Sm.) (p. 28).
5 ' Forewing with costa brown like remainder of wing, not evenly colored; costal
shading not extended to cubital vein near base of wing but separated from it by
several light and dark lines (Figs. 28, 29)
E. agema (Stkr.) (p. 28)
Quaest. Ent., 1981, 17 (1,2)
12
Lafontaine
6 (30 Forewing with black basal dash (Figs. 49, 50)
E. laetificans (Sm.) (p. 36).
6 7 Forewing without black basal dash . 7
7 (60 Right sacculus extension more than 1.50 times length of right harpe; forewing
dark brown with copper colored basal and subterminal areas (Fig. 48)
E. infracta (Morr.) (p. 35).
7 7 Right sacculus extension less than 1.50 times length of right harpe; forewing
light brown or gray (Fig. 47)
E.dodi Mc.D. (p. 34).
8 (20 Clavus prominent, sclerotized, much longer than wide; dorsal margin of
sacculus invaginated at base of clavus (Fig. 93) ... 9
8 Clavus inconspicuous and at most only slightly sclerotized; dorsal margin of
sacculus not invaginated at base of clavus . 12
9 (8) Forewing without veins Cu\ and M< pale-lined and projected into terminal area
(Fig. 61); distributed from Great Plains eastward to Great Lakes region
E. detersa personata (Morr.) (p. 45).
9' Forewing with veins Cu\ and M< pale-lined and projected into terminal area;
distributed either in eastern North America, or from western Great Plains
westward . . 10
10 (9') Distributed in eastern North America
E. d. detersa (Wlk.) (p. 45).
10 7 Distributed from western Great Plains westward .11
1 1 (100 Pale areas of forewing white, cream-colored, or pale orange-brown (Figs. 62,
63); distributed throughout intermontane region and western Great Plains (Fig.
184)
E. cicatriocosa (Grt. & Rob.) (p. 45).
11 ' Pale areas of forewing yellow (Fig. 64); distributed in southern and western
portions of intermontane region (Fig. 185)
E. recula (Harv.) (p. 47).
12 (80 Forewing without basal dash, not longitudinally streaked .. 13
12 7 Forewing either with black basal dash, or longitudinally streaked, particularly
on veins Cu, Cu\ and M< or both .. 18
13 (12) Right sacculus extension about as long as, or shorter than, right harpe (Fig. 80)
E. teleboa (Sm.) (p. 31).
13 7 Right sacculus extension longer than right harpe .. 14
14 (137) Basal and subterminal areas of forewing orange or yellow, median area gray
(Fig. 46)
E. murdocki (Sm.) (p. 34).
147 Basal, subterminal and median areas of forewing essentially same color .. 15
15 (147) Uncus slightly constricted at about 0.25 from apex; right harpe with 10-15 setae
(Fig. 79); hind wing white in most specimens (Figs. 33-38) . 16
15 7 Uncus dilated at middle, then tapered to apex; right harpe with 30-60 setae
(Fig. 82); hind wing brown (Figs. 41-44) . . 17
16 (15) Forewing yellow
E. citricolor (Grt.) (p. 29).
The Euxoa detersa group (Lepidoptera: Noctuidae)
13
16 '
17 (15')
17'
18 (120
18 '
19 (18)
W
20 (190
20'
21 (200
21 '
Forewing white or cream-colored with black dusting
E. tronella (Sim) (p. 30).
Vesica with small nipple-like diverticulum sub-basally, in adition to normal
foot-like sub-basal diverticulum; right harpe with about 30 setae; hind wing
gray-brown, paler toward base in most specimens; distributed in Great Plains
and Great Basin (Fig. 172)
E. moerens (Grt.) (p. 32).
Vesica with only normal foot-like diverticulum sub-basally; right harpe with
50-60 setae; hind wing dark brown; distributed along Cascades-Sierra Nevada
axis from Washington southward to southern California, and in La Sal
Mountains, Utah (Fig. 173)
E. latro (B. & Benj.) (p. 33).
Veins Cuj and M3 pale-lined, contrasting, projected into terminal area
Veins Cux and M3 not pale-lined or contrasting; cubital vein pale-lined to
reniform spot in some specimens but no pale-lined veins projected into terminal
area
Right sacculus extension at least 1.50 times length of right harpe (Fig. 90).
E. niveilinea (Grt.) (p. 40).
Right sacculus extension, at most 1.33 times length of right harpe
Vesica with small nipple-like sub-basal diverticulum on left side in addition to
normal foot-like sub-basal diverticulum; right harpe 0.9- 1.0 mm long (Fig. 91)
E. dargo (Stkr.) (p.42 ).
Vesica with only normal foot-like diverticulum sub-basally; right harpe 1.1-1. 3
mm long
Forewing with costal area pale, evenly colored, pale shade extended
uninterrupted to cubital vein on basal quarter of wing (Fig. 51)
E. quadridentata (Grt. & Rob.) (p. 37).
Forewing with costal area not particularly pale, or if so, then streaked and
unevenly colored; costal shade interrupted from cubital vein on basal quarter of
wing by a black line
Ground color of forewing brown with fine black streaks on veins (Fig. 53) ....
E. inscripta Laf. n. sp. (p. 38).
Ground color of forewing gray streaked with pale silver-gray (Fig. 15)
E. brevipennis (Sm.) (p. 22).
Hind wing white with thin black terminal line (Fig. 15)
E. brevipennis (Sm.) (p. 22).
Hind wing shaded with brown, at least on marginal third of wing
Hind wing pale with sharply contrasted dark marginal band on outer third
Hind wing smoky-brown, paler near base in many specimens
Right sacculus extension as stout as, or thinner than, right harpe (Fig. 72) ....
E. redimicula (Morr.) (p. 24).
Right sacculus extension stouter than right harpe
Basal half of prothoracic collar, and costa of forewing, pale yellow-brown;
forewing with transverse lines conspicuous; known only from vicinity of
Saskatoon, Saskatchewan (Fig. 57)
E. unica McD. (p. 39).
19
23
20
21
22
24
25
27
26
Quaest. Ent., 1981, 17 (1,2)
14
Lafontaine
26'
27 (240
21'
28 (27)
28 '
29 (280
29/
30 (270
30'
31 (30)
31'
32 (30')
32'
33 (32')
Prothoracic collar and costa of forewing essentially same color as remainder of
forewing; forewing with transverse lines obscure, veins overlaid with black
scales; distributed from southern Montana southward to Colorado and westward
to Lake Tahoe, California (Fig. 53)
E. inscripta Laf. n. sp. (p. 38).
Forewing with transverse lines conspicuous; costal, subterminal and basal areas
paler than median area . 28
Forewing with transverse lines inconspicuous; subterminal and basal areas not
paler than median area; costal area paler than remainder of forewing in many
specimens . . 30
Terminal area of forewing blurred and streaked into subterminal area;
transverse posterior line not scalloped between veins (Figs. 17-19)
E. servita (Sm.) (p. 23).
Terminal area of forewing with smooth, regular inner margin; transverse
posterior line scalloped between veins . 29
Distributed in northern Cordilleran region from British Columbia southward to
south-central Colorado, northern Utah, central Nevada and southern California
(Fig. 164)
E. auripennis Laf. (p. 25).
Distributed in southern mountains from southwestern Colorado and southern
Utah southward to central New Mexico, central Arizona and southern
California (Fig. 164)
E. arizonensis Laf. (p. 26).
Forewing shaded with maroon or red .31
Forewing shaded with buff, brown or gray . . 32
Forewing light red with contrasting silver-gray reniform and orbicular spots and
costa (Fig. 1); distributed from British Columbia southward to southern
California (Fig. 158)
E. costata (Grt.) (p. 18).
Forewing maroon dusted with dark scales; reniform and orbicular spots either
not silver-gray, or if so, then with cubital vein pale as well (Fig. 3); distributed
from western Canada southward in the Rocky Mountain region to eastern
Arizona and northern New Mexico (Fig. 159)
E. castanea Laf. n. sp. (p. 19).
Forewing largely black with contrasting silver-gray reniform and orbicular spots
and costa (Fig. 5); distributed from northern Utah southward to Arizona and
New Mexico (Fig. 161
E. foeminalis (Sm.) (p. 20).
Forewing brown or with extensive brown shading, almost black in some
specimens from Rocky Mountain region of British Columbia and Montana 33
Forewing, including terminal area, yellow-buff; series of black, sagittate spots
on proximal side of subterminal line; orbicular spot round; hind wing unevenly
shaded with light and dark brown, dark median line in most specimens (Fig.
14); distributed in Great Plains in Alberta and Montana (Fig. 161)
E. clausa McD. (p. 22).
Forewing dark brown in most specimens; if light buff, then terminal area dark
33'
The Euxoa detersa group (Lepidoptera: Noctuidae)
15
brown with dark streaks on proximal side of subterminal line; orbicular spot oval
in most specimens; hind wing evenly colored, slightly paler near base (Figs.
7-12); widespread (Fig. 160)
E. idahoensis (Grt.) (p. 21).
Key to females of E. detersa group
1 Ovipositor valve with stout setae dorsally or apically . . . 2
T Ovipositor valve without stout setae .7
2 (1) Stout setae on dorsal margin of valve; ovipositor valve without definite row of
long sub-basal setae .3
2' Stout setae primarily on apical third of valve; ovipositor valve with definite row
of long sub-basal setae . 5
3 (2) Stout setae scattered on dorsal margin of valve, not in single row (Fig. 1 55) ... 4
3' Stout setae in single row on dorsal margin of valve (Fig. 156); distributed from
western Great Plains westward (Fig. 184)
E. cicatricosa (Grt. & Rob.)(p. 45).
4 (3) Distributed east of Rocky Mountains (Fig. 183)
E. detersa (Wlk.) (p. 43).
4' Distributed in western and southern intermontane region (Fig. 185)
E. recula (Harv.) (p. 47).
5 (2') Long sub-basal setae on ovipositor valve stout and spike-like (Fig. 1 53)
E. dargo (Stkr.) (p. 42).
5' Long sub-basal setae on ovipositor valve thin and hair-like .6
6 (57) Long sub-basal setae stouter on ventral portion of valve than those on dorsal
portion (Fig. 154); forewing with cubital vein pale (Fig. 59)
E. melura McD.(p. 42).
6' Long sub-basal setae thin and hair-like on both ventral and dorsal portions of
valve (Fig. 133); forewing with cubital vein not pale (Fig. 18)
E. servita (Sm.)(p. 23).
7 (F) Sclerotized plates in wall of ductus bursae short, not extended as far anteriorly
as anterior apophyses (Fig. 118)
E. quadridentata (Grt. & Rob.)(p. 37).
1' Sclerotized plates in wall of ductus bursae longer, extended farther anteriorly
than anterior apophyses 8
8 (7') Ovipositor valve with prominent, smooth, flange-like projection at apex .9
8' Ovipositor valve without flange, or at most, trace of flange at apex, its surface
covered with minute setae . 22
9 (8) Forewing either with black basal dash, or prominently streaked longitudinally,
or both . 10
9' Forewing without black basal dash and without longitudinal streaks 17
10 (9) Orbicular spot elongate, almost bar-like; junction of ductus bursae and corpus
bursae anterior to middle of corpus bursae in most specimens 11
10' Orbicular spot round or oval; junction of ductus bursae and corpus bursae at
middle, or posterior to middle. . 13
11 (10) Orbicular spot buff-filled, not outlined in white; basal dash prominent; costa
Quaest. Ent., 1981, 17 (1,2)
16
Lafontaine
11'
12 (110
12'
13 (100
13'
14 (13)1
14'
15 (130
15'
16 (150
16'
17 (90
\r
18 (17)
18'
19 (170
19'
evenly colored, buff or pale gray (Fig. 32)
E. oblongistigma (Sm.) (p. 28).
Orbicular spot with dark center in most specimens, dark center outlined in
white; basal dash thin and inconspicuous; costa not evenly colored but marked
with white or brown (Figs. 25-29) . . 12
Forewing predominantly various shades of brown; hind wing dark brown (Fig.
29)
E. agema (Stkr.) (p. 28).
Forewing streaked with gray and white in most specimens; hind wing gray or
brown, paler at base (Figs. 25-28)
E. olivalis (Grt.) (p. 27).
Forewing streaked longitudinally; basal dash inconspicuous if present . . 14
Forewing not streaked longitudinally; basal dash prominent . . 15
Cubital vein and costa pale and contrasting (Fig. 56)
E. niveilinea (Grt.) (p. 40).
Cubital vein and costa not particularly pale (Fig. 54)
E. inscripta Laf. n. sp. (p. 38).
Flange-like projection at apex of ovipositor valve rounded, about as long as wide
(Fig. 135)
E. auripennis Laf. (p. 25).
Flange-like projection at apex of ovipositor valve finger-like, longer than wide
(Fig. 134) .16
Forewing with median area heavily dusted with black (Fig. 24). distributed
from Colorado and New Mexico westward (Fig. 164)
E. arizonensis Laf. (p. 26).
Forewing with median area light gray or brown (Fig. 21); distributed from
Montana and Wyoming eastward (Fig. 163)
E. redimicula (Morr.) (p. 24).
Anterior end of ovipositor valve markedly produced ventrally, depth of valve
greater than its length (Fig. 140) . . 18
Anterior end of ovipositor valve, at most, slightly produced ventrally, depth of
valve less than its length . . 19
Forewing yellow
E. citricolor (Grt.)(p. 29).
Forewing cream-colored or pale buff, dusted with black
E. tronella (Sm.) (p. 30).
Ovipositor valve with raised, fin-like, sclerotized plate on posterior half of dorsal
margin of valve; dorsal edge of plate straight or slightly concave, curved
abruptly at anterior end to valve (Fig. 142); sclerotized plate in ventral wall of
ductus bursae extended as far anteriorly as plate in dorsal wall (Fig. Ill) ...
E. teleboa (Sm.) (p. 31).
Ovipositor valve with rounded, or finger-like, sclerotized projection at apex, this
curved around dorsal margin of valve, its dorsal edge convex and tapered
anteriorly in most specimens, tapered abruptly in some specimens with
‘The female of E. unica McD. is unknown, however, it would probably key out to this couplet.
The Euxoa detersa group (Lepidoptera: Noctuidae)
17
sclerotized plate in dorsal wall of ductus bursae extened farther anteriorly than
plate in ventral wall (Fig. 143) . . 20
20 (190 Forewing wiht median area pale gray, basal and subterminal areas orange (Fig.
46)
E. murdocki (Sm.) (p. 34).
20' Forewing with basal median and subterminal areas similar in color, if not in
shade .21
21 (200 Forewing dark brown, heavily dusted with gray in most specimens; hind wing
dark brown to base (Fig. 45); distributed along Cascades-Sierra Nevada axis
from Washington southward to southern California; disjunct population in La
Sal Mts., Utah (Fig. 173)
E. latro (B. & Benj.) (p. 33).
2 Y Forewing light gray, light brown, or orange; hind wing gray, paler toward base
in most specimens (Figs. 41-43); distributed throughout Great Plains and Great
Basin (Fig. 172)
E. moerens (Grt.) (p. 32).
22 (80 Forewing with prominent, black, basal dash . . 23
22' Forewing without basal dash . 29
23 (22) Ovipositor valve with short, conical setae apically; forewing with veins and
M3 not pale-lined . . 24
23' Ovipositor valve with fine-tipped setae apically, or with veins Cuj and M3
pale-lined, pale streaks projected into terminal area . . 28
24 (23) Forewing shaded with maroon or reddish brown . . 25
24' Forewing shaded with buff, brown or gray . . 26
25 .(24) Ovipositor valve rounded or somewhat truncate apically; forewing light reddish
brown with contrasted silver-gray costa, reniform spot and orbicular spot;
cubital vein not pale (Fig. 2); distributed from British Columbia southward to
southern California (Fig. 158)
E. costata (Grt.) (p. 18).
25' Ovipositor valve acutely angled apically; forewing maroon or .chestnut-brown,
dusted with black; costa, reniform spot and orbicular spot either without
silver-gray shading, or with cubital vein pale as well (Fig. 3); distributed from
western Canada southward in the Rocky Mountain region to eastern Arizona
and northern New Mexico (Fig. 159)
E. castanea Laf n. sp. (p. 19).
26 (24') Forewing extensively black with contrasted silver-gray reniform and orbicular
spots and costa (Fig. 6); distributed from northern Utah southward .to New
Mexico and Arizona (Fig. 161)
E. foeminalis (Grt.) (p. 20).
26' Forewing brown or streaked with brown, almost black in some specimens from
Rocky Mountain region of British Columbia and Montana . . 27
27 (26') Forewing, including terminal area, yellow-buff, series of black sagittate spots on
proximal side of subterminal line; orbicular spot round; hind wing unevenly
shaded with light and dark brown, dark median line in most specimens (Fig.
14); distributed in Great Plains of Alberta and Montana (Fig. 161)
E. clausa McD. (p. 22).
Quaest. Ent., 1981, 17 (1,2)
18
Lafontaine
21' Forewing brown in most specimens; if light buff, then terminal area of forewing
dark brown with dark streaks on proximal side of subterminal line; orbicular
spot oval in most specimens; hind wing evenly colored, slightly lighter toward
base (Figs. 7-12); widely distributed (Fig. 160)
E. idahoensis (Grt.) (p. 21).
28 (230 Hind wing mottled gray-brown with median line on ventral surface; forewing
with inconspicuous, irregular, basal dash; forewing with dark streaks extended
from terminal area into subterminal area interrupted by pale subterminal line
(Fig. 16)
E. brevipennis (Sm.) (p. 22).
28' Hind wing evenly colored, pale gray-brown, slightly lighter at base, without
trace of median line on ventral surface; forewing with conspicuous basal dash
extended to transverse anterior line; forewing with dark streaks extended
uninterrupted from terminal area into subterminal area, subterminal line not
evident (Fig. 50)
29 (220
29'
30 (290
E. laetificans (Sm.) (p. 36).
Forewing marked with silver-gray; inner edge of terminal area with W-mark
where pale-lined veins Cuj and M3 project into it (Fig. 16)
E. brevipennis (Sm.) (p. 22).
Forewing brown or gray, without silver-gray streaks; inner edge of terminal area
without W-mark, veins Cuj and M3 not pale-lined 30
Forewing with median area pale gray-brown, basal and subterminal areas pale
gray (Fig. 47)
E. dodi McD. (p. 34).
Forewing with median and terminal areas dark brown, basal and subterminal
areas paler brown with copper suffusion (Fig. 48)
E. infracta (Morr.) (p. 35).
Species of the Euxoa detersa group
Euxoa costata (Grote)
Figs. 1, 2, 65,96, 126, 158
Agrotis costata Grote, 1876, p. 80; Vancouver Island, B.C.
Rhynchagrotic costata; Smith, 1890b, p. 38; redescription.
Rhynchagrotis costata; Smith, 1893, p. 55.
Triphaena costata ; Hampson 1903, p. 631.
Euxoa costata ; Barnes and McDunnough, 1 9 1 7, p. 43.
Euxoa costata ; Draudt, 1924, p. 48; redescription.
Description. — Vestiture of head and thorax and ground color of forewing reddish brown. Transverse lines obscure.
Reniform and orbicular spots and costa shaded with silver-gray. Hind wing smoky-brown.
Expanse: 36.1 ± 1.3 mm. Length of forewing: 1 7.0 ± 0.8 mm (50 specimens).
Male genitalia (Fig. 65). Sacculus extensions slightly longer than harpes and about as stout. Sub-basal diverticulum of
vesica large; median diverticulum positioned about 1 /3 distance from sub-basal bend in vesica to apex. Vesica projected to
right apically.
Length of right sacculus extension: 1 .7 1 ± 0.08 mm (6 specimens).
Length of left sacculus extension: 1 .62 ± 0.08 mm (7 specimens).
Length of right harpe: 1.61 ±0.04 mm (7 specimens).
The Euxoa detersa group (Lepidoptera: Noctuidae)
19
Length of right sacculus: 1.52 ±0.09 mm (7 specimens).
Female genitalia (Figs. 96, 126). Ductus bursae attached to corpus bursae 0.33-0.50 way along right side from
posterior end. Ovipositor valves with short, conical setae near apex. Ovipositor valve rounded or somewhat truncate
apically.
Type material. - The holotype of Agrotis costata Grote is a female in the collection of the
British Museum (Natural History). The specimen has been dissected.
Distribution and period of flight.. - Euxoa costata occurs from southern British Columbia
southward in the Cascades and Sierra Nevada Mountains to southern California (Fig. 158).
Fifty-eight specimens from 21 localities were examined.
Specimens have been collected from mid-July until early September.
Habitat. - This species inhabits dry conifer forests, especially those dominated by
Douglas-fir ( Pseudotsuga menziesii (Mirbel) Franco).
Remarks. - Male specimens of Euxoa costata may be distinguished from those of other
similar species by the longer sacculus extensions and harpes of the male genitalia. In external
appearance, specimens of E. costata are most easily confused with those of E. castanea; for
distinguishing characters see discussion under that species.
Euxoa castanea Lafontaine, new species
Figs. 3,4, 66, 97, 127, 159
Description. — Antenna of male biserrate and bifasciculate. Antenna of female filiform. Frontal tubercle present.
Vestiture of head and thorax dark reddish brown.
Forewing ground color chestnut brown dusted with black. Transverse anterior and posterior lines indistinct in most
specimens. Subterminal area slightly paler than median area in most specimens. Terminal area dark gray-brown. Costa,
reniform spot, and orbicular spot paler than median area, shaded with buff or silver-gray. Orbicular spot oval in most
specimens. Area between reniform and orbicular spots, and proximal to orbicular spot, black in most specimens. Cubital
vein pale and contrasting from wing base to reniform spot. Basal dash and claviform spot black.
Fiind wing of both male and female gray-brown; discal spot present.
Expanse: 36.4 ± 0.7 mm. Length of forewing: 16.8 ± 1.6 mm. (100 specimens).
Male genitalia (Fig. 66). Different from those of E. costata only in proportion. Harpe and sacculus extension in males
of E. castanea shorter than those of E. costata males.
Length of right sacculus extension: 1.33 ±0.10 mm (25 specimens).
Length of left sacculus extension: 1 .24 ±0.10 mm (25 specimens).
Length of right harpe: 1 .28 ± 0.06 mm (25 specimens)
Length of right sacculus: 1.35 ±0.07 mm (25 specimens).
Female genitalia (Figs. 97, 127). Ovipositor valve triangular in outline, pointed apically. Small flange-like projection
on ovipositor valve of female specimens from southern Rocky Mountains in Colorado and New Mexico.
Type material. - Holotype male: Golden, B.C., 2 mi E., 3000 ft, 21 July 1960 (D.F.
Hardwick), Type No. 15939 in the Canadian National Collection, Ottawa. Paratypes: 78
males, 49 females: ALBERTA: Banff, 30 July-7 Aug. 1922 (C.B.D. Garrett); Exshaw, 5 mi
N.E., 4225 ft, 18 July 1961 (D.F. Hardwick); Horseshoe Canyon, Drumheller, 2750 ft, 17 July
1960 (D.F. Hardwick); Lake Louise, 4 mi SE., 6000 ft, 20 July 1960 (D.F.. Hardwick);
Nordegg, 17 July-8 Aug. 1921 (J. McDunnough) BRITISH COLUMBIA: Crow’s Nest, 5 mi
NW., 4300 ft, 26 July 1960 (D.F. Hardwick); Fernie, 1 1 mi NE., 3600 ft, 27 July 1960 (D.F.
Hardwick); Golden, 2 mi E., 3000 ft, 21 July 1960 (D.F. Hardwick); Radium Hot Springs, 16
mi N., 2900 ft, 22 July 1960 (D.F. Hardwick); Radium Hot Springs, 28 mi S., 23 July 1960
(D.F. Hardwick).
Distribution and flight period. - Euxoa castanea occurs across the aspen parkland and
southern boreal forest of western Canada, southward in the Rocky Mountains to New Mexico.
It occurs in the northern Great Plains in suitable forested habitat. Approximately 700
specimens from 87 localities were examined.
Quaest. Ent., 1981, 17 (1,2)
20
Lafontaine
Specimens were collected from early July until late August.
Habitat. - This species is found most abundantly in dry conifer forests of Douglas-fir
(Pseudotsuga menziessi (Mirbel) Franco), and lodgepole pine ( Pinus contorta Dough). It is
also found-in spruce and aspen forests, particularly in the northeastern portion of its range.
Remarks. - Specimens of E. castanea may be distinguished from those of E. costata and E.
idahoensis by the dark chestnut-brown color of the forewing, and also from the former by the
pale cubital vein. In specimens of E. costata the forewing is light reddish brown without dark
dusting; the cubital vein is not pale. Specimens of E. idahoensis have gray or brown forewings;
the cubital vein is pale in many specimens.
Euxoa foeminalis (Smith)
Figs. 5,6,67,98, 128, 161
Carneades foeminalis Smith, 1 900, p. 454; Colorado.
Paragrotis foeminalis ; Dyar, 1902, p. 148.
Euxoa foeminalis ; Hampson, 1903, p. 296.
Euxoa foeminalis ; Draudt, 1924, p. 48; redescription.
Carneades foeminalis-, Rindge, 1955, p. 112; type material.
Description. — Vestiture of head and thorax black and gray. Ground color of forewing dark silver-gray dusted with
black. Transverse lines obscure. Subterminal area, costa, reniform spot, and orbicular spot pale silver-gray, contrasted with
darker ground color of remainder of forewing. Cubital vein not pale-lined. Hind wing smoky-brown.
Expanse: 36. 4± 1.4 mm. Length of forewing: 1 6.9 ± 0.9 mm. (40 specimens).
Male genitalia (Fig. 67). Indistinguishable from those of E. idahoensis.
Length of right sacculus extension: 1 .51 ±0.07 mm (20 specimens).
Length of left sacculus extension: 1 .45 ± 0.08 mm (20 specimens).
Length of right harpe: 1.39 ±0.04 mm (20 specimens).
Length of right sacculus: 1 .49 ± 0.05 mm (20 specimens).
Female genitalia (Figs. 98, 128). Ovipositor valves more broadly rounded than those of E. idahoensis females.
Type material. - Carneades foeminalis was described by Smith on the basis of two nominal
male (one is a female) and two female specimens from Garfield County, Colorado. The male
specimen is in the collection of the American Museum of Natural History, New York; three
female specimens are in the collection of the United States National Museum (type No. 4784).
A lectotype will be designated by Todd (1982).
Distribution and period of flight. - Euxoa foeminalis occurs in southwestern United States
from north-central Utah and western Colorado southward to northern New Mexico and
Arizona (Fig. 161). Forty specimens were examined from 6 localities.
Specimens have been collected from late May until mid-July.
Habitat. - The habitat of this species is inferred from relatively few localities to be conifer
forests. This species, however, was collected at one locality in an open oak-pine forest.
Remarks. - Euxoa foeminalis specimens resemble dark specimens of E. idahoensis from
montane areas of British Columbia and Montana; they differ from these in that the forewing is
shaded with black rather than dark brown and the costa, reniform spot, orbicular spot, and
subterminal area are all shaded with silver-gray, the cubital vein is not pale. In dark specimens
of E. idahoensis the subterminal area is not pale, the cubital vein is pale in most specimens
(compare Figs. 5 and 10). Specimens of E. idahoensis from the area of sympatry with E.
foeminalis are shaded with pale brown and the cubital vein is pale in most specimens.
The Euxoa detersa group (Lepidoptera: Noctuidae)
21
Euxoa idahoensis (Grote)
Figs. 7,8, 9, 10, 11, 12, 68,99, 129, 130, 160
Agrotis idahoensis Grote, 1878, p. 171; Idaho.
Carneades idahoensis : Smith, 1890b, p. 197; redescription.
Carneades idahoensis; Smith, 1893, P. 105; catalogue.
Paragrotis idahoensis: Dyar, 1902, p. 148.
Euxoa idahoensis ; Hampson, 1903, p. 295.
Euxoa costata idahoensis ; Barnes and McDunnough, 1917, p. 43.
Euxoa costata form idahoensis ; Draudt, 1924, p. 48.
Euxoa costata idahoensis ; McDunnough, 1938, p. 60.
Agrotis furtivus Smith, 1890a, p. 56; California.
Carneades furtivus. Smith, 1890b, p. 197.
Carneades furtivus\ Smith, 1893, p. 105; catalogue.
Paragrotis furtivus ; Dyar 1902, p. 148.
Euxoa furtiva\ Hampson, 1903, p. 297 .
Euxoa furtiva\ Barnes and McDunnough,
1917, p. 43; = Euxoa costata idahoensis.
Euxoa furtiva\ Draudt, 1924, p. 48; = Euxoa costata form idahoensis.
Description. — Vestiture of head and thorax gray-brown. Color and markings of forewing extremely variable; most
common and widespread form (Figs. 7, 8) with forewing pale brown with dark brown streaks. Terminal space and sagittate
spots proximal to subterminal line dark gray-brown. Costa, reniform spot, orbicular spot, and cubital vein paler than
remainder of forewing. Orbicular spot oblong in most specimens.
Specimens from Great Plains region pale gray (Fig. 9), those from montane areas of British Columbia and Montana
dark brown (Fig. 10). Some specimens from Black Hills, South Dakota, and from montane areas of Colorado heavily
streaked (Fig. 12) and resembling specimens of E. oblongistigma. Many specimens from northern Great Basin without
pale costa (Fig. 1 1 ) and frequently without pale-filled spots.
Hind wing smoky-brown, paler toward base; discal spot present.
Expanse: 36. 2± 1.7 mm. Length of forewing: 1 6.3 ± 0.8 mm (100 specimens).
Male genitalia (Fig. 68). Indistinguishable from those of E. castanea, E. foeminalis , and E. clausa.
Length of right sacculus extension: 1 .42 ±0.09 mm (30 specimens).
Length of left sacculus extension: 1 .33 ± 0.09 mm (30 specimens).
Length of right harpe: 1 .39 ± 0.06 mm (30 specimens).
Length of right sacculus: 1 .37 ±0.07 mm (30 specimens).
Female genitalia (Figs. 99, 129, 130). Ovipositor valve of females from northern and western portion of range rounded
apically (Fig. 129), those of females from Wyoming Basin and southern Rocky Mountains more acutely angled, small
flange-like projection in some specimens (Fig. 130).
Type material. - The holotype of Agrotis idahoensis is a male in the collection of the British
Museum (Natural History) from Idaho. Agrotis furtivus was described by Smith on the basis
of three nominal females (one specimen is a male) from California. A lectotype will be
designated by Todd (1982) from the syntypes in the collection of the United States National
Museum.
Distribution and period of flight. - Euxoa idahoensis is found in western North America as
far east as south-central Saskatchewan and western South Dakota. Approximately 600
specimens were examined from 96 localities.
Adults have been collected from mid-June until late August, however, most were collected
in the latter half of July.
Habitat. - Although this species is most common in dry conifer forests of pine, fir, and
Douglas-fir, it is also found occasionally in pinon-juniper woodland in the Great Basin, and in
deciduous thickets in the Great Plains.
Remarks. - Euxoa idahoensis is probably the most variable species of any in the E. detersa
group with numerous color forms occurring in different parts of its range. The most widespread
form (Figs. 7 and 8) grades imperceptibly into the other more localized geographic forms where
they occur together.
Quaest. Ent., 1981, 17(1,2)
22
Lafontaine
Specimens of Euxoa idahoensis cannot be distinguished from those of the other members of
the E. costata -E. foeminalis -E. clausa series by structural characters. They can be
distinguished from specimens of these species by characters in key.
Euxoa clausa McDunnough
Figs. 13, 14, 69, 100, 131, 161
Euxoa clausa McDunnough, 1923, p. 163; Lethbridge, Alberta.
Euxoa costata form clausa ; Draudt, 1924, p. 48.
Euxoa clausa ; McDunnough, 1 938, p. 6 1 .
Description. — Male antenna markedly biserrate, serrations deeper than those of E. idahoensis males. Vestiture of
head and thorax light yellowish brown. Ground color of forewing pale yellow-brown overlaid with dusting of brown scales.
Transverse lines present in most specimens but obscure. Terminal area concolorous or sightly darker than remainder of
forewing. Subterminal area with series of black sagittate spots on proximal side of subterminal line in most specimens (Fig.
14). Costa, orbicular spot, and reniform spot not particularly pale. Hind wing unevenly colored with smoky-brown shading
on outer third of wing, on median line, and on discal spot.
Expanse: 34.9 ± 2.3 mm. Length of forewing: 1 5.8 ± 1 .0 mm (40 specimens).
Male genitalia (Fig. 69). Indistinguishable from those of E. idahoensis males.
Length of right sacculus extension: 1.40 ±0.09 mm (20 specimens).
Length of left sacculus extension: 1 .33 ±0.08 mm (20 specimens).
Length of right harpe: 1 .43 ± 0.06 mm (20 specimens).
Length of right sacculus: 1 .40 ± 0.05 mm (20 specimens).
Female genitalia (Figs. 100, 131). Indistinguishable from those of E. idahoensis.
Type material. - The holotype of E. clausa is a male from Lethbridge, Alberta, type No.
598 in the Canadian National Collection, Ottawa. The specimen has not been dissected.
Distribution and period of flight. - Specimens of E. clausa have been collected only in the
northwestern Great Plains (Fig. 161). Fifty specimens from six localities were examined; these
were collected from mid-July until mid-August.
Habitat. - From limited available data, it appears that this species inhabits open shortgrass
prairie.
Remarks. - Euxoa clausa occurs sympatrically with a pale form of E. idahoensis (Fig. 9) in
the northern Great Plains. Specimens of these two species may be distinguished by characters
in key.
Euxoa brevipennis (Smith)
Figs. 15, 16, 70, 101, 132, 168
Agrotis brevipennis Smith, 1887, p. 455; California, Colorado, Nevada.
Carneades brevipennis ; Smith, 1890b, p. 140; redescription.
Carneades brevipennis ; Smith, 1893, p. 89; catalogue.
Paragrotis brevipennis, Dyar, 1902, p. 140.
Euxoa brevipennis ; Hampson, 1903, p. 208.
Euxoa brevistriga Smith, 1910b, p. 257; Colorado, probably Denver.
Euxoa brevipennis brevistriga ; Barnes and McDunnough, 1917, p. 41.
Euxoa brevipennis form brevistriga; Draudt, 1924, p. 37.
Euxoa brevistriga; Rindge, 1955, p. 103; type material.
Euxoa angulirena Smith, 1910b, p. 257; Colorado.
Euxoa brevipennis ab. angulirena; Barnes and McDunnough, 1917, p. 41.
Euxoa angulirena; Rindge, 1955, p. 101; type material.
Description. — Vestiture of head and thorax gray, or mixture gray and black scales. Ground color of forewing gray,
streaked longitudinally with white and dark gray in most specimens; some specimens with little or no'pale streaking.
Transverse anterior and posterior lines barely evident or absent. Subterminal area and posterior half of median area
streaked in most specimens with veins dark, interveinal area pale. Terminal area dark gray, its inner margin streaked into
subterminal area; streaks interrupted by pale subterminal line in many specimens. Dark terminal shade indented by pale
The Euxoa detersa group (Lepidoptera: Noctuidae)
23
W-mark where pale-lined veins Cuj and M3 project into it. Costa pale in most specimens. Reniform and orbicular spots
with pale line between black outline and gray-filled center. Black, basal dash in some specimens. Hind wing of male white
with narrow gray terminal line and in some specimens on veins as well. Hind wing of female pale gray with dusting of
darker gray scales on outer third of wing, on median line, and on veins.
Expanse: 35.1 ± 1.5 mm. Length of forewing: 17.1 ± 0.8 mm (100 specimens).
Male genitalia (Fig. 70). Distinguishable from those of other species in E. detersa group by characters of vesica.
Vesica with bend, or slight twist in it before it bends to project dorsally. Sub-basal diverticulum curved to left apically.
Sacculus extension not straight or incurved as are those of most other species but slight S-curved, bent away from
cucullus near apex. Sacculus extensions stouter than those of males of E. costata - E. foeminalis -E. clausa series.
Length of right sacculus extension: 1.45 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1.39 ±0.10 mm (20 specimens).
Length of right harpe: 1.25 ±0.06 mm (20 specimens).
Length of right sacculus: 1.42 ±0.06 mm (20 specimens).
Female genitalia (Figs. 101, 132). Similar to those of E. dodi (Fig. 115) and E. infracta (Fig. 116) but corpus
bursae slightly constricted mesially in most females of E. brevipennis.
Type material. - It is not known how many specimens Smith based Agrotis brevipennis on.
The only type material located is two specimens in the collection of the United States National
Museum. One of these will be selected as lectotype by Todd (1982).
Smith described Euxoa brevistriga from two males collected in Colorado (probably near
Denver according to Smith). One of these males, in the collection of the American Museum of
Natural History, will be selected as lectotype by Todd (1982).
The holotype of Euxoa angulirena Smith is a female from Colorado in the collection of the
American Museum of Natural History. The specimen has been dissected.
Distribution and period of flight. - Euxoa brevipennis occurs throughout the intermontane
region and in the western Great Plains (Fig. 168). Approximately 350 specimens from 53
localities were examined.
This is a late-flying species; adults have been collected from early September until late
October in the north and November in the south.
Habitat. - This species inhabits dry, open sagebrush areas; it also occurs in open
pinon-juniper woodland where sagebrush is abundant.
Remarks. - A combination of large size, pattern of forewing streaking, lack of transverse
lines, and pale hind wing allow specimens of E. brevipennis to be distinguished from those of
other species in the E. detersa group. The larva has been reared on tumblemustard
{Sisymbrium spp.), hare’s-ear mustard {Conringia orientalis (L.) Dumort), Russian thistle
{Salsola kali L.) and sweet clover {Melilotus spp.) in Montana (Cook, 1930b).
Euxoa servita (Smith)
Figs. 17, 18, 19,71, 102, 133, 162
Carneades servitus Smith, 1895, p. 336; Colorado; Calgary, Alberta.
Paragrotis servitus-, Dyar, 1902, p. 148.
Euxoa servita-, Hampson, 1903, p. 308.
Euxoa servita-, Dod, 1905, p. 148; probably aberration of E. redimicula.
Euxoa redimicula ab. servita-, Barnes and McDunnough, 1917, p. 43.
Euxoa servita ; McDunnough, 1949, p. 5; good species.
Euxoa servita-, McDunnough, 1950, p. 391; bibliography.
Agrotis servita-, Forbes, 1954, p. 42.
Carneades servitus-, Rindge, 1955, p. 130; type material.
Euxoa servita-, Lafontaine, 1974a, p. 409; redescription.
Euxoa servita novangliae McDunnough, 1950, p. 391; Franconia, New Hampshire.
Description. — Vestiture of head and thorax, gray. Dorsal half of prothoracic collar reddish brown or gray. Ground
color of forewing various shades of gray or brownish gray, darker in specimens from forested areas. Maculation complete
but appearing streaked and blurred in most specimens. Transverse posterior line slightly scalloped between veins, series of
Quaest. Ent., 1981, 17 (1,2)
24
Lafontaine
small, dark, sagittate spots on proximal side extended into median area. Terminal area dark gray, its inner margin
streaked into subterminal area Several small, black streaks extended from terminal area into subterminal area opposite
reniform spot. Hind wing of both sexes smoky-brown.
Expanse: 3 1 .4 ± 1.9 mm. Length of forewing: 15.4 ±0.7 mm (100 specimens).
Male genitalia (Fig. 71). Distinguishable from those of E. redimicula . E. auripennis, and E. arizonensis only by
numerical methods (see Lafontaine, 1974a).
Length of righ sacculus extension: 1.51 ±0.09 mm (40 specimens).
Length of left sacculus extension: 1.41 ±0.1 1 mm (40 specimens).
Length of right harpe: 1.24 ±0.06 mm (40 specimens).
Length of right sacculus: 1.39 ±0.06 mm (40 specimens).
Female genitalia (Figs. 102, 133). Distinguishable from those of other three members of E. servita-E. arizonensis
series by form of ovipositor valve. Apical third of ovipositor valve clothed with stout setae, appearing mace-like; those of
other species of series clothed with fine setae, sclerotized flange-like projection at apex of each valve.
Type material. - Carneades servitus was described by Smith from two specimens: a male
from Colorado in the collection of the American Museum of Natural History, New York; a
female from Calgary, Alberta, in the collection of the United States National Museum. A
lectotype will be designated by Todd (1982).
The holotype of Euxoa servita novangliae is a female from Franconia, New Hampshire in
the collection of the American Museum of Natural History. The specimen has been dissected.
Distribution and period of flight. - Euxoa servita occurs from Nova Scotia westward across
southern Canada and northern United States to British Columbia and southward in the Rocky
Mountain region to central Arizona and New Mexico (Fig. 162). It has not been collected in
the Cascades or Sierra Nevada Mountains. Approximately 1000 specimens from 147 localities
were examined.
Adults have been collected from mid-July until late August.
Habitat. - East of the Rocky Mountains this species inhabits forested areas dominated by
spruce and aspen. In British Columbia and in the Rocky Mountain region it is found in aspen
and Douglas-fir ( Pseudotsuga menziesii (Mirbel) Franco) forests, and more uncommonly, in
areas dominated by lodgepole pine ( Pinus contorta Dougl.) and pondersoa pine ( Pinus
ponderosa Dougl.). In general, this species occurs in more mesic habitats than E. redimicula ,
E. auripennis , and E. arizonensis.
Remarks. - Female specimens of E. servita are easily distinguished from those of E.
redimicula , E. auripennis , and E. arizonensis by the stout setae rather than apical flange-like
projection on the ovipositor valve. Separation of males of the four species was discussed by
Lafontaine (1974a). Most male specimens can be distinguished by characters in the key.
Euxoa servita adults have an earlier flight period than do those of the other three species,
however, there is usually some overlap.
The subspecific epithet novangliae McDunnough has been used for dark brown specimens
from populations in eastern North America. I do not consider eastern populations to be
subspecifically distinct because dark brown forms also occur in mesic conifer forest areas in
western North America. Darker wing color appears to be associated with more mesic habitats.
Euxoa redimicula (Morrison)
Figs. 20,21,72, 103, 134, 163
Agrotis redimacula Morrison, 1874, p. 165; Colorado, New York, Massachusett.
Agrotis redimicula ; Morrison, 1875b, p. 57; spelling correction.
Carneades redimicula ; Smith, 1890b, p. 202; redescription.
Carneades redimicula-. Smith, 1893, p. 107; catalogue.
Paragrotis redimicula-. Dyar, 1902, p. 148.
Euxoa redimicula-. Hampson, 1903, p. 306.
The Euxoa detersa group (Lepidoptera: Noctuidae)
25
Euxoa redimicula ; McDunnough, 1949, p. 5; type limitation.
Euxoa redimicula ; McDunnough, 1950, p. 390; bibliography.
Agrotis redimicula ; Forbes, 1954, p. 42; redescription.
Euxoa redimicula ; Lafontaine, 1974a, p. 412; redescription; lectotype selection.
Agrotis redimicula ; Lafontaine, 1981; correction of original spelling.
Description. — Vestiture of head and thorax gray-brown. Upper half of prothoracic collar gray or reddish brown.
Ground color of forewing pale brown or brownish-gray. Transverse lines present but obscure in most specimens. Basal,
median, and subterminal areas essentially concolorous in most specimens; subterminal area slightly paler in some
specimens. Terminal area gray-brown, darker than ground color. Area between reniform and orbicular spots, and proximal
to orbicular spot, black and contrasted with ground color. Claviform spot small. Hind wing of male with dark terminal
band; basal two-thirds of wing pale buff, outer third smoky-brown. Hind wing of female smoky-brown to base.
Expanse: 3 1 .6 ± 1.8 mm. Length of forewing: 1 5.6 ± 0.8 mm (60 specimens).
Male genitalia (Fig. 72). Distinguishable from those of E. servita , E. auripennis and E. arizonensis males only by
numerical methods (see Lafontaine, 1974a).
Length of right sacculus extension: 1 .39 ±0.13 mm (22 specimens).
Length of left sacculus extension: 1 .30 ±0.16 mm (22 specimens).
Length of right harpe: 1 . 1 9 ± 0.05 mm (22 specimens).
Length of right sacculus: 1 .38 ±0.07 mm (22 specimens).
Female genitalia (Figs. 103, 1 34). Distinguishable from those of E. servita female by the presence of apical flange-like
projection and from those of E. auripennis female by elongate finger-like flange-like projection rather than rounded,
ear-like projection. Not distinguishable from those of E. arizonensis female although flange-like projection of E.
redimicula female larger than that of E. arizonensis female (compare figures 134 and 136).
Type material. - The lectotype of Agrotis redimicula is a female from Cambridge,
Massachusetts, in the collection of Michigan State University, East Lansing. The specimen has
been dissected.
Distribution and period of flight. - Euxoa redimicula occurs from Nova Scotia southward
to New Jersey and westward to western Saskatchewan, central Montana and western
Wyoming. Approximately 120 specimens from 49 localities were examined.
Specimens have been collected from late July until mid-September.
Habitat. - In the east, Euxoa redimicula inhabits dry deciduous forests. In the Great
Plains, it occurs in groves of aspen, cottonwood, and ponderosa pine.
Remarks. - Females of E. redimicula may be distinguished from those of E. servita and E.
auripennis by the long, finger-like flange-like projection on the ovipositor valve and by lack of
contrast between median and subterminal areas of the forewing. Males of E. redimicula may
be distinguished by hind wing color; this is pale buff with a dark terminal band. Hind wings of
males of E. servita E. auripennis , and E. arizonensis are dark, slightly paler near base in many
specimens.
Euxoa auripennis Lafontaine
Figs. 22, 23,73, 104, 135, 164
Euxoa auripennis Lafontaine, 1974a, p. 412; Cranbrook B.C.
Description. — Vestiture of head and basal half of prothoracic collar white; upper half of prothoracic collar reddish
brown in most specimens, gray in some specimens Vestiture of remainder of thorax gray or brown laterally, white dorsally.
Ground color of forewing various shades of gray. Transverse lines prominent. Transverse posterior line scalloped between
veins, black shade of teeth extended on veins in subterminal area. Basal and subterminal areas silver-gray. Median area
gray-brown, contrasted with paler coloration of basal and subterminal areas. Terminal area dark gray, its inner margin not
streaked into subterminal area. Reniform and orbicular spots large, pale-filled. Basal dash black, prominent. Hind wing
smoky-brown, paler near base.
Expanse: 31.3 ± 1 .5 mm. Length of forewing: 1 5.4 ± 0.7 mm ( 1 00 specimens).
Male genitalia (Fig. 73). Distinguishable from those of E. servita , E. redimicula , and E. arizonensis only by numerical
methods (see Lafontaine, 1974a).
Length of right sacculus extension: 1 .35 ± 0. 1 0 mm (34 specimens).
Length of left sacculus extension: 1 .23 ±0.14 mm (34 specimens).
Length of right harpe: 1 . 1 2 ± 0.04 mm (34 specimens).
Quaest. Ent., 1981, 17 (1,2)
26
Lafontaine
Length of right sacculus: 1.30±0.05 mm (34 specimens).
Female genitalia (Figs. 104, 135). Distinguishable from those of E. servita female by sclerotized flange-like
projection on ovipositor valve and from those of E. redimicula and E. arizonensis females by rounded, ear-like, rather
than elongate, finger-like sclerotized projection.
Type material. - The holotype of E. auripennis Lafontaine is a female (type No. 13191) in
the Canadian National Collection, Ottawa.
Distribution and period of flight. - Euxoa auripennis occurs in southern Canada and
northern United States from southern Manitoba and northeastern North Dakota westward to
British Columbia and Washington. In western North America it occurs southward in the
Rocky Mountain region to south-central Colorado and southward in the Cascades-Sierra
Nevada mountains to southern California (Fig. 164). Approximately 400 specimens from 88
localities were examined.
Specimens have been collected from late July until late September.
Habitat. - In most of its range Euxoa auripennis occurs in ponderosa pine ( Pinus
ponderosa Dougl.) and lodgepole pine ( Pinus contorta Dough) parkland. In the northern Great
Plains region it occurs in aspen woodland.
Remarks. - Females of Euxoa auripennis may be distinguished from those of E. servita, E.
redimicula and E. arizonensis by characters described above and in key. Males of E. servita
and E. redimicula may be distinguished from E. auripennis male by wing color and maculation
characters given in key; those of E auripennis and E. arizonensis can frequently be
distinguished only by numerical methods (see Lafontaine, 1974a) and by range. Where ranges
of the two species overlap in southern Colorado, they are separated by habitat: E. auripennis
occurs in more mesic habitats, such as pine forests, than does E. arizonensis which occurs in
xeric pinon-juniper woodland. Habitat data are not available for the southern California
collections of the two species.
Euxoa arizonensis Lafontaine
Figs. 24, 74, 105, 136, 164
Euxoa arizonensis Lafontaine, 1974a, p. 416; White Mts., Arizona.
Description. — Maculation and color almost identical to that of E. auripennis specimens but forewing darker in
most E. arizonensis specimens. Basal and subterminal areas silver-gray, dusted with black scales. Median area dark gray,
without pale streak distal to claviform spot in most specimens. Reniform and orbicular spots silver-gray, dusted with black
scales.
Expanse: 29.5 ± 1 .9 mm. Length of forewing: 1 4.9 ± 0.7 mm (40 specimens).
Male genitalia (Fig. 74). Distinguishable from those of E. servita, E. redimicula , and E. auripennis only by numerical
methods (see Lafontaine, 1974a).
Length of right sacculus extension: 1 .43 ±0.07 mm (25 specimens).
Length of left sacculus extension: 1 . 1 9 ± 0.08 mm (25 specimens).
Length of right harpe: 1.06 ±0.04 mm (25 specimens).
Length of righ sacculus: 1 .28 ± 0.05 mm (25 specimens).
Female genitalia (Figs. 105, 136). Distinguishable from those of E. servita female by the presence of apical flange-like
projection on ovipositor valve and from those of E. auripennis by elongate, finger-like flange-like projection rather than
rounded, ear-like projection. Indistinguishable from those of E. redimicula female although flange-like projection of E.
arizonensis female smaller than that of E. redimicula female (compare figures 1 34 and 1 36).
Type material. - The holotype of E. arizonensis is a female (type No. 13192) in the
Canadian National Collection, Ottawa.
Distribution and period of flight. - Euxoa arizonensis occurs in southwestern United States
from central Colorado and western New Mexico westward to central Nevada and southern
California (Fig. 164). Approximately 100 specimens from 21 localities were examined.
The Euxoa detersa group (Lepidoptera: Noctuidae)
27
Specimens have been collected from early August until mid-September
Habitat. - In the southern portion of its range Euxoa arizonensis occurs in ponderosa pine
(Pinus ponderosa Dougl.) parkland. In Colorado, Nevada, and possibly in California, E.
arizonensis occurs in pinon-juniper woodland.
Remarks. - Euxoa arizonensis is very similar to E. auripennis in both markings and
structural characters (see remarks under E. auripennis and the two might better be treated as
conspecific subspecies. They are ranked as species here because differences in ovipositor valves
suggests that females of the two species may oviposit in different soil types (i.e. are ecologically
distinct) and differences in ovipositor valve shape are most pronounced in the area of sympatry
viz. central Colorado.
Euxoa olivalis (Grote)
Figs. 25,26, 27,75, 106, 137, 165
Agrotis olivalis Grote, 1879, p. 43; Colorado.
Carneades olivalis-. Smith, 1890b, p. 141; redescription.
Carneades olivalis-. Smith, 1893, p. 89; catalogue.
Paragrotis olivalis-, Dyar, 1902, p. 140.
Euxoa olivalis-, Hampson, 1903, p. 211.
Euxoa olivalis-, Draudt, 1924, p. 38; ffr E. plagigera (Morr.).
Euxoa mcdunnoughi Cook, 1930a, p. 147; Jefferson Co., Montana.
Euxoa mcdunnoughi', McDunnough, 1932, p. 230; = E. olivalis.
Description. — Vestiture of head and thorax gray, brown and black scales; transverse band of white scales posterior
to prothoracic collar. Ground color of forewing brown or gray, longitudinally streaked with black, pale gray, and dark
brown. Transverse lines essentially absent. Terminal area slightly darker than ground color of forewing. Series of black,
sagittate spots on proximal side of subterminal line in most specimens. Costa not pale, or with dark and pale lines crossing
it where transverse lines would be. Color of costa separated from pale-lined cubital vein near wing base by black line.
Orbicular and reniform spots outlined in white. Orbicular spot elongate and rectangular, or slightly oval, about twice as
long as wide. Black basal dash thin and inconspicuous in most specimens. Hind wing of male pale buff basally with
smoky-brown shading on outer quarter of wing. Hind wing of female similar to that of male, or smoky-brown to wing base.
Expanse: 3 1 .9 ± 1 .3 mm. Length of forewing: 1 5.9 ± 0.6 mm ( 1 00 specimens).
Male genitalia (Fig. 75). Distinguishable from those of E. agema and E. oblongistigma by longer sacculus extensions.
Harpe pubescent.
Length of right sacculus extension: 1 .57 ± 0.07 mm (25 specimens)
Length of left sacculus extension: 1 .43 ± 0.08 mm (25 specimens).
Length of right harpe: 1.03 ±0.04 mm (25 specimens).
Length of right sacculus: 1 .30 ±0.05 (25 specimens).
Female genitalia (Figs. 106, 137). Indistinguishable from those of E. oblongistigma and E. agema females. Those of
these three species may be distinguished from those of females of other species in the E. detersa group by the rounded or
pear-shaped rather than elongate corpus bursae.
Type material. - The holotype of Agrotis olivalis Grote is a female in the collection of the
British Museum (Natural History). The specimen has been dissected.
The holotype of Euxoa mcdunnoughi Cook is a male in the Montana Experiment Station
collection, Bozeman.
Distribution and period of flight. - Euxoa olivalis occurs in western North America from
southern Saskatchewan westward to southern British Columbia and southward to northern
New Mexico, Arizona and southern California (Fig. 165). Approximately 1 100 specimens from
98 localities were examined.
Specimens have been collected from late June until early September; the flight period is
later, on the average, in the south than it is in the north.
Habitat. - Euxoa olivalis occurs in a variety of habitats. In the Great Basin it occurs in
pinon-juniper woodland; in montane areas it occurs in open ponderosa pine ( Pinus ponderosa
Quaest. Ent., 1981, 17 (1,2)
28
Lafontaine
Dougl.) and lodgepole pine ( Pinus contorta Dougl.) parkland; in the Great Plains it has been
collected in sagebrush areas. In general, E. olivalis is found in habitats intermediate between
xeric habitats of E. oblongistigma and mesic ones of E. agema.
Remarks. - Most specimens of E. olivalis are distinguished from those of E. oblongistigma
and E. agema by characters in the key. Specimens from populations of E. olivalis from more
mesic, higher elevation, localities are darker and have more brown coloration than do specimens
from more xeric, lower elevation, localities and are similar in color and pattern to E. agema
specimens. Although males of the two species are distinguished by characters of the genitalia,
some females from more mesic localities are identified by association with males only.
Euxoa agema (Strecker)
Figs. 28,29, 76, 138, 166
Agrotis agema Strecker, 1899, p. 5; Colorado.
Paragrotis agema; Dyar, 1902, p. 140.
Euxoa agema; Hampson. 1903, p. 666.
Euxoa agema; Barnes and McDunnough, 1917, p. 41; = E. oblongistigma.
Description. — Head, thorax, and pattern of forewing similar to that of E. olivalis specimens. Ground color of
forewing dark brown longitudinally streaked with light brown; forewing darker and without gray streaks of most specimens
of E. olivalis. Hind wing smoky-brown, slightly paler toward base.
Expanse; 32.4 ± 1.2 mm. Length of forewing: 1 5.8 ± 0.7 mm (100 specimens).
Male genitalia (Fig. 76). Distinguishable from those of E. olivalis male by shorter sacculus extensions. In male of E.
agema sacculus extensions appear to be about as long as harpes; in male of E. olivalis they are markedly longer than
harpes. Male genitalia of E. agema indistinguishable from those of E. oblongistigma male.
Length of right sacculus extension: 1.19 ±0.06 mm (25 specimens).
Length of left sacculus extension: 1 .08 ± 0.07 mm (25 specimens).
Length of right harpe: 1.1 3 ±0.06 mm (25 specimens).
Length of right sacculus: 1 .26 ± 0.05 mm (25 specimens).
Female genitalia (Figs. 107, 138). Indistinguishable from those of E. olivalis , and E. oblongistigma females.
Type material. - The holotype of Agrotis agema Strecker is a female in the collection of the
Field Museum of Natural History, Chicago.
Distribution and period of flight. - Euxoa agema occurs in mountain ranges that surround
the Great Basin and Colorado Plateau (Fig. 166). Within the Great Basin it has been collected
only in the Ruby Mountains, Nevada. Approximately 350 specimens from 38 localities have
been examined.
Adults have been collected from mid-July until early September.
Habitat. - This species inhabits forested areas of spruce, aspen, and lodgepole pine ( Pinus
contorta Dough). Although specimens of E. agema are sometimes collected with those of E.
olivalis and E. oblongistigma , they are generally found in more mesic habitats than are those
of the other two species.
Remarks. - Specimens of E. agema may be distinguished from those of E. olivalis and E.
oblongistigma by characters in key (also see remarks under E. olivalis ).
Euxoa oblongistigma (Smith)
Figs. 30,31,32, 77, 108, 139, 167
Agrotis oblongistigma Smith, 1887, p. 454; Montana, Black Hills, South Dakota.
Carneades oblongistigma; Smith, 1890b, p. 140; redescription.
Carneades oblongistigma; Smith, 1893, p. 89; catalogue.
Paragrotis oblongistigma; Dyar, 1902, p. 140.
Euxoa oblongistigma; Hampson, 1903, p. 210.
The Euxoa detersa group (Lepidoptera: Noctuidae)
29
Euxoa oblongistigma', Draudt, 1924, p. 38; redescription.
Description. — Vestiture of head and thorax brown and black scales; transverse band of white scales posterior to
prothoracic collar in some specimens. Ground color of forewing buff or pale gray streaked longitudinally with dark brown
and black. Transverse lines essentially absent. Terminal area slightly darker than remainder of forewing. Series of black,
sagittate spots on proximal side of subterminal line in most specimens. Costa pale, evenly colored, without cross-lines; pale
color of costa extended to cubital vein near wing base, not separated from it by black line. Reniform and orbicular spots
pale, without white line between center of spot and black outline. Orbicular spot elongate and rectangular or slightly oval,
about twice as long as wide. Black basal dash prominent in most specimens. Hind wing of male pale buff, darker
smoky-brown terminal band present in many specimens. Hind wing of female similar to that of male but smoky-brown
shading more extensive.
Expanse: 32.0 ± 1 .2 mm. Length of forewing: 1 5.7 ± 0.8 mm ( 1 00 specimens).
Male genitalia (Fig. 77). Indistinguishable from those of E. agema male. Distinguishable from those of E. olivalis
male by shorter sacculus extensions.
Length of right sacculus extension: 1 .32 ± 0.07 mm (25 specimens).
Length of left sacculus extension; 1 . 1 9 ± 0. 1 0 mm (25 specimens).
Length of right harpe: 1 . 1 5 ± 0.06 mm (25 specimens).
Length of righ sacculus: 1 .28 ± 0.07 mm (25 specimens).
Female genitalia ( Figs. 1 08, 139). I ndistinguishable from those of E. olivalis and E. agema females.
Type material. - Agrotis oblongistigma was described by Smith from four female
specimens from Montana and South Dakota, of which two have been located, in the collection
of the United States National Museum. One of these will be selected as lectotype by Todd
(1982).
Distribution and period of flight. - Euxoa oblongistigma occurs in the western Great Plains
from southern Saskatchewan and Alberta southward to northern Colorado, and in the
intermontane region from southern British Columbia southward to northern New Mexico,
southern Utah, central Nevada, and central California (Fig. 167). Approximately 700
specimens from 75 localities were examined.
Habitat. - Euxoa oblongistigma inhabits dry, sagebrush areas and open conifer woods
where sagebrush is plentiful.
Remarks. - Specimens of E. oblongistigma are distinguished from those of E. olivalis and
E. agema by characters in the key. Generally, the pale, evenly colored costa, reniform spot and
orbicular spot allow specimens of E. oblongistigma to be readily distinguished from those of the
other two species.
Euxoa citricolor (Grote)
Figs. 33, 34, 35, 78, 109, 140, 169
Agrotis citricolor Grote, 1 880b, p. 1 54; Colorado.
Carncades citricolor ; Grote, 1883, p. 26.
Carncades citricolor ; Smith, 1890b, p. 155; redescription.
Carneades citricolor. Smith, 1893, p. 92; catalogue.
Paragrotis citricolor, Dyar, 1 902, p. 1 42.
Euxoa citricolor, Draudt, 1924, p. 39; redescription.
Euxoa citricolor ab. postmcdialis Strand, 1915, p. 144; Prescott, Arizona; infrasubspecific name.
Euxoa citricolor form postmcdialis Draudt, 1924, p. 39; validation of name.
Euxoa citricolor ab. postmcdialis ; McDunnough, 1938; checklist.
Description. — Vestiture of head and thorax yellow. Ground color of forewing yellow, overlaid with dusting of
brown scales in many specimens. Transverse lines obscure or absent. Terminal space, reniform spot, and in many
specimens orbicular spot, dark gray-brown, contrasting. Hind wing white in most specimens, a few specimens with some
dark shading on outer third of wing or along veins.
Expanse: 35.3 ± 1.3 mm. Length of forewing: 16.8 ±0.8 mm (100 specimens).
Male genitalia (Fig. 78). Distinguishable from those of E. tronclla by shape of harpe and sacculus extension, and by
smaller sub-basal diverticulum of vesica. In specimens of E. citricolor, harpe and sacculus not markedly curved near base
but appear slightly squeezed together; in specimens of E. tronella, harpe and sacculus extension markedly curved near
base, an even C-shape, or ‘U’, where they meet.
Quaest. Ent., 1981, 17 (1,2)
30
Lafontaine
Length of right sacculus extension: 1.78 ±0.10 mm (22 specimens).
Length of left sacculus extension: 1.78 ±0.11 mm (22 specimens).
Length of right harpe: 1 .32 ± 0.06 mm (22 specimens).
Length of right sacculus: 1.48 ±0.07 mm (22 specimens).
Female genitalia (Figs. 109, 140). Indistinguishable from those of E. tronella female. The genitalia of these two
species distinguishable from those of other species in E. detersa group by characters in key.
Type material. - Grote described A. citricolor on the basis of two specimens, a male and a
female. Only the male has been located, this in the collection of the British Museum (Natural
History). This specimen is here designated lectotype. The specimen, a male in good condition, is
labelled “Colorado, Grote Coll., 81-116”, “type”. A genitalic slide has been prepared (No.
6330). The holotype of E. postmedialis Draudt is in the collection of the British Museum
(Natural History). The specimen, a female from Prescott, Arizona, has been dissected.
Distribution and period of flight. - Euxoa citricolor occurs from southern Washington
southward to southern California and eastward to central Wyoming, Colorado and New
Mexico. It has also been collected in the northern Great Plains at Watford City, North Dakota,
at Mitchell, South Dakota and at Dinosaur Provincial Park, Alberta (see figure 169).
Approximately 400 specimens from 72 localities were examined.
Specimens have been collected from late August until early October.
Habitat. - This species inhabits open aridlands where the flora is dominated by big
sagebrush ( Artemisia tridentata Nutt.). It is most common in areas where the soil is loose and
coarse. In the Great Plains it has been collected in areas of extensive erosion such as badlands
and river terraces.
Remarks. - Specimens of E. citricolor may be distinguished from those of E. tronella by the
color of the forewing, as well as by differences in male genitalia given above. In specimens of E.
citricolor the forewing is yellow; in those of E. tronella , the forewing is cream-colored, or pale
buff, with a dusting of darker scales.
Euxoa tronella (Smith)
Figs. 36, 37,38, 79, 110, 141, 170
Carneades tronellus Smith, 1903a, p. 1 1; Stockton, Utah.
Euxoa tronella ; Hampson, 1903, p. 432.
Euxoa tronellus\ Barnes and McDunnough, 1917, p. 41; = E. citricolor.
Euxoa tronellus ; Draudt, 1924, p. 39; = E. citricolor.
Carneades tronellus ; Rindge, 1955, p. 134; type material.
Description. — Vestiture of head and thorax cream-colored or pale, buffy gray. Ground color of forewing
cream-colored or pale buff, dusted with darker scales in most specimens. Transverse lines obscure or absent. Terminal area
gray, darker than ground color in most specimens. Reniform and orbicular spots partially filled with gray shading or
concolorous with ground color. Hind wing white in most specimens; smoky-brown shading on terminal margin of wing and
on median line in some specimens.
Some geographical variation evident. Specimens from Mojave Desert, California, cream-colored, without darker scales
or maculation. Specimens from Saskatchewan and North Dakota buff-colored, heavily dusted with dark scales; hind wing
with terminal half shaded with smoky-brown. Specimens from Great Basin and Wyoming fall in between these extremes in
an east-west dine.
Expanse: 34.0 ± 1 .7 mm. Length of forewing: i 6.0 ± 0.6 mm ( 1 00 specimens).
Male genitalia (Fig. 79). Distinguishable from those of E. citricolor male by shape of harpe and sacculus extension,
and by larger sub-basal diverticulum of vesica. In male of E. tronella , harpe and sacculus extension markedly curved near
base, an even C-shape, or ‘U’ formed where they meet; in male of E. citricolor , harpe and sacculus extension not markedly
curved near base but appear slightly squeezed together.
Length of right sacculus extension: 1 .6 1 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1 .60 ± 0.09 mm (20 specimens).
Length of right harpe: 1 ,30±0.07 mm (20 specimens).
Length of right sacculus: 1 .37 ±0.07 mm (20 specimens).
The Euxoa detersa group (Lepidoptera: Noctuidae)
31
Female genitalia (Figs. I 10, 141). Indistinguishable from those of E. citricolor female.
Type material. - Carneades tronellus was described by Smith on the basis of six specimens
from Stockton, Utah. Of these six, only three specimens have been located, a male in the
collection of the American Museum of Natural History, and a male and female in the
collection of the United States National Museum. A lectotype will be designated by Todd
(1982).
Distribution and period of flight. - Euxoa tronella occurs in the western Great Plains from
southern Saskatchewan and Alberta southward to western South Dakota and southern
Wyoming, and in the intermontane region from southern Washington southward to northern
New Mexico and southern California (Fig. 170). Approximately 600 specimens from 53
localities were examined.
Specimens have been collected from late August until early October.
Habitat. - Euxoa tronella inhabits open aridlands where big sagebrush ( Artemisia
tridentata Nutt.) is abundant. This species is apparently similar to E. citricolor in habitat
requirements in the intermontane region; specimens of the two species are frequently collected
together. In the Great Plains, E. tronella is not restricted to areas of extensive erosion as is E.
citricolor.
Remarks. - Specimens of E. tronella may be distinguished from those of E. citricolor by
forewing color, and by differences in male genitalia as given under E. citricolor.
Euxoa teleboa (Smith)
Figs. 39, 40, 80, 111, 142, 171
Carneades teleboa Smith, 1890b, p. 219; Las Vegas, New Mexico.
Carneades teleboa ; Smith, 1893, p. 99; catalogue.
Paragrotis teleboa-, Dyar, 1902, p. 145.
Euxoa teleboa ; Flampson, 1903, p. 261.
Euxoa teleboa-, Draudt, 1924, p. 45; redescription.
Carneades pedalis Smith, 1890b, p. 220; Colorado.
Carneades pedalis-, Smith, 1893, p.98; catalogue.
Paragrotis pedalis-, Dyar, 1902, p. 145.
Euxoa pedalis-, Hampson, 1903, p. 261.
Euxoa pedalis ; Draudt, 1924, p. 45; redescription.
Euxoa pedalis\ Lafontaine, 1982, = E. teleboa.
Carneades recticincta Smith, 1895, p. 334; Calgary, Alberta.
Paragrotis recticincta-, Dyar, 1902, p. 145.
Euxoa recticincta-, Hampson, 1903, p. 261.
Euxoa recticincta-, Draudt, 1924, p. 45; redescription.
Euxoa recticincta-, Lafontaine, 1982; = E. teleboa.
Description. — Vestiture of head and thorax, and ground color of forewing reddish orange or cream-colored.
Transverse lines, except median line, obscure or absent. Median line black, prominent. Terminal area and distal portion of
subterminal area dusted with black scales. Reniform spot dark-filled, forming part of dark median line in most specimens.
Hind wing of male white; hind wing of female white or dusted with gray-brown scales.
Expanse: 3 1 .6 ± 1.5 mm. Length of forewing: 1 5.7 ± 0.8 mm. ( 1 00 specimens).
Male genitalia (Fig. 80). Sacculus extension short, about as long as harpe in most specimens. Vesica with small
nipple-like sub-basal diverticulum in Addition to normal sub-basal diverticulum.
Length of right sacculus extension: 1 . 1 7 ± 0. 1 1 mm (20 specimens).
Length of left sacculus extension: 1 .07 ±0.13 mm (20 specimens).
Length of right harpe: 1 .25 ±0.06 mm (20 specimens).
Length of right sacculus: 1 .29 ± 0.07 mm (20 specimens).
Female genitalia (Figs. Ill, 142). Sclerotized plate in ventral wall of ductus bursae extended as far anteriorly as plate
in dorsal wall. Sclerotized flange-like projection on ovipositor valve rectangular, fin-like plate on posterior half of dorsal
margin of valve. Flange-like projection pointed at posterior end, convex on dorsal margin and tapered abruptly to valve at
anterior end.
Quaest. Ent., 1981, 17 (1,2)
32
Lafontaine
Type material. - The holotype of Carneades teleboa is a female in the collection of the
United States National Museum. The specimen has been dissected.
The holotype of Carneades pedalis Smith is a male in the collection of the United States
National Museum. The specimen is without an abdomen.
The holotype of Carneades recticincta Smith is a female in the collection of the United
States National Museum. The specimen has been dissected.
Distribution and period of flight. - Euxoa teleboa occurs in the western Great Plains and
eastern intermontane region from southern Saskatchewan and Alberta southward to northern
Texas, central New Mexico, northern Arizona an east-central Nevada. A worn specimen in the
Canadian National Collection is labelled Peachland, B.C., 26 July, 1912, J.B. Wallis .
Locality and collecting date both suggest that this specimen is mislabelled. Approximately 200
specimens from 42 localities were examined
Adults have been collected from mid-August until late September.
Habitat. - In the intermontane region, and in the Rocky Mountain region, Euxoa teleboa
occurs in dry pinon-juniper woodland. In the Great Plains it inhabits arid areas in which the
flora is dominated by sages ( Artemisia spp.).
Remarks. - Specimens of E. teleboa are most similar to those of E. tronella in color and
markings, however, they may be distinguished readily by the broad, dark median line present in
specimens of E. teleboa. Structurally, males of E. teleboa may be distinguished from those of
E. tronella by the shorter sacculus extension and by the additional sub-basal diverticulum of
the vesica. Shape of female ovipositor valve and shape of sclerotized, flange-like projection on
valve may be used to distinguish females of the two species.
Euxoa moerens (Grote)
Figs. 41, 42, 43,81, 112, 143, 172
Carneades moerens Grote, 1 883, p. 4; Arizona.
Carneades moerens ; Smith, 1890b, p. 156; redescription.
Carneades moerens'. Smith, 1893, p. 95; catalogue.
Paragrotis moerens ; Dyar, 1902, p. 142.
Euxoa moerens ; Hampson, 1903, p. 224.
Euxoa moerens', Draudt, 1924, p. 39; redescription.
Agrotis luteola Smith, 1887, p. 457; Arizona.
Carneades luteola'. Smith, 1890b, p. 160; redescription.
Carneades luteola'. Smith, 1893, p. 94; catalogue.
Paragrotis luteola', Dyar, 1902, p. 143.
Euxoa luteola', Hampson, 1903, p. 229.
Euxoa luteola ; Barnes and McDunnough, 1 9 1 7, p. 4 1 ; = E. moerens.
Description. — Antenna of male slightly biserrate. Vestiture of head and thorax brownish gray. Ground color of
forewing pale gray, pale brown, or orange-brown, dusted with darker scales. Transverse lines indistinct. Terminal area
gray-brown, slightly darker than ground color of forewing. Reniform and orbicular spots concolorous with ground color in
most specimens, outlined in black. Hind wing buff with dusting of pale smoky-brown scales on outer third of wing and in
many specimens on median line as well. Hind wing smoky-brown, slightly paler near base, in some specimens.
Expanse: 29.8 ± 1 .2 mm. Length of forewing: 1 4.6 ± 0.7 mm ( 1 00 specimens).
Male genitalia (Fig. 81). Uncus with stout setae mesially, brush-like in appearance. Harpe curved dorsally near apex;
harpe with about 30 scattered setae. Cucullus strap-like, not markedly constricted subapically Vesica with small
nipple-like sub-basal diverticulum in addition to normal foot-like sub-basal diverticulum.
Length of right sacculus extension: 1 .53 ± 0.07 mm (20 specimens).
Length of left sacculus extension: 1 .52 ± 0.07 mm (20 specimens).
Length of right harpe: 1.06 ± 0.04 mm (20 specimens).
Length of right sacculus: 1 .23 ± 0.06 mm (20 specimens).
Female genitalia (Figs. 1 12, 143). Corpus bursae oval; ductus bursae entering corpus bursae about one-quarter from
posterior end. Sclerotized plate in dorsal wall of ductus bursae extended farther anteriorly than plate in ventral wall.
The Euxoa detersa group (Lepidoptera: Noctuidae)
33
Sclerotized flange-like projection on ovipositor valve elongate, fin-like, tapered abruptly to valve at anterior end in many
specimens.
Type material. - The holotype of Carneades moerens Grote is a male in the collection of the
United States National Museum. The specimen has been dissected.
The holotype of Agrotis luteola Smith is a female in the collection of the United States
National Museum. The specimen has been dissected.
Distribution and period of flight. - Euxoa moerens occurs in the western Great Plains from
southern Saskatchewan and Alberta southward to central Colorado, and in the intermontane
region from southern Idaho and central Oregon southward to central Colorado, northern
Arizona and central Nevada. Approximately 250 specimens from 37 localities were examined.
Specimens have been collected from mid-August until late September.
Habitat. - Euxoa moerens inhabits open pinon-juniper woodland where sagebrush is
abundant. It also occurs in mixed conifer forests of pinon pine ( Pinus spp.), Utah juniper
(Juniperus osteosperma (Torr.) Little) and ponderosa pine ( Pinus ponderosa Dougl.). In the
Great Plains region it occurs in open aridlands where sage ( Artemisia spp.) is abundant.
Remarks. - Specimens of Euxoa moerens are most similar to those of E. latro in wing
markings and structural characters. Specimens of the two species may be distinguished by
characters in key.
Euxoa latro (Barnes and Benjamin)
Figs. 44, 45,82, 113, 144, 173
Agrotis latro Barnes and Benjamin, 1926, p. 305; Truckee, California.
Euxoa latro ; McDunnough, 1938, p. 58.
Description. — Antenna of male slightly biserrate, Vestiture of head and thorax mixture of gray, brown and black
scales. Ground color of forewing brown or gray-brown markedly dusted with black. Transverse lines present; median line
conspicuous in most specimens. Orbicular spot concolorous with ground color. Reniform spot pale, cream-colored or pale
buff in many specimens. Hind wing smoky-brown.
Expanse: 30.9 ± 1 .3 mm. Length of forewing: 1 5.3 ± 0.8 mm (80 specimens).
Male genitalia (Fig. 82). Uncus with stout setae mesially, brush-like in appearance. Harpe evenly incurved throughout
its length, not curved dorsally near apex. Harpe bristly in appearance, its surface covered with 50-60 setae. Cucullus
bat-like, markedly constricted subapically. Vesica only with normal, foot-like sub-basal diverticulum in most specimens,
trace of extra nipple-like diverticulum in some specimens.
Length of right sacculus extension: 1 .55 ± 0.09 mm (20 specimens).
Length of left sacculus extension: 1 .52 ± 0.09 mm (20 specimens).
Length of right harpe: 0.96 ± 0.05 mm (20 specimens).
Length of right sacculus: 1 .34 ± 0.06 mm (20 specimens).
Female genitalia (Figs. 1 13, 144). Indistinguishable from those of E. moerens female.
Type material. - The holotype of Agrotis latro Barnes and Benjamin is a male in the
collection of the United States National Museum. The specimen has been dissected.
Distribution and period of flight. - Euxoa latro occurs in the Cascades and Sierra Nevada
Mountain systems from central Washington southward to southern California. A disjunct
population occurs in the La Sal Mountains, Utah (Fig. 173). Approximately 200 specimens
from 18 localities were examined.
Most specimens were collected between mid- August and mid-September. One specimen was
collected in mid-October.
Habitat. - Euxoa latro inhabits conifer forests of pine and fir.
Remarks. - Specimens of Euxoa latro may be distinguished from those of E. moerens by
characters in key. In general, the much darker coloration of Euxoa latro specimens allow them
to be readily distinguished from E. moerens specimens. Although the two species rarely occur
Quaest. Ent., 1981, 17 (1,2)
34
Lafontaine
together, they have been collected together in the Sierra Nevadas where the ponderosa pine
zone meets the pinon-juniper zone.
Euxoa murdocki (Smith)
Figs. 46,83, 114, 145, 174
Agrotis murdocki Smith, 1890a, p. 49; British Columbia, Utah.
Carneades murdocki ; Smith, 1890b, p. 174; redescription.
Carneades murdocki\ Smith, 1893, p. 99; catalogue.
Paragrotis murdocki’, Dyar, 1902, p. 145.
Euxoa murdocki ; Hampson, 1903, p. 247.
Euxoa murdocki ; Draudt, 1924, p. 45; redescription.
Description. — Antenna of male slightly biserrate. Vestiture of head and thorax orange-brown or yellow-brown.
Ground color of forewing orange in basal and subterminal areas, gray in median and terminal areas. Transverse lines
prominent ; median line dark gray. Reniform and orbicular spots partially outlined in black, inconspicuous. Reniform spot
dark gray, concolorous with median line. Orbicular spot pale gray, concolorous with median area Hind wing smoky-brown,
paler near base in most specimens.
Expanse: 29.0 ± 1 .5 mm. Length of forewing; 1 4. 1 ± 0.7 mm ( 1 00 specimens).
Male genitalia (Fig. 83). Similar to those of E. moerens male but sacculus larger and vesica with normal, foot-like
sub-basal diverticulum only.
Length of right sacculus extension: 1 .48 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1 .48 ± 0.09 mm (20 specimens).
Length of right harpe: 1 .05 ± 0.05 mm (20 specimens).
Length of right sacculus: 1 .38 ± 0.05 mm (20 specimens).
Female genitalia (Figs. 1 14, 145). Indistinguishable from those of E. moerens and E. latro females.
Type material. - Agrotis murdocki Smith was described from a male from British
Columbia and a female from Utah. Both specimens are in the collection of the United States
National Museum. A lectotype will be designated by Todd (1982).
Distribution and period of flight. - Euxoa murdocki occurs from south-central British
Columbia southward to west-central Colorado, central Nevada and southern California (Fig.
174). I have seen only one specimen collected east of the Continental Divide, this from Banff,
Alberta. Approximately 250 specimens from 41 localities were examined.
Specimens have been collected from late August until mid-October.
Habitat. - Euxoa murdocki is found most commonly in conifer forests of Douglas-fir
( Pseudotsuga menziesii (Mirbel) Franco) and ponderosa pine ( Pinus ponderosa Dough). In
the Great Basin it occurs in pinon-juniper woodland.
Remarks. - Although Euxoa murdocki specimens are structurally similar to those of E.
moerens and E. latro , they are unlike those of any other species in the genus in wing color. The
combination of orange basal and subterminal areas and gray median and terminal areas make
this species one of the easiest to recognize in Euxoa.
Euxoa dodi McDunnough
Figs. 47, 84, 115, 146, 175
Euxoa dodi McDunnough, 1923, p. 163; Lethbridge, Alberta.
Euxoa dodi ; Draudt, 1924, p. 43; redescription.
Description. — Antennae of male biserrate. Vestiture of head and thorax buff or brown. Ground color of forewing
brown or pale gray-brown. Transverse lines all prominent except median line. Basal and subterminal areas slightly paler
than median area. Terminal area dark gray-brown. Reniform orbicular and claviform spots outlined in black, concolorous
with basal and subterminal areas. Hind wing pale smoky-brown, slightly darker on outer quarter of wing.
Expanse: 30.3 ± 1 .2 mm. Length of forewing: 1 4.9 ± 0.7 mm (80 specimens).
Male genitalia (Fig. 84). Uncus with long, thin setae mesially. Harpe pubescent, in addition to longer scattered setae.
Right sacculus extension less than 1.50 times length of right harpe. Vesica without extra nipple-like sub-basal
The Euxoa detersa group (Lepidoptera: Noctuidae)
35
diverticulum.
Length of right sacculus extension: 1.59 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1.45 ±0.12 mm (20 specimens).
Length of right harpe: 1.17 ±0.05 mm (20 specimens).
Length of right sacculus: 1.31 ±0.06 mm (20 specimens).
Female genitalia (Figs. 1 15, 146). Ductus bursae and corpus bursae similar to those of E. moerens , E. latro and E.
murdocki females. Ovipositor valve clothed with fine setae, without sclerotized projection at apex.
Type material. - The holotype of Euxoa dodi McDunnough is a male (type No. 608) in the
Canadian National Collection, Ottawa. The specimen has been dissected.
Distribution and period of flight. - Euxoa dodi occurs from southern Saskatchewan and
Alberta southward to southern North Dakota, southern New Mexico and northern Arizona
(Fig. 175). Approximately 110 specimens from 36 localities were examined.
Specimens have been collected from mid-August until early September.
Habitat. - Euxoa dodi occurs in areas of dry, shortgrass prairie where sage ( Artemisia spp.)
is abundant. It also occurs in the Rocky Mountain foothills where prairie habitat occurs in open
pine parkland.
Remarks. - Specimens of E. dodi may be distinguished from those of E. infracta by paler
coloration and lack of copper suffusion of the forewing and by the male genitalia. The sacculus
extension of E. dodi male is less than 1.50 times length of harpe; it is more than 1.50 times
length of harpe in E. infracta male. In general, Euxoa dodi occurs in more open xeric habitats
than does E. infracta. Specimens of E. dodi from central Colorado are darker than are those
from the Great Plains. At present it is not known whether E. dodi and E. infracta hybridize in
this region or if these specimens represent a dark form of E. dodi.
Euxoa infracta (Morrison)
Figs. 48,85, 116, 147, 176
Agrotis infracta Morrison, 1875a, p. 115; Colorado, Texas.
Carneades infracta ; Smith, 1890b, p. 160; redescription.
Carneades infracta ; Smith, 1 893, p. 94; catalogue.
Paragrotis infracta ; Dyar, 1902, p. 142.
Euxoa infracta ; Hampson, 1903, p. 228.
Euxoa infracta ; Draudt, 1924, p. 40; redescription.
Description. — Antenna of male biserrate. Vestiture of head and thorax dark blackish brown. Ground color of
forewing brown; basal and subterminal areas brown with a copper suffusion, median and terminal areas dark blackish
brown. Transverse lines all prominent except median line. Reniform, orbicular and claviform spots outlined in black,
slightly paler than median area. Hind wing smoky-brown, slightly paler near base in many specimens.
Expanse: 3 1 .4 ± 1.3 mm. Length of forewing: 1 5.7 ± 0.8 mm ( 1 00 specimens).
Male genitalia (Fig. 85). Similar to those of E. dodi male but sacculus extension longer and harpe shorter in E.
infracta male. Right sacculus extension longer than 1.50 times length of right harpe.
Length of right sacculus extension: 1 .62 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1 .60 ±0.10 mm (20 specimens).
Length of right harpe: 0.99 ±0.06 mm (20 specimens).
Length of right sacculus: 1.36 ±0.07 mm (20 specimens).
Female genitalia (Figs. 1 16, 147). Indistinguishable from those of E. dodi female.
Type material. - Agrotis infracta Morrison was described from two specimens, one from
Colorado and one from Texas. Only the specimen from Colorado has been located. This
specimen, a female in good condition except for missing antenna, is labelled “Col.”, “type”,
“ Carneades infracta Morr. 1782”, “ Euxoa slide MSU 3”. This specimen, in the collection of
! Michigan State University, East Lansing, is here designated lectotype.
Distribution and period of flight. - Euxoa infracta occurs from southern Manitoba
westward to British Columbia and southward to northern New Mexico, east-central Arizona,
Quaest. Ent., 1981, 17 (1,2)
36
Lafontaine
central Nevada and east-central California (Fig. 176). Approximately 450 specimens from 97
localities were examined.
Specimens have been collected from mid-July until late September.
Habitat. - Euxoa infracta is found most commonly in conifer forests of pine and fir. In the
intermontane region it also occurs in pine and aspen forests.
Remarks. - Specimens of Euxoa infracta may be distinguished from those of E. dodi by
characters in the key (see also remarks under E. dodi).
Euxoa laetificans (Smith)
Figs. 49, 50, 86, 117, 150, 177
Carneades laetificans Smith, 1894, p. 48; Glenwood Springs, Colorado.
Paragrotis laetificans ; Dyar, 1 902, p. 1 40.
Euxoa laetificans ; Hampson, 1903, p. 208.
Euxoa laetificans ; Draudt, 1924, p. 38; redescription.
Carneades masculinus Smith, 1903b, p. 6; Silver Bow Co., Montana; Yakima, Washington; southern Utah.
Euxoa masculinus ; Barnes and McDunnough, 1917, p. 41; = E. laetificans.
Carneades masculinus-, Rindge, 1955, p. 120; type material.
Description. — Sexually dimorphic. Vestiture of head gray; vestiture of prothoracic collar reddish brown (male) or
gray (female); vestiture of thorax gray or reddish gray. Ground color of forewing light reddish brown (male) or gray
(female). Transverse lines absent in most specimens, trace of lines in some specimens. Median area around reniform and
orbicular spots shaded with black. Terminal area dark gray, streaked into subterminal area. Costa and cubital vein paler
than remainder of forewing, yellow-brown (male) or silver-gray (female). Cubital vein pale from wing base to reniform
spot. Reniform and orbicular spots with pale line between black outline and gray center; orbicular spot fused with costal
shade in some specimens. Basal dash and claviform spot black, prominent. Hind wing of male white or pale buff with some
smoky-brown shading on terminal margin. Hind wing of female smoky-brown to base.
Expanse: 30.6 ±1.5 mm. Length of forewing: 1 4.5 ± 0.7 mm ( 1 00 specimens).
Male genitalia (Fig. 86). Sacculus extensions and harpes long; harpe pubescent. Vesica without nipple-like sub-basal
diverticulum, only foot-like diverticulum sub-basally. Median diverticulum with basal bulge in many specimens.
Length of right sacculus extension: 1 .7 1 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1 .65 ±0.12 mm (20 specimens).
Length of right harpe: 1.45 ±0.05 mm (20 specimens).
Length of right sacculus: 1.45 ±0.05 mm (20 specimens).
Female genitalia (Figs. 1 17, 150). Corpus bursae oval or rectangular enlarged posterolaterally on left side. Ovipositor
valve clothed with short, fine setae, without sclerotized projection at apex. Valve triangular, pointed toward apex.
Type material. - Carneades laetificans was described by Smith from two specimens, a male
and a female from Colorado. Both specimens are in the collection of the United States National
Museum. A lectotype will be selected by Todd (1982).
Carneades masculinus Smith was described from seven males from Montana, Washington
and Utah. Type material is in the collections of the American Museum of Natural History and
the United States National Museum. A lectytype will be designated by Todd (1982).
Distribution and period of flight. - Euxoa laetificans occurs in western North America
from southern Saskatchewan westward to southern British Columbia and southward to
southern Colorado, Utah, Nevada, and east-central California (Fig. 177). Approximately 700
specimens from 71 localities were examined.
Specimens have been collected from late July until mid-September.
Habitat. - Euxoa laetificans is found most commonly in open pine forests of ponderosa and
lodgepole pine ( Pinus ponderosa Dougl. and P. contorta Dough). It is collected in areas of
coarse, gravelly soil. In the intermontane region E. laetificans occurs in pinon-juniper woodland
as well as in pine forests.
Remarks. - Specimens of Euxoa laetificans are similar to those of E. quadridentata but
may be distinguished from them by lack of pale-lined veins in subterminal area and by
The Euxoa detersa group (Lepidoptera: Noctuidae)
37
characters of genitalia. The harpe of E. laetificans male is pubescent and harpe and sacculus
extensions are much longer than are those of E. quadridentata male. Ovipositor valve of E.
laetificans female is narrow and pointed apically; that of E. quaridentata female is broad and
rounded apically.
Larvae have been reared on alfalfa ( Medicago sativa L.), sweet clover ( Melilotus spp.),
beet {Beta vulgaris L.), lettuce {Lactuca sativa L.) and Russian thistle {Salsola kali L.)
(Cook, 1930b).
Euxoa quadridentata (Grote and Robinson)
Figs. 51, 52, 87, 118, 148, 149, 178
Agrotis quadridentata Grote and Robinson, 1865, p. 491; Colorado.
Carneades quadridentata. Smith, 1890b, p. 139; redescription.
Carneades quadridentata ; Smith, 1893, p. 89; catalogue.
Paragrotis quadridentata ; Dyar, 1 902, p. 1 40.
Euxoa quadridentata', Hampson, 1903, p. 209.
Euxoa quadridentata', Draudt, 1924, p. 38; redescription.
Carneades pugionis Smith, 1900, p. 419; Denver, Colorado; Calgary, Alberta.
Paragrotis pugionis', Dyar, 1902, p. 141.
Euxoa pugionis', Hampson, 1903, p. 209.
Euxoa pugionis', Barnes and McDunnough, 1917, p. 41; = E. quadridentata.
Carneades pugionis', Rindge, 1955, p. 127; type material.
Euxoa flutea Smith, 1910b, p. 255; Sierra Nevada, California.
Euxoa quadridentata flutea ; Barnes and McDunnough, 1917, p. 41.
Euxoa quadridentata flutea-, Draudt, 1924, p. 38; redescription.
Euxoa flutea ; Rindge, 1955, p. 127; type material.
Description. — Sexually dimorphic. Vestiture of head gray; upper half of prothoracic collar gray-brown, basal half
yellow (male) or pale gray (female). Vestiture of thorax mixture of brown, black and gray scales. Ground color of forewing
dark brown streaked with yellow (male) or silver-gray (female). Transverse lines indistinct. Veins Cuj and M3 pale-lined
with silver-gray in both sexes and projected into terminal area; cubital vein silver-gray. In male, costal area, posterior
margin of wing, and area distal to claviform spot pale yellow or yellow-buff; subterminal area streaked with yellow or pale
buff. In female, pale yellow shading replaced by silver-gray. Terminal area dark gray-brown, streaked into subterminal
area; apices of streaks in subterminal area black. Reniform and orbicular spots filled with yellow-buff, or with yellow-buff
between gray center and black outline. Black basal dash present, inconspicuous in some specimens. Claviform spot black.
Hind wing in most males white basally with smoky-brown shading on outer third of wing. Hind wing of female and of male
from Sierra Nevada populations dark smoky-brown.
Expanse: 30.7 ±1.1 mm. Length of forewing: 1 5.2 ± 0.6 mm ( 1 00 specimens).
Male genitalia (Fig. 87). Harpe and sacculus extension much shorter than those of E. laetificans male; harpe not
pubescent. Vesica without extra nipple-like sub-basal diverticulum.
Length of right sacculus extension: 1 .22 ±0.13 mm (24 specimens).
Length of left sacculus extension: 1 . 1 8 ± 0. 1 3 mm (24 specimens).
Length of right harpe: 1 . 1 8 ± 0.05 mm (24 specimens).
Length of right sacculus: 1 .35 ± 0.07 mm (24 specimens).
Female genitalia (Figs. 1 18, 148, 149). Corpus bursae somewhat rectangular. Sclerotized plates in dorsal and ventral
wall of ductus bursae short, not extended farther anteriorly than anterior apophyses. Ovipositor valve clothed with fine
setae, a short, sclerotized flange-like projection at valve apex in females from populations of nominate subspecies (see
below).
Type material. - Agrotis quadridentata Grote and Robinson was described from an
undetermined number of males and one female. Of the type material, only one male has been
located, this in the collection of the Academy of Natural Sciences of Philadelphia. This
specimen, labelled “Col.”, “type No. 7596, Agrotis quadridentata 6, A.R. Grote & Rob.”,
“genitalia slide Euxoa 5202 3”, is here designated lectotype.
Carneades pugionis Smith was described from seven males. Type material is in the
collections of the American Museum of Natural History and the United States National
Museum. A lectotype will be designated by Todd (1982).
Quaest. Ent„ 1981, 17 (1,2)
38
Lafontaine
Euxoa flutea Smith was described from two females; these are in the collection of the
American Museum of Natural History. A lectotype will be designated by Todd (1982).
Distribution and period of flight. - Euxoa quadridentata occurs in western North America
from eastern North Dakota and southern Manitoba westward to British Columbia and
southward to southern New Mexico, Arizona and California (Fig. 178). Approximately 1400
specimens from 138 localities were examined.
Specimens have been collected from mid-August until early October.
Habitat. - Euxoa quadridentata occurs in the Great Plains and Great Basin in open prairie
or sagebrush areas and in open pinon-juniper woodland. In the Cascades and Sierra Nevada
Mountains it occurs in lodgepole pine ( Pinus contorta Dougl.) forests.
Remarks. - Specimens of Euxoa quadridentata may be distinguished from those of E.
laetificans by characters in the key (see also remarks under E. laetificans ).
Larvae have been reared on wheat Triticum aestivum L.) in Montana (Cook, 1930b).
Populations of E. quadridentata may be arranged in two subspecies. Differences in
ovipositor valves of the two subspecies suggest that females may oviposit in different soil types.
Hybridization experiments and longer series of females from eastern Nevada would help to
clarify the status of the two subspecies. Relative scarcity of females of this species has
hampered such studies. Of 1400 specimens examined, about 150 were females. Only four
females were located from eastern Nevada. The subspecies of E. quadridentata are as follows.
Euxoa quadridentata quadridentata (Grote and Robinson)
Agrotis quadridentata Grote and Robinson, 1865, p. 491 .
Carneades pugionis Smith, 1900, p. 419.
The nominate subspecies occurs in the Great Plains, the Rocky Mountain region and the
eastern intermontane region as far west as eastern British Columbia, western Montana and
Wyoming, eastern Nevada and northwestern Arizona. Specimens from populations of this
subspecies can be distinguished from those of the western sub-species only by form of female
ovipositor valve. The ovipositor valve of E. q. quadridentata female has a short, sclerotized,
flange-like projection at apex of valve (Fig. 148). That of E. q. flutea female lacks this
sclerotized projection (Fig. 149).
Euxoa quadridentata flutea Smith
Euxoa flutea Smith, 1910b, p. 255.
This subspecies occurs in the Cascades and Sierra Nevada Mountains, and in the
intermontane region as far east as south-central British Columbia, western Idaho and eastern
Nevada. Specimens of E. q. flutea, unlike those of E. q. quadridentata, have darker hind wings
in more mesic habitats.
Euxoa inscripta Lafontaine, new species
Figs. 53, 54,88, 119, 151, 179
Description. — Antenna of male biserrate and bifasciculate. Antenna of female filiform. Frontal tubercle present.
Vestiture of head and thorax mixture of gray and brown scales.
The Euxoa detersa group (Lepidoptera: Noctuidae)
39
Ground color of forewing brown dusted with black. Basal line, transverse anterior and transverse posterior lines, and
subterminal line indistinct. Median line absent. Cubital vein pale, remainder of veins lined with black. Terminal area
dark gray-brown, streaked into subterminal area; portion of streaks in subterminal area black in most specimens.
Reniform and orbicular spots with pale brown shading between dark brown central spot and black outline. Orbicular
spot oval in most specimens Claviform spot black, prominent in most specimens.
Hind wing of male white basally with sharply defined smoky-brown band on terminal third or quarter of wing.
Hind wing of female smoky-brown.
Expanse: 30.6 ± 1.4 mm. Length of forewing: 1 5.2 ± 0.6 mm (80 specimens).
Male genitalia (Fig. 88). Right and left sacculus extensions similar in length, longer than harpes. Vesica without
extra nipple-like sub-basal diverticulum; median diverticulum with basal bulge.
Length of right sacculus extension: 1.52 ±0.09 mm (20 specimens).
Length of left sacculus extension: 1.53 ±0.10 mm (20 specimens).
Length of right harpe: 1.18 ±0.04 mm (20 specimens).
Length of right sacculus: 1.46 ±0.08 mm (20 specimens).
Female genitalia (Figs. 119, 151). Corpus bursae somewhat rectangular; junction of ductus bursae and corpus
bursae about 0.25 from posterior end of corpus bursae. Ovipositor valve with fine setae, sclerotized flange-like projection
at apex.
Type material. - Holotype male, Craig, Colo., 22 mi N., 7100 ft, 15 Aug. 1965 (D.F.
Hardwick), type No. 15940 in the Canadian National Collection, Ottawa. Paratypes: 70 males,
33 females: COLORADO: Craig, 22 mi N., 7100 ft, 15 Aug. 1965 (D.F. Hardwick); Dinosaur,
4 mi NE., 7000 ft, 18 Aug. 1965 (D.F. Hardwick); Maybell, 6 mi SE., 6200 ft, 17 Aug. 1965
(D.F. Hardwick); Sapinero, 5 mi ENE., 7700 ft, 15 Aug. 1971 (D.F. Hardwick); Sargents, 3
mi NE., 9700 ft, 14 Aug. 1971 (D.F. Hardwick). MONTANA: NE side Howie Rd., 1/4 mi
from Gibson Rd., 4500 ft, Sweet Grass Co., 21 Aug. 1969 (J.G. Franclemont). UTAH: Salina,
13 mi SE., 7400 ft, 26 Aug. 1965 (D.F. Hardwick). WYOMING: Alcova, 4 mi SW ., 5500 ft,
14 Aug. 1965 (D.F. Hardwick); Kemmerer, 8 mi SW., 6800 ft, 23 Aug. 1964 (D.F. & V.J.
Hardwick); Lander 10 mi SE., 5600 ft, 31 Aug. 1964 (D.F. & V.J. Hardwick); Sage, 6 mi N.,
6200 ft, 22 Aug. 1964 (D.F. Hardwick).
The holotype and most of the paratypes are in the Canadian National Collection. One
paratype is in the collection of J.G. Franclemont, Ithaca, New York.
Distribution and period of flight. - Euxoa inscripta occurs from southern Montana
southward to western Colorado and central Utah; it also occurs in west-central Nevada and
east-central California (Fig. 179). One hundred and fourteen specimens from 15 localities were
examined.
Specimens have been collected from mid-August until late August.
Habitat. - Euxoa inscripta occurs in arid areas where sagebrush {Artemisia spp.) is
abundant. It is found in areas of coarse, gravelly soil.
Remarks. - In wing pattern, specimens of E. inscripta are similar to those of E. olivalis but
may be distinguished from them by rounded orbicular spot and by lack of silver-gray streaking.
In structural characters, males of E. inscripta are most similar to those of E. unica but may be
distinguished by longer, symmetrical sacculus extensions.
The specific epithet inscripta refers to the fine, black lines on wing veins and maculation.
Specimens from California and Nevada have darker, more gray forewings than do those
from more easterly populations and pale shading between central dark spot and black outline of
reniform and orbicular spots is white in many specimens rather than pale brown.
Euxoa unica McDunnough
Figs. 57,89, 179
Euxoa unica McDunnough, 1940, p. 192; Saskatoon, Saskatchewan.
Quaest. Ent., 1981, 17 (1,2)
40
Lafontaine
Description. — Male antenna markedly biserrate. Vestiture of head, upper half of prothoracic collar and thorax
brown; basal half of prothoracic- collar yellowish brown. Ground color of forewing light brown dusted with darker brown,
especially around reniform and orbicular spots. Transverse lines present. Terminal area dark brown, blackish brown
sagittate spots on proximal side of subterminal line. Costa and cubital vein yellowish buff. Reniform and orbicular spots
pale buff. Claviform spot black. Basal dash small, black. Hind wing white with smoky-brown shading on terminal quarter
of wing and on veins.
Expanse: 31,33 mm. Length of forewing: 15,16 mm (2 specimens).
Male genitalia (Fig. 89). Similar to those of E. inscripta male but sacculus extensions shorter and asymmetrical, left
sacculus extension much shorter than right.
Length of right sacculus extension: 1 .52, 1 .38 mm (2 specimens).
Length of left sacculus extension: 1 .28, 1 .08 mm (2 specimens).
Length of right harpe: 1 .38, 1 .25 mm (2 specimens).
Length of right sacculus: 1 .58, 1 .48 mm (2 specimens).
Female genitalia. Female genitalia of E. unica unknown but would probably be similar to those of E. inscripta female.
Type material. - The holotype of Euxoa unica McDunnough is a male (type No. 5071) in
the Canadian National Collection, Ottawa.
Distribution and period of flight. - Euxoa unica is known from two males, both collected in
the vicinity of Saskatoon, Saskatchewan. Specimens were collected 23 Aug. 1937 (holotype)
and 26 Aug. 1942.
Remarks. - The rarity of Euxoa unica is puzzling. The northern Great Plains region,
including central Saskatchewan, has been well collected in August and September; a light trap
was run for many years at Saskatoon and only two specimens of E. unica were collected. While
there are species known from comparably restricted, formerly glaciated, areas, other
explanations for its apparent rarity must be considered.
Specimens of E. unica differ from those of other species in the E. detersa group in that male
antenna is more prominently biserrate and male sacculus extensions are markedly
asymmetrical. In these features, E. unica males resemble those of E. intrita (Morrison).
Specimens of E. intrita , however, differ from those of E. unica and other species in the E.
detersa group in many ways: antenna more prominently biserrate; forewing shorter and
broader; male sacculus extensions much stouter and more markedly asymmetrical than are
those of E. unica ; harpes longer; sacculus stouter; sub-basal diverticulum of vesica T-shaped
with projections directed both dorsally and ventrally.
Specimens of E. intrita are variable in wing markings; of about 600 specimens in the
Canadian National Collection, only about 10 have wing markings that resemble those of E.
unica specimens; these are from Saskatoon.
If E. unica specimens are of hybrid origin, Euxoa niveilinea (Grote) and E. dargo
(Strecker) appear to be the two most likely candidates for being the other parent in an E.
intrita group-2?, detersa group cross.
The E. intrita group is well-removed from the E. detersa group in the reconstructed
phylogeny of Euxoa (see figure 186) and successful intergroup hybridization in Euxoa is at
present unknown. While hybrid origin for E. unica specimens cannot be demonstrated, further
work in this area would be worthwhile
Euxoa niveilinea (Grote)
Figs. 55, 56, 90, 120, 152
Agrotis niveilinea Grote, 1882, p. 216; Arizona.
Carneades niveilinea-, Smith, 1890b, p. 139; redescription.
Carneades niveilinea-. Smith, 1893, p. 89; catalogue.
Paragrotis niveilinea-, Dyar, 1902, p. 140.
Euxoa niveilinea-, Hampson, 1903, p. 210.
The Euxoa detersa group (Lepidoptera: Noctuidae)
41
Euxoa niveilinea ; Draudt, 1924, p. 37; redescription.
Euxoa rabiata Smith, 1910b, p. 255; Volga, South Dakota; Colorado; Calgary, Alberta.
Euxoa niveilinea rabiata ; Barnes and McDunnough, 1917, p. 41.
Euxoa niveilinea rabiata ; Draudt, 1924, p. 38; redescription.
Euxoa niveilinea rabiata ; McDunnough, 1950, p. 374; redescription.
Agrotis rabiata ; Forbes, 1954, p. 38; good species.
Euxoa rabiata ; Rindge, 1955, p. 128; type material.
Euxoa rabiata ; Lafontaine, 1982; = E. niveilinea.
Description. — Vestiture of head and thorax brownish gray; transverse band of white scales on thorax posterior to
prothoracic collar. Ground color of forewing dark brownish gray dusted with black. Transverse lines obscure in most
specimens. Basal and median areas concolorous; subterminal area slightly paler than median area; terminal area dark
gray, series of black, sagittate spots in sub-terminal area on proximal side of subterminal line in many pecimens. Costa
extensively dusted with silver-gray in many specimens. Veins Cu, Cuj and M3 pale-lined, latter two projected into
terminal area. Reniform and orbicular spots concolorous with subterminal area. Claviform spot black, obscure. Basal
dash obscure or absent. Hind wing of both sexes white with smoky-brown shading on terminal line and veins in most
specimens.
Male genitalia (Fig. 90). Harpe markedly bent mesially, basal half of right harpe forming an angle of about 90° with
right sacculus extension in most males. Right sacculus extension long, curved toward valve apically. Vesica with extra,
nipple-like sub-basal diverticulum and with basal bulge on median diverticulum.
Length of right sacculus extension: 1.78 ±0.10 mm (20 specimens).
Length of left sacculus extension: 1.75 ±0.10 mm (20 specimens).
Length of right harpe: 1 . 1 1 ± 0.07 mm (20 specimens).
Length of right sacculus: 1.39 ±0.07 mm (20 specimens).
Female genitalia ( Figs. 1 20, 1 52). Corpus bursae slightly constricted mesially. Ovipositor valve clothed with fine setae,
sclerotized flange-like projection at apex.
Type material. - Agrotis niveilinea Grote was described from nearly a dozen specimens. Of
these, four males have been located in the collection of the Unites States National Museum. A
male labelled “Arizona”, “Col. B Neumoegen”, “type No. 33736 U.S.N.M.”, “ Agrotis
niveilinea Grote $ type”“ 6 genitalia on slide Apr. 1966 ELT 2172” is here designated
lectotype.
Euxoa rabiata Smith was described from six males and two females. Four male syntypes
are in the collection of the American Museum of Natural History. A lectotype will be selected
by Todd (1982).
Distribution and period of flight. - Euxoa niveilinea occurs in the Great Plains region from
southern Saskatchewan and Alberta southward to northern Texas and southern New Mexico.
Its range extends eastward from the great Plains region into the “prairie peninsula” of central
Michigan and westward in the intermontane region to central Arizona. An apparently disjunct
population occurs in relict prairie habitat in southeastern Texas (Fig. 180). Approximately 700
specimens from 66 localities were examined.
Specimens have been collected from late August until late October.
Habitat. - Euxoa niveilinea inhabits dry, sandy prairie areas where sage ( Artemisia spp.) is
abundant.
Remarks. - Specimens of Euxoa niveilinea are most easily confused with those of E. dargo
and E. cicatricosa. Males of E. niveilinea may be distinguished from those of E. dargo and E.
cicatricosa by long sacculus extension; females may be distinguished by flange-like projection
rather than stout setae on ovipositor valve. Most specimens of E. niveilinea can be distinguished
from those of E. dargo by pale rather than dark hind wing and from those of E. cicatricosa by
presence of transverse band of white scales on the thorax posterior to the prothoracic collar
lacking in E. cicatricosa specimens.
Quaest. Ent., 1981, 17 (1,2)
42
Lafontaine
Euxoa dargo (Strecker)
Figs. 58,91, 121, 153, 181
Agrotis dargo Strecker, 1 898, p. 6; Loveland, Colorado.
Paragrotis dargo\ Dyar, 1902, p. 140.
Euxoa dargo\ Barnes and McDunnough, 1917, p. 41.
Euxoa dargo\ Draudt, 1924, p. 37; redescription.
Agrotis dargo\ Forbes, 1954, p. 38; redescription.
Carneades rumatana Smith, 1903c, p. 203; Volga, South Dakota; Calgary, Alberta.
Euxoa rumatana ; Barnes and McDunnough, 1917, p. 41; = E. dargo.
Carneades rumatana ; Rindge, 1955, p. 129; type material.
Description. — Head, thorax and pattern of forewing similar to that of E. niveilinea but ground color dark brown.
Claviform spot with pale buff streak distal to it. Hind wing of most males buff basally with sharply delimited smoky-brown
band on terminal third or half of wing. Hind wing of some males smoky-brown almost to base. Hind wing of female
smoky-brown, slightly paler at base.
Expanse: 27. 2± 1.5 mm. Length of forewing: 13.7 ±0.8 mm (100 specimens).
Male genitalia (Fig. 91 ). Harpe and sacculus extension short. Vesica with extra nipple-like sub-basal diverticulum and
with basal bulge on median diverticulum.
Length of right sacculus extension: 1 . 1 6 ± 0. 1 0 mm (20 specimens).
Length of left sacculus extension: 1 .09 ±0.10 mm (20 specimens).
Length of right harpe: 0.93 ± 0.08 mm (20 specimens).
Length of right sacculus: 1 . 1 0 ± 0.06 mm (20 specimens).
Female genitalia (Figs. 121, 153). Corpus bursae oval, slightly invaginated on left side in most specimens. Ovipositor
valve with stout setae apically; sub-basal row of long setae stout and spike-like.
Type material. - Agrotis dargo was described by Strecker from “ a number of examples”.
Of this material, two males and two females are in the collection of the Field Museum of
Natural History, Chicago. A male labelled “Col.”, “A. dargo type Streck.”, is here designated
lectotype. The specimen is in good condition; it has been dissected.
Carneades rumatana Smith, was described from sixteen males and one female. Type
material is in the collections of the American Museum of Natural History, the United States
National Museum and the Canadian National Collection. A lectotype will be designated by
Todd (1982).
Distribution and period of flight. - Euxoa dargo occurs in western North America from
southern Manitoba westward to southern British Columbia and southward to central Nebraska,
northern New Mexico, southern Idaho and east-central Oregon (Fig. 181). Approximately 350
specimens from 60 localities were examined.
Specimens have been collected from late August until early October.
Habitat. - Euxoa dargo inhabits dry prairie or sagebrush ( Artemisia spp.) areas. In the
southern portion of its range it is found in areas where prairie habitat occurs in open pine
parkland.
Remarks. - Specimens of E. dargo resemble those of E. niveilinea and E. cicatricosa but
may be distinguished from them by characters in the key (see also remarks under E.
niveilinea). Most specimens of E. dargo can be distinguished from those of E. niveilinea by
darker hind wing and by pale streak distal to claviform spot.
Euxoa melura McDunnough
Figs. 59,92, 122, 154, 182
Euxoa melura McDunnough, 1932, p. 231; Eureka, Utah.
Description. — Head, thorax and wing markings almost identical to those of E. olivalis (p. 73) specimens except
claviform spot larger, filled with dark gray and orbicular spot round or slightly oval in most specimens rather than elongate
and bar-like. Specimen illustrated is one of few with elongate orbicular spot (see Fig. 59).
The Euxoa detersa group (Lepidoptera: Noctuidae)
43
Expanse: 3 1 .2 ± 1.4 mm (20 specimens). Length of forewing: 15.5 ±0.5 mm (32 specimens).
Male genitalia (Fig. 92). Unlike those of any other species in E. detersa group. Harpe elongate, almost straight.
Sacculus extension short, about 0.66 length of harpe. Vesica with extra sub-basal diverticulum; without basal pouch on
median diverticulum.
Length of right sacculus extension: 0.93 ±0.08 mm (1 1 specimens).
Length of left sacculus extension: 0.83 ±0.10 mm (10 specimens).
Length of right harpe: 1.37 ±0.05 mm (11 specimens).
Length of right sacculus: 0.99 ±0.05 mm (1 1 specimens).
Female genitalia (Figs. 122, 154). Similar to those of E. dargo but corpus bursae not invaginated on left side;
ovipositor valve with sub-basal row of long setae mostly thin and hair-like, only 3 or 4 setae at anteroventral corner of
valve stout.
Type material. - The holotype of Euxoa melura McDunnough is a male in the Canadian
National Collection, Ottawa (type No. 3363).
Distribution and period of flight. - The distribution' of E. melura is poorly known. It has
been collected in southern Washington and from southwestern Montana and southern Idaho
southward to west-central Colorado and southern Idaho (Fig. 182). Forty specimens from 8
localities were examined
Specimens have been collected from late May until early July. This species has the earliest
flight period of those in the E. detersa group.
Habitat. - No available data.
Remarks. - Specimens of E. melura are similar in wing markings to those of E. olivalis.
Specimens of E. melura can usually be distinguished from those of E. olivalis by the rounded
orbicular spot and by earlier collecting date. Specimens of the two species are easily
distinguished by genitalia. Males of E. melura have short sacculus extensions and long harpes;
those of E. olivalis have long sacculus extensions and short harpes. Ovipositor valve of E.
melura female has stout setae apically; that of E. olivalis female has fine setae and a
sclerotized flange at apex.
Euxoa detersa (Walker)
Figs. 60,61,93, 123, 155, 183
Charaeas detersa Walker, 1856, p. 212; Nova Scotia.
Agrotis detersa; Grote, 1873, p. 82.
Carneades detersa ; Smith, 1 893, p. 94.
Paragrotis detersa; Dyar, 1902, p. 142.
Euxoa detersa; Hampson, 1903, p. 229.
Euxoa detersa ; Draudt, 1924, p. 40; redescription.
Euxoa detersa; McDunnough, 1949, p. 3.
Euxoa detersa; McDunnough, 1950, p. 371; redescription.
Agrotis detersa; Forbes, 1954, p. 38.
Euxoa detersa; Rings and Johnson, 1976, 1-16; bibliography.
Agrotis pitychrous Grote, 1873, p. 82; Long Island, New York.
Carneades pitychrous; Smith, 1890b, p. 159; redesciption.
Carneades pitychrous; Smith, 1893, p. 94; = E. detersa.
Agrotis personata Morrison, 1876, p. 238; Illinois.
Agrotis personata; Grote, 1880a, p. 187; = E. pitychrous.
Carneades personata; Smith, 1890b, p. 159; = E. pictychrous.
Paragrotis personata; Dyar, 1902, p. 142; = E. pitychrous.
Euxoa personata; Hampson, 1903, p. 229; = E. pitychrous.
Euxoa personata; Smith 1910a, p. 106; good species.
Euxoa detersa personata; Barnes and McDunnough, 1917, p. 41.
Euxoa detersa personata; Draudt, 1924, p. 40; redescription.
Agrotis personata; Forbes, 1954, p. 38; good species.
Euxoa detersa personata; Lafontaine, 1982.
Agrotis az//Strecker, 1898, p. 6; Clyde, New York.
Quaest. Ent., 1981,17(1,2)
44
Lafontaine
Paragrotis azif ; Dyar, 1902, p. 143
Euxoa detersa azif, ; Barnes and McDunnough, 1917, p. 41.
Euxoa detersa personata form azif Draudt, 1 924, p. 40.
Euxoa detersa azif McDunnough, 1938, p. 58.
Euxoa azif Lafontaine, 1982, = E. detersa personata.
Description. — Frontal tubercle markedly reduced, absent in many specimens. Vestiture of head and thorax gray
and brown. Ground color of forewing pale gray, buff or brown. Basal and subterminal areas slightly paler than median
area in some specimens. Terminal area gray-brown, darker than ground color of forewing. Reniform and orbicular spots
slightly paler than median area Claviform spot black, inconspicuous. Costa and veins Cu, Cux and M3 pale-lined and pale
streak distal to claviform spot in most specimens from Atlantic seaboard and lower St. Lawrence River region. Hind wing
of male varied from buff basally with smoky-brown shading on outer third of wing to entirely smoky-brown. Hind wing of
female smoky-brown.
Expanse: 30. 6± 1.2 mm. Length of forewing: 1 5.3 ± 0.8 mm (100 specimens).
Male genitalia (Fig. 93). Harpe markedly curved toward valve mesially and apically. Sacculus invaginated at base of
clavus; clavus large, sclerotized. Vesica without extra nipple-like sub-basal diverticulum and without basal bulge on
median diverticulum.
Length of right sacculus extension: 1 .43 ±0.11 mm (23 specimens).
Length of left sacculus extension: 1.31 ±0.12 mm (23 specimens).
Length of right harpe: 1 .08 ± 0.06 mm (23 specimens).
Length of right sacculus: 1.30 ±0.07 mm (23 specimens).
Female genitalia (Figs. 123, 155). Corpus bursae oval, slightly invaginated on left side. Ovipositor valve with stout
setae dorsally and apically; stout setae scattered on dorsal margin of valve, not in a row. A few long setae on ovipositor
valve sub-basally but no row of long sub-basal setae.
Type material. - Charaeas detersa Walker was described from four specimens; these are in
the collection of the British Museum (Natural History). A male labelled “type”, “Nova
Scotia”, “2. Charaeas ? detersa ”, “Noctuidae, Brit. Mus. slide No. 6307”, is here designated
lectotype.
Agrotis pitychrous Grote was described from an unknown number of specimens from Long
Island, New York. Type material, stated to be in the Lintner collection, now deposited in the
collection of the New York State Museum, Albany, has not been located. Characters given in
the original description, however, are sufficient to allow the names E. detersa and E.
pitychrous to be synonymized.
The nominal holotype of Agrotis personata Morrison is a female in the collection of
Michigan State University, East Lansing. The origin of the holotype was stated to be Illinois,
however, the specimen is labelled “Ohio”.
The holotype of Agrotis azif Strecker is a female in the collection of the Field Museum of
Natural History, Chicago.
Distribution and period of flight. - Euxoa detersa occurs on the Atlantic seaboard from
Gaspe Peninsula, Quebec southward to South Carolina. Its range extends westward through
the St. Lawrence River region and the Great Lakes region to the Great Plains. In central North
America E. detersa occurs from north-central Manitoba and south-central Alberta southward
to southern Minnesota and central Nebraska. It has alsao been collected at Fort Smith,
Northwest Territories (Fig. 183). Reports of this species from Colorado (e.g. Forbes, 1954) are
apparently based on misidentified specimens of E. cinereopallida (Smith), and on mislabelled
specimens of the Atlantic seaboard form of E. detersa. Approximately 1000 specimens from
105 localities were examined.
Specimens have been collected from early August until early October.
Habitat. - This species inhabits areas of loose, shifting sand such as beach and dune areas.
Remarks. - Specimens of E. detersa may be distinguished from those of E. cicatricosa by
characters in the key (see also remarks under E. cicatricosa ).
The larva of E. detersa is commonly called the sandhill cutworm. It is a locally important
economic pest of crops grown in sandy soil including corn, tobacco, potato, strawberry, oats,
The Euxoa detersa group (Lepidoptera: Noctuidae)
45
wheat and rye (Rings and Johnson, 1976).
Populations of E. detersa are arranged in two subspecies as follows:
Euxoa detersa detersa (Walker)
Charaeas detersa Walker, 1856, p. 212.
Agrotis pitychrous Grote, 1873, p. 82.
The nominate subspecies occurs in eastern North America from the Atlantic seaboard
westward in the St. Lawrence River region as far west as the eastern Great Lakes region. In
most specimens of this subspecies the forewing is streaked longitudinally with costa, and veins
Cu, Cux and M3 pale-lined; the claviform spot has a pale strerak distal to it (Fig. 60).
Euxoa detersa personata (Morrison)
Agrotis personata Morrison, 1876, p. 238.
Agrotis ozz/Strecker, 1898, p. 6.
This subspecies occurs from central North America eastward through the Great Lakes region
to the St. Lawrence River region. In specimens of this subspecies the forewing is not streaked
longitudinally (Fig. 61).
Specimens from the upper St. Lawrence River region, the Ottawa Valley and upper state
New York are extremely varied; both streaked and non-streaked forms occur as well as every
combination of forms between these two extremes.
Euxoa cicatricosa (Grote and Robinson)
Figs. 62,63,94, 124, 156, 184
Agrotis cicatricosa Grote and Robinson, 1 865. p. 492; Colorado.
Carneades cicatricosa ; Smith, 1890b, p. 1 38; redescription.
Carneades cicatricosa ; Smith, 1893, p. 88; catalogue.
Paragrotis cicatricosa, Dyar, 1902, p. 140.
Euxoa cicatricosa ; Hampson, 1903, p. 205.
Euxoa cicatricosa, Draudt, 1924, p. 37; redescription.
Euxoa cicatricosa ; McDunnough, 1949, p. 3.
Carneades neomexicana Smith, 1890b, p. 218; New Mexico.
Carneades neomexicana-, Smith, 1893, p. 88; catalogue.
Paragrotis neomexicana, Dyar, 1902, p. 140.
Euxoa neomexicana-, Hampson, 1903, p. 206.
Euxoa cicatricosa neomexicana-, Barnes and McDunnough, 1917, p. 41 .
Euxoa cicatricosa neomexicana-, Draudt, 1924, p. 37; redescription.
Carneades neomexicana-, Rindge, 1955, p. 122; type material.
Euxoa neomexicana-, Lafontaine, 1982; = E. cicatricosa.
Setagrotis ducalis Smith, 1907, p. 128; Stockton, Utah.
Euxoa ducalis ; Barnes and McDunnough, 1 9 1 7, p. 4 1 .
Euxoa ducalis-, Draudt, 1924, p. 37; redescription.
Euxoa ducalis-, McDunnough, 1925, p. 244; near E. perolivalis.
Euxoa ducalis-, Lafontaine, 1982 = E. cicatricosa.
Euxoa teplia Smith, 1910b, p. 253; Colorado; Stockton, Utah.
Euxoa teplia\ Barnes and McDunnough, 1917, p. 41; = E. cicatricosa.
Euxoa teplia ; Rindge, 1955, p. 133; type material.
Description. — Frontal tubercle present. Vestiture of head and thorax gray, brown, and white scales. Ground color
of forewing gray-brown with pale streaks. Subterminal area, costa, reniform and orbicular spots, streak distal to claviform
Quaest. Ent„ 1981, 17 (1,2)
46
Lafontaine
spot, and veins Cu, Cuj and M3 silver-gray, white, or orange-brown, paler than remainder of forewing. Veins Cuj and M3
pale-lined, projected into terminal area. Small, black basal dash in some specimens. Claviform spot black, small. Hind
wing of male white, light buff shading near teminal margin in some specimens. Hind wing of female buff with
smoky-brown shading on veins in many specimens.
Expanse: 29. 8± 1.4 mm. Length of forewing: 1 4.8 ± 0.7 mm (100 specimens).
Male genitalia (Fig. 94). Similar to those of E. detersa male; most structures smaller, on average, than those of E.
detersa male.
Length of right sacculus extension: 1.32 ±0.1 2 mm (25 specimens).
Length of left sacculus extensison: 1.16 ±0.1 3 mm (25 specimens).
Length of right harpe: 0.99 ±0.05 mm (25 specimens).
Length of right sacculus: 1.24 ±0.08 mm (25 specimens).
Female genitalia (Figs. 124, 156). Similar to those of E. detersa female but may be distinguished from them by
arrrangement of stout setae on ovipositor valve. Stout setae on ovipositor arranged in a row on dorsal margin of valve.
Type material. - The holotype of Agrotis cicatricosa Grote and Robinson is a fdmale (type
No. 7595) in the collection of the Academy of Natural Sciences of Philadelphia.
Carneades neomexicana Smith was described from several specimens. Syntypes are in the
collections of the United States National Museum and the American Museum of Natural
History. A lectotype will be designated by Todd (1982).
The holotype of Setagrotis ducalis Smith is a female in the collection of the United States
National Museum. The specimen has been dissected.
Euxoa teplia Smith was described from one male and one female. The specimens are in the
collection of the American Museum of Natural History. A lectotype will be designated by Todd
(1982).
Distribution and period of flight. - Euxoa cicatricosa occurs from central Saskatchewan
westward to south-central British Columbia and southward to northern Texas, southern New
Mexico, central Arizona and southern California (Fig. 184). Approximately 1300 specimens
from 101 localities were examined.
Specimens have been collected from late August until late October.
Habitat. - Euxoa cicatricosa occurs in very arid areas where vegetation is sparse and soil is
loose, coarse sand or gravel.
Remarks. - Specimens of E. cicatricosa are similar to those of E. niveilinea in wing
markings and color. Males of E. cicatricosa may be distinguished from those of E. niveilinea by
shorter sacculus extension of the genitalia; females may be distinguished by stout setae rather
than sclerotized flange on ovipositor valve. Most specimens of E. cicatricosa can be
distinguished from those of E. niveilinea by lack of transverse white band on thorax posterior to
prothoracic collar and by presence of a pale streak distal to claviform spot on forewing.
In structural characters, E. cicatricosa specimens are most similar to those of E. detersa and
E. recula. Stout setae on ovipositor of E. cicatricosa female form a single row on dorsal margin
of valve; these setae are scattered on dorsal margin of valve, not in a row, in females of E.
detersa and E. recula.
Specimens of E. cicatricosa may be distinguished from those of E. recula by pattern and
color of forewing. In specimens of E. cicatricosa the forewing is dark gray-brown with fine
silver-gray streaks. Reniform and orbicular spots have dark central spots. In specimens of E.
recula , pale areas of forewing are more extensive and are yellow. Reniform and orbicular spots
and a broad streak extended from claviform spot to subterminal line are yellow.
Most specimens of E. cicatricosa may be distinguished from those of E. detersa personata
by presence of longitudinal streaks on the forewing, particularly pale-lined veins Cu2 and M3
projected into terminal area and by pale hind wing. A form of E. cicatricosa occurs at some
localities in the Great Plains, particularly in south-central Saskatchewan, with the forewing not
The Euxoa detersa group (Lepidoptera: Noctuidae)
47
longitudinally streaked. Forewings of these specimens are similar to those of E. detersa
specimens from the same region and these specimens may be the result of hybridization. In
other characters, however, the specimens are not intermediate; setal arrangement on ovipositor
valves, hind wing color, and frontal tubercle size are normal for E. cicatricosa specimens.
Similar unstreaked forms also occur in Texas, Kansas and Colorado outside of the known range
of E. detersa.
Euxoa cicatricosa specimens are varied in wing markings. The most widespread form (Fig.
63) occurs throughout the range of the species except for the Great Basin and southwestern
United States. A pale form, with more extensive white streaks (Fig. 62) occurs in the Great
Basin. Pale markings on specimens from New Mexico and Arizona are orange-brown rather
than white or silver-gray.
Euxoa recula (Harvey)
Figs. 64,95, 125, 157, 185
Agrotis recula Harvey, 1876, p. 37; Oregon.
Carneades recula\ Smith, 1 890b, p. 1 38; redescription.
Carneades recula ; Smith, 1893, p. 88; catalogue.
Paragrotis recula\ Dyar, 1902, p. 140.
Euxoa recula ; Hampson, 1903, p. 207; redescription.
Euxoa cicatricosa recula ; Barnes and McDunnough, 1917, p. 41 .
Euxoa cicatricosa recula\ Draudt, 1924, p. 37; redescription.
Euxoa recula ; Lafontaine, 1982; good species.
Description. — Frontal tubercle present. Vestiture of head and thorax yellow-buff and gray scales. Ground color of
forewing brownish gray with extensive yellow shading. Basal and subterminal areas, costa, reniform and orbicular spots,
and area distal to claviform spot shaded with yellow in most specimens. Some specimens with silver-gray shading on costa
and in subterminal area. Veins Cu, Cuj and M3 pale-lined, latter two projected into subterminal area. Hind wing of male
white; hind wing of female pale buff with smoky-brown shading on veins.
Expanse: 30.8 ± 1 .5 mm. Length of forewing: 1 5.4 ± 0.7 mm ( 1 00 specimens).
Male genitalia (Fig. 95). Similar to those of E. detersa and E. cicatricosa males but most structures, on average,
smaller.
Length of right sacculus extension: 1 . 1 2 ± 0. 1 2 mm (20 specimens).
Length of left sacculus extension: 1 .02 ±0.12 mm (20 specimens).
Length of right harpe: 0.92 ±0.04 mm (20 specimens).
Length of right sacculus: 1 .20 ±0.04 mm (20 specimens).
Female genitalia (Figs. 125, 157). Indistinguishable from those of E. detersa female. May be distinguished from those
of E. cicatricosa female by arrangement of stout setae on dorsal margin of ovipositor valve, these scattered, not in a single
row as in E. cicatricosa female.
Type material. - Agrotis recula was described by Harvey from two females collected in
Oregon. One of these has been located in the collection of the British Museum (Natural
History). The specimen, labelled “type”, “Oregon, Grote Coll. 81-116”, “5969 Oregon”,
“ Agrotis recula Harvey”, “ Agrotis recula Harvey type”, “Noctuidae Brit. Mus. slide No.
6359”, is here designated lectotype. The specimen is a female in fair condition.
Distribution and period of flight. - Euxoa recula occurs from central Oregon southward to
southern California and westward through southern Arizona to central New Mexico (Fig. 185).
Approximately 400 specimens from 32 localities were examined.
Specimens have been collected from early September until late October.
Habitat. - Euxoa recula occurs in habitats similar to those inhabited by E. cicatricosa. The
two species are frequently collected together in Nevada and Oregon. In southern Arizona and
in the Mojave Desert, California, however, E. recula is found in areas where E. cicatricosa does
not occur.
Quaest. Ent., 1981, 17 (1,2)
48
Lafontaine
Remarks. - Specimens of E. recula may be distinguished from those of E. cicatricosa and
E. deter sa by characters in the key (see also remarks under E. cicatricosa).
PHYLOGENY
Introduction
The following chapter presents a discussion of two phylogenies, one of the subgenera and
species groups of Euxoa (Fig. 186), a of the species of the E. detersa group (Fig. 187).
Construction of both phylogenies was based on Hennigian principles as outlined by numerous
recent workers (Schlee, 1969; Kavanaugh, 1972; Whitehead, 1972; Munroe, 1974) by
establishing sister group relationships on the basis of shared derived character states
(synapomorphies of Hennig, 1966). Morphological uniformity among Euxoa species, and
extensive intraspecific variability frequently made it necessary to characterize groups on the
basis of overall similarity and group trends. This parallels the situation described by Whitehead
(1976) for the weevil genus Rhinochenus
Before attempting a phylogenetic reconstruction of the E. detersa group, it was necessary to
trace character states and trends through the rest of the genus in order to determine ancestral
and derived conditions in the E. detersa group and to evaluate their usefulness for phylogenetic
purposes. Such a comparison of Euxoa character states has not been done previously. In
addition, other genera of the Noctuinae were examined to determine ancestral character states
for the genus.
The structural uniformity within the family Noctuidae, especially among the assemblage of
subfamilies collectively known as trifid noctuids, makes the formulation of phylogenetic
reconstructions particularly difficult. Richards (1932) and Kiriakoff (1963), for example,
studied the structure of the thoracic ear in an attempt to shed light on the phylogeny of the
Noctuidae and related families. They were, however, unable to find any characters that would
aid in distinguishing the “trifid” subfamilies. Phylogenies have been constructed for the genera
of the Noctuinae (Forbes, 1933); Sterrhini (Hogue, 1963); Heliothidinae (Hardwick, 1970b);
and Plusiinae (Eichlin and Cunningham, 1978).
Alternatives to a strictly morphological approach to phylogenetic reconstruction in the
noctuids have barely been considered. The possibility of using serological studies of proteins
was investigated by Martin and Cotner (1934) and discussed by Hudson (1973). Although the
results of these investigations seem promising, a more critical and comprehensive study has not
been made.
Relation of Euxoa to other noctuine genera
In a revision of the genera of the Agrotinae ( = Noctuinae), McDunnough (1928) stated that
Euxoa was most closely related to Protexarnis, Loxagrotis, and Pseudorthosia although he did
not include a detailed phylogeny. Forbes (1933), using a phenetic approach to reconstruct a
phylogeny for the Noctuinae, provided the most reasonable hypothesis of the phyletic position
of Euxoa in the subfamily. Forbes considered Peridroma to be the genus retaining the largest
number of ancestral character states within the Noctuinae. This position is further supported
by the work of Birch (1972a, 1972b) on British noctuids who found that Peridroma is the only
genus in the Noctuinae that has retained male abdominal scent-brushes.
The Euxoa detersa group (Lepidoptera: Noctuidae)
49
According to Forbes’ hypothesis, a Peridroma- like ancestor gave rise to two lineages, one in
which the fore-tibial setae of adults were progressively reduced, and one in which they were
enlarged into digging claws. Also in this latter group, the length of the fore-tibia is reduced and
the frons is modified, this usually either bulging and roughened or with a projecting frontal
tubercle. These three derived character states are related to the subterranean pupation habit of
the larvae, apparently being adaptations that aid the adults in escaping from the soil upon
emergence. Genera belonging to this group include Euxoa, Agrotis, Feltia, and Loxagrotis.
Ancestral character states of these genera are: male antennae biserrate and bifasciculate; juxta
with a projecting spine; male valve with sacculus large; sacculus with an apical bulge; digitus
present on cucullus; harpe short, projected dorsally; vesica with one or more cornuti near base;
female with bursa copulatrix bisaccate (i.e. cervix bursae present, see Callahan and Chapin,
1960); ovipositor valves without stout setae or sclerotized projections. According to Forbes,
Euxoa is the sister group of a large number of genera (e.g. Agrotis, Feltia, Loxagrotis) that
share the same tarsal and frontal modifications as Euxoa. Although Forbes does not mention
which characters were used in reconstructing this phylogeny, the shared, derived character
state of Euxoa is development in males of a sacculus extension projected along the ventral edge
of the cucullus. Also the digitus on the cucullus is lost in Euxoa. A shared, derived character
state of the sister group of Euxoa is a long, sclerotized projection extended onto the vesica from
the apex of the aedeagus. Males of most genera of this group, however, are more easily
recognized by the position and shape of the harpe, which is reduced and is extended along the
inner surface of the cucullus. The female genitalia, not used by Forbes, offer excellent derived
character states for these two lineages. The ancestral condition of the ductus bursae is one in
which the walls are sclerotized and form a sheath. The sclerotized portion varies from
occupying the posterior third to extending almost the full length of the ductus. In the female
genitalia of Euxoa the sclerotized portion is reduced to two plates, one in the dorsal wall and
one in the ventral wall of the ductus. In females of the sister group of Euxoa the ductus bursae
is membranous.
Fletcher (1961) considered the African genus Euxootera to be closely related to Euxoa.
This assumption, however, is based primarily on a sacculus extension in males. This represents
a situation similar to that of Protexarnis which was discussed by Forbes (.1933). Lack of tarsal
and frontal modifications in adults of these genera suggests that the sacculus extensions are
convergent with those of Euxoa males.
Phylogeny of species groups of Euxoa
The ancestral character states for the genus are: those of the ancestral states of the Euxoa
group of genera (see above) plus, male genitalia with uncus tapered to apex; sacculus extension
short and rounded; harpe without pubescence; vesica curved in a loop above apex of aedeagus to
project dorsally; vesica with a single sub-basal diverticulum.
Reconstruction of a phylogeny of Euxoa was difficult and results must be considered
tentative. Three major problems encountered were structural uniformity of the species,
extensive parallelism in some character states, and scattered distribution of a number of derived
character states. Many of the character states used are not discrete derived character states but
are similarities in shape and proportion created by modifications of a number of genitalic
structures. Species groups were combined on the basis of shared, derived character states into
progressively more inclusive lineages until all species groups were included in the phylogeny.
Quaest. Ent., 1981, 17 (1,2)
50
Lafontaine
The structures and the distribution of derived character states among the subgenera and
species groups of Euxoa, which are used in the phylogenetic analysis, are shown in Table 1 .
Parallelism and secondary losses
The structural uniformity among Euxoa species presents a serious obstacle to phylogenetic
analysis. Many of the derived character states listed in table 1 are changes in shape or size and
are susceptible to parallel evolution. For example, trends toward a reduction in the size of the
sacculus and a reduction in the length of the sacculus extension occur independently in a
number of lineages.
Some character states are restricted to a lineage but are not found in all species groups of
the lineage, nor are they restricted to a monophyletic grouping within it. Apparently, a number
of character states (e.g. #6, 8, 13) have been derived only once but have subsequently been lost
a number of times.
Other character states, such as presence of a second sub-basal diverticulum on the male
vesica, are present in a number of species groups but cannot be homologized from group to
group. As a result, it has not been possible to determine how many times this character state
has been derived and whether or not it has been lost secondarily by some species groups.
In spite of the hazards of using characters susceptible to parallelism and secondary loss, the
groupings obtained appear to be monophyletic. Evidence of this was discovery of additional
characters and trends that were revealed by grouping taxa not formerly associated. An example
of this was discovery of the value of the shape and setal arrangement of the male uncus as
useful taxonomic and phylogenetic characters.
Interpretation of character states
Some of the character states and trends listed in table 1 are explained as a necessary
antecedent for reviewing the reconstructed phylogeny of subgenera and species groups shown in
figure 186.
Frontal tubercle ( character 2). - The frontal tubercle has been lost three times, each in
species associated with sandy areas.
Uncus (character 3). - An apically spatulate uncus has either been derived independently in
the E. cinereopallida-E. mitis lineage and the E. luctuosa group, or, more probable, it has been
derived once, in their common ancestor, and lost in Orosagrotis.
Sacculus extension and harpe ( characters 6 & 7). - A form of the male valve, which
converges on the Agrotis- like condition, with the harpe along the inner surface of the cucullus
parallel to its dorsal margin and with the sacculus extension reduced has been derived three
times: lineage 5; lineage 6 ( E . edictalis group of Pleonectopoda)\ and lineage 13 E.
cinereopallida-Orosagrotis lineage).
Vesica loop ( character 8). - The loop in the vesica has been lost four times (lineages 1, 3, 4,
5-13), each time in a different way.
Vesica - second sub-basal diverticulum ( character 10). - A second sub-basal diverticulum is
present in the vesica in thirteen species groups of subgenus Euxoa. The second diverticulum is a
useful phylogenetic structure within species groups. It is, however, of little use for intergroup
relationships because the shape and position of the diverticulum differs between groups and
cannot be homologized.
Vesica-cornuti (character 11). - Loss of the cornuti from the vesica is not a useful character
state for phylogenetic groupings. Cornuti have been lost in fourteen species groups and in most
Table 1 . Structures and Evolutionary Classification and Distribution of Character States among Subgenera and Species Groups of Euxoa.
The Euxoa detersa group (Lepidoptera: Noctuidae)
51
x
<D
Cd
T3 ^
00
0 ^
c do
•-
to <L>
5 5
„ *
CO
C/3
(continued on next page)
Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by
lineage (Fig. 186)
52
Lafontaine
T3
C 1
'o' 3
c
<D
■9
(continued on next page)
Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by
lineage (Fig. 186)
The Euxoa detersa group (Lepidoptera: Noctuidae)
53
OO IT)
ro
cd
H
<d
t/3
CD
3-<
a.
X
aJ
2
’> _
<D cd
X
2 2
cd ^3
T3 cd
a,
1/3
O
O c
G i
o *
cd
G
«
a,
x
W
*2
s
O)
Ex. - ex-group comparisons
In. - in-group comparisons
54
Lafontaine
others, their presence or absence varies from species to species and often from specimen to
specimen.
Vesica shape ( not in table). - The most useful phylogenetic character for associating species
and species groups is the overall shape of the vesica. The shape is usually constant within a
species group; species-specific characters of the vesica normally involve shape and position of
the accessory pouches, primarily the sub-basal diverticulum.
Bursa (character 12). - Shape of the bursa copulatrix is a very useful phylogenetic
character. A unisaccate bursa (i.e. cervix bursa lost) has been derived eight times. A unisaccate
bursa which is constricted mesially with the ductus bursae entering the corpus bursae at the
posterior end has been derived four times: lineage 1; lineage 3 ( E . olivia group); lineage 6; and
lineage 9 ( E . tessellata- E. choris groups). This type of unisaccate bursa probably evolved from
a bisaccate type similar to that of lineage 5 (compare bursa of E. camalpa and that of E.
serotina in Lafontaine, 1976b). An oval unisaccate bursa in which the ductus bursae enters the
corpus bursae on the right side has been derived four times: lineage 4 (in E. westermanni group
of (Pleonectopoda); lineage 9 ( E . rufula-E. terrena groups); lineage 10 (E. pallipennis group);
and lineages 12 and 13.
Ovipositor flanges ( character 13). - Sclerotized flange-like projections are present on the
ovipositor valves in nineteen species groups of the subgenus Euxoa (lineages 10-13), and in its
derivative Orosagrotis. The limited distribution of this structure within Euxoa and its
restriction to this genus suggests that it was derived only once, this being in the common
ancestor of lineages 10-13. Within these lineages, the flange-like structure has been lost many
times and probably reacquired occasionally. This structure is apparently quite plastic, its
presence or absence being related to oviposition habits.
Ovipositor with sclerotized rim ( character 15). - This character state is found only in the E.
rufula-E. serricornis lineage; it has been secondarily lost in E. rufula and E. intrita.
Phytogeny of the genus Euxoa
The following is a brief explanation of the phylogeny presented in figure 186.2 A detailed
survey of the distribution of character states in the genus will not be made, nor will an attempt
be made to show all branching points in the phylogeny of lineages 8, 9 and 1 1 .
An evolutionary classification, and distribution by lineage, of the character states used in
the following discussion is given in table 1. The purpose of this section is to establish a rough
working hypothesis of the phylogeny of the genus in order to put character states used in the
phylogeny of the E. detersa group into perspective, and to show the relationship of the E.
detersa group to the other species groups.
Dichotomy 1. - The shared, derived character states of lineage 1 are the unique shapes of
the female corpus bursae and the male vesica (see Hardwick, 1970a). Lineages 2-13 share the
derived state of character 4.
Dichotomy 2. - Lineage 2, while one of the most easily distinguished groups in the genus
(see Lafontaine, 1975a), is difficult to define in terms of uniquely derived character states
because of the large number of ancestral character states retained by its members. Structurally,
species in this group are more similar to the hypothetical common ancestor of the genus than
2It will eventually be necessary to create a subgenus for both lineage 2 and 3. This would result
in the genus consisting of six monophyletic subgenera with the subgenus Euxoa remaining
paraphyletic with respect to the very distinctive subgenus Orosagrotis.
Table 2. Structures and Evolutionary Classification and Distribution of Character States among Adults of the Euxoa detersa Group.
The Euxoa detersa group (Lepidoptera: Noctuidae)
55
X>
<D
cS
4J ^
T3 r-
00
0 T
c ob
3
X> <3->
•r
£ cs
S a
Quaest. Ent., 1981, 17(1,2)
Structure/ Character Ancestral State Derived State Basis for classf n Times derived Times lost Distribution of derived state by
lineage (Fig. 187)
56
Lafontaine
<Z3
C
O
.Sr
’>
O
1 3 Ovipositor - conical absent present
setae
(continued on next page)
Structure/ Character Ancestral State Derived State Basis for classfn Times derived Times lost Distribution of derived state by
lineage (Fig. 187)
The Euxoa detersa group (Lepidoptera: Noctuidae)
57
<N
7
LO
<n
I
— 1
"O
<D
00
Quaest. Ent., 1981, 17 (1,2)
58
Lafontaine
Table 3. Distribution of Derived Character States among Adults of the Euxoa detersa
Group
(continued on next page)
The Euxoa detersa group (Lepidoptera: Noctuidae)
59
Table 3 (continued)
Lineage No. Species Fig. No. Character State Number
(see table 2)
Male Genitalia
123456789 10 11
= not present in all specimens
- = derived character state secondarily lost
Quaest. Ent., 1981, 17 (1,2)
60
Lafontaine
Table 3 (continued)
(continued on next page)
The Euxoa detersa group (Lepidoptera: Noctuidae)
61
Table 3 (continued)
* = not present in all specimens
those of any other group. The shape of the bursa copulatrix in females of this lineage is unlike
that of females in any other species group. Lineages 3-13 share the derived state of characters
6a and 7.
Dichotomy 3. - The shared, derived character state of lineage 3 is the asymmetrical valves
of the male genitalia (see Lafontaine, 1976d). Lineages 4-13 share the derived character state
of having lost the loop in the vesica. This has occurred independently in lineage 1 and lineage 3
( E . olivia and E. septentrionalis groups).
Dichotomy 4. - The loss of the vesica loop in the remaining sub-genera and species groups
(lineages 4-13) has occurred in one of two ways. The vesica has a submedian twist or coil in
Quaest. Ent., 1981, 17 (1,2)
62
Lafontaine
males of the subgenus Pleonectopoda and ancestrally in Longivesica males (see Hardwick
(1970a). In Longivesica this is still evident in males of the E. divergens group. In the subgenus
Euxoa (as restricted above) and Orosagrotis , the vesica has a simple bend sub-basally without
a loop or twist in it.
Dichotomies 5-8. - Within the subgenus Euxoa , the affinities of four lineages remain
uncertain with presently available information. The remaining lineages (9-13) have been
associated on the basis of the pubescence on the harpe (character 8). Although this character
state is present in less than one-third of the species groups of lineages 9-13, its restriction to
members of the subgenus Euxoa suggests that it has been derived only once. On this admittedly
tenuous basis, lineages 9-13 are considered to form a monophyletic group.
Dichotomy 9. - The species within lineage 9 share a number of uniquely derived character
states but none of these is exhibited by all species groups. A markedly incurved C-shaped harpe
is found in most species groups and is therefore probably a shared, derived character state of
this lineage. Other derived character states are: a basally enlarged sacculus, this tending to be
teardrop-shaped in males of the E. declarata-E. simulata lineage rather than crescentic as in
males of other species groups; the sub-basal diverticulum of the vesica rather than being
elbowed and foot-shaped, is T-shaped in males of the E. rufula-E. setonia lineage. Another
character state uniquely derived in lineage 9 is the presence of a sclerotized rim around the
posterior margin of the ovipositor valve (character 15). This is found in females of the E.
rufula-E. serricornis lineage but has been lost in females of the otherwise structurally similar
E. rufula- E. intrita lineage. The bisaccate bursa copulatrix has been lost from females of the
E. rufula-E. terrena lineage. Lineages 10-13 share the derived state of character 13. This has
been lost independently at least eighteen times.
Dichotomy 10. - Derived character states of lineage 10 are the shape of the vesica and the
derived state of character 6c. Lineages 11-13 share the derived state of character 5.
Dichotomy 11. — Members of lineages 11-13 have descended from a common ancestor with
a bisaccate bursa copulatrix which was probably similar in shape to that found in females of the
E. punctigera group (see Lafontaine, 1974c). A simple reduction of the cervix bursae (the left
sac) resulted in the corpus bursae (the right sac) being oval, or oblong, with the ductus bursae
entering it at about one-third of the way along the right side from the posterior end. This type
of bursa characterizes females of lineages 12 and 13, and inependently, females of the E.
pallipennis group. A modified type of bisaccate bursa copulatrix characterizes females of the
E. obeliscoides-E. basalis lineage of lineage 1 1 in which the corpus bursae is enlarged at each
end giving it a dumbbell, or figure 8 appearance (see Hardwick, 1973a, 1973b, and Hardwick
and Lefkovitch, 1973). Reduction of the cervix bursae would result in a unisaccate bursa
copulatrix that is very different in shape from that which characterizes females of lineages 12
and 13. In females of the E. tessellata-E. albipennis lineage the corpus bursae appears swollen
posteriorly; the ductus bursae enters at the posterior end. In females of lineages 12 and 13 the
corpus bursae is oval; the ductus bursae enters the corpus bursae on the right side. An exception
to this is that the corpus bursae is oval in females of the E. albipennis group. This group is
included in lineage 1 1 on the basis of characters of the male genitalia.
Lineage 1 1 is also characterized by derived group trends in the male genitalia. The sacculus
tends to be reduced, the cucullus lengthened and the apical portion of the cucullus enlarged and
foot-shaped, frequently with a well developed ‘heel’ at the posteroventral corner. The apical
two-thirds of the harpe tends to be straight rather than evenly incurved or S-shaped.
The Euxoa detersa group (Lepidoptera: Noctuidae)
63
Dichotomy 12. - Within lineages 12 and 13, the most difficult group to characterize in
terms of uniquely derived character states is lineage 12 (the E. detersa group). In lineage 13,
the sacculus extension is reduced in males of most species, being much shorter than the harpe.
In females of lineage 13, the sclerotized flanges on the ovipositor valves tend to be fused
together. In males of the E. cinereopallida- Orosagrotis lineage the harpe is along the inner
surface of the cucullus. This character state, together with reduction of the sacculus extension,
converges on the form of the valve characteristic of males of Agrotis and Feltia. Another
derived character state of males of the E. cinereopallida-Orosagrotis lineage, but lost from
Orosagrotis males, is an apically swollen uncus. In females of the E. luctuosa group, the ductus
seminalis has shifted anteriorly to the middle of the left side of the corpus bursae (see
Lafontaine, 1976c). This trend is continued in Orosagrotis females with the ductus seminalis
arising at the anterior end of the bursae ( E . wilsoni group), or on the right side ( E . ridingsiana
group) (see Hardwick, 1970a).
Lineage 12 (the E. detersa group), on the other hand, is characterized by the lack of the
derived character states and trends described above for lineage 13. It is possible then, that the
E. detersa group is paraphyletic in that lineage 13 may have evolved from a lineage within the
E. detersa group.
Although many details of the phylogeny presented in figure 199 have been omitted, the
above discussion is sufficient to elucidate the position of the E. detersa group within the genus
and to allow ancestral character states to be inferred.
Phylogeny of the Euxoa detersa group
Ancestral character states for adults of the E. detersa group are: maculation normal (i.e
transverse lines all present, longitudinal streaking absent); male antenna moderately biserrate;
frontal tubercle present; male genitalia with sacculus extension about 1.25 times length of
harpe; vesica with single sub-basal diverticulum, this containing several cornuti, and with
median diverticulum situated about one-third of way from sub-basal bend to apex; female
genitalia with ovipositor valve clothed with fine setae, a sclerotized flange at apex of each valve,
and a row of long setae sub-basally; ductus bursae entering corpus bursae on right side about
one-third of way from posterior end.
The 31 species in the E. detersa group are easily arranged in 12 monophyletic lineages (see
figure 187) on the basis of both structural and wing marking similarity. Many of the species
within these lineages are distinguished only by numerical techniques or subtle differences in
wing markings. Most of these lineages were defined and grouped into more inclusive
monophyletic units on the basis of discrete, shared, derived character states. Unfortunately,
structural uniformity within the E. detersa group made it necessary to associate some lineages
on the basis of overall structural similarity.
Structures and distribution of derived character states used in the phylogenetic analysis of
the E. detersa group are shown in tables 2 and 3.
The reconstructed phylogeny of the Euxoa detersa group, and the numbers of characters
used in grouping the lineages, are shown in figure 187. Each circled number represents one of
the 12 lineages.
Early differentiation produced two stocks (lineages 1-5; and lineages 6-12), which have
become almost equally diverse. In general, the pattern of character differentiation has been
conservative, consisting of slight modifications (with frequent reversals) of various elements of
the genitalia, in contrast to a more linear type of character modification resulting from
Quaest. Ent., 1981, 17 (1,2)
64
Lafontaine
progressive changes of a single character. Nevertheless, lineages 1-5 are linked by a
modification of the bursa, and in general exhibit more modifications in the female genitalia
than are exhibited by females of lineages 6-12. Conversely, lineages 6-12 are linked by
modifications of the male vesica, and generally exhibit more changes in the male genitalia than
are exhibited by males of lineages 1-5.
Lineages 1 and 3 share the derived character state of the vesica projected to the left rather
than dorsally above the apex of the aedeagus (character 6). This is most likely a derived
character state shared by lineages 1-3 that has been lost by lineage 2. The vesica of specimens
of lineage 2 ( E . brevipennis ) has a slight twist in the sub-basal bend of the vesica (character 7)
returning the direction of its projection to the ancestral position.
The shared, derived character state linking lineages 6-12 is presence in males of an
additional, small, sub-basal diverticulum (character 8) situated dorsolaterally on the left side
opposite the normal, foot-like sub-basal diverticulum. This has apparently been lost from
lineages 9 and 12 and from males of some species of lineages 7 and 8.
The shared, derived character state uniting lineages 7-12 is that the median diverticulum of
males has a bulge at its base and the basal half of the diverticulum is projected dorsally parallel
to the axis of the vesica (characater 10). This has been lost in lineage 12 and by males of E.
melura and some specimens of E. laetificans. Ancestrally this diverticulum is horn-shaped and
projects from the vesica at right angles.
The sequence of evolutionary changes of male vesica is shown in diagram 1 (page 67).
Lineage 7 contains adults whose forewings are not streaked longitudinally while those of
lineages 8-12 are streaked (character 1). The^non-streaked pattern is considered to be the
ancestral condition for the E. detersa group since this is the pattern that predominates in the
genus. A similar streaked pattern, however, does occur independently in a number of species
groups. Such a pattern would be expected to arise many times since it is cryptic for individuals
that rest among grasses during the day. Although the streaked and non-streaked patterns seem
susceptible to parallel evolution, a dichotomy based on these does appear to be justified on the
basis of the corresponding overall similarity of the genitalia of species within each group.
Lineages 8, 9 and 10 are each considered to be the sister group of the remaining lineages on
the basis that after each dichotomy, species in the remaining lineages are more similar to each
other in both genitalia and wing pattern.
Alternative hypothesis
Structural uniformity among species of the Euxoa detersa group presents an obvious,
obstacle to phylogenetic analysis. Many structural characters that are present have been
independently lost or derived in several lineages, this further weakening the credibility of the
reconstructed phylogeny. Considering the shortcomings of available data, it is advisable to
discuss some alternative hypotheses and problem areas of the reconstructed phylogeny of the
Euxoa detersa group in the hope that discovery of additional characters will shed light on these
problem areas.
Problem areas in a reconstructed phylogeny can be of three types; incorrect assignment of
species to a lineage; incorrect reconstructed phylogeny of species within a lineage; incorrect
arrangement of lineages.
The only apparent possibility of a problem of the first type is with lineage 9. The sister group
relationship of species in lineage 9 would be incorrect if it could be shown that Euxoa unica
McD. is not a species but the product of interspecific hybridization (see remarks under E.
The Euxoa detersa group (Lepidoptera: Noctuidae)
65
unica).
Of the second type, problems associated with reconstructing the phylogeny of lineages 1 and
3 are discussed in the next section.
Many problems were encountered in arranging lineages into larger monophyletic groups,
these resulted from a lack of characters. General appearance of wing markings and genitalia of
members of lineages 2 and 5 are incongruent with those of lineages with which they have been
associated. Undoubtedly some incongruency results from different habitat preferred by
members of these two lineages, however, the possibility of convergence cannot be ruled out. The
larger number of derived character states shared by members of lineage 2 and those of
associated lineages as compared to that of members of lineage 5 makes the possibility of
convergence in lineage 2 less likely than it is in lineage 5.
Arrangement of lineages 8-10 is based on increasing overall similarity of their members to
those of lineages 11 + 12. Discovery of additional characters could change this arrangement.
In the Euxoa detersa group characters suitable for phylogenetic analysis are so few, and the
possibility of convergent evolution so great, that discovery of additional characters could
change the position of some lineages in any number of ways. For these reasons, no attempt is
made to suggest where lineages discussed above would be placed in a reconstructed phylogeny
after discovery of additional characters.
Phylogeny of the species of the Euxoa detersa group
Of the five lineages that include more than two species (i. e. #1, 3, 4, 7 and 12), lineage 1
presented the most difficulties in reconstruction of phylogenetic relationships. The species in
this lineage are so uniform structurally that only distribution patterns, wing color and
maculation can be used in reconstructing a phylogeny. Euxoa costata and E. castanea are
considered sister species on the basis of the characteristic reddish-brown, or chestnut, color of
the forewings. The assumption that these two species share a recent common ancestry is further
supported by the distribution patterns of the two species. Euxoa castanea is distributed in the
Rocky Mountain region while E. costata is distributed along the Cascade-Sierra Nevada axis,
the two being sympatric only in the north-west (see figures 158 and 159). The phylogen;y of the
remaining three species in the lineage was constructed from more circumstantial evidence.
Euxoa clausa and E. foeminalis both have restricted distributions which are contained within
the distribution of E. idahoensis. Euxoa idahoensis is a widespread and polytypic species with
features which are suitably ancestral for either one, whereas E. clausa and E. foeminalis are
forms which would fall at opposite ends of the range of variation of E. idahoensis. It thus seems
more likely that E. clausa and E. foeminalis are more closely related to E. idahoensis than to
each other. Euxoa foeminalis is tentatively associated more closely with E. idahoensis than is
E. clausa because of the similarities in wing markings.
In lineage 3 E. servita is considered to be the sister species of the E. redimicula-E.
arizonensis lineage because E. servita is the most divergent of the four species in the lineage in
terms of both genitalia structures and maculation and is sympatric with the other three species
(see Lafontaine, 1974a). The three species in the E. redimicula-E. arizonensis lineage are
largely allopatric and may be the result of trichotomous speciation. The evidence, however,
suggests otherwise. Adults of E. auripennis and E. arizonensis are more similar to each other
than they are to those of E. redimicula. It is possible that E. arizonensis and E. auripennis
speciated from a common ancestor during the Wisconsinan while E. redimicula appears to have
arisen in pre-Wisconsinan times. Both E. auripennis and E. redimicula occur in disjunct
Quaest. Ent., 1981, 17(1,2)
66
Lafontaine
woodland areas within the Great Plains and the two may have been widely sympatric in this
area during the Wisconsinan when large portions of the Great Plains were forested.
In the phylogeny of lineage 4, E. agema is considered to be the sister species of E.
oblongistigm. Although adults of E. agema are more similar to those of E. olivalis in terms of
wing markings than to adults of E. oblongistigma, the genitalia of males of E. agema and E.
oblongistigma share the derived character state of having a shortened sacculus extnsion and a
lengthened harpe. These structural characters probably reflect phylogenetic affinities more
accurately than do wing markings.
In lineage 1\ males of E. moerens, E. latro, and E. murdocki share a derived condition of
the uncus setae. In these species, setae on the middle portion of the uncus are short and stout,
and have a brush-like appearance. Normally only setae on the apical third of the uncus are
stouter and these are not erect and brush-like; setae on the middle portion of the uncus are thin
and hair-like. Also the antennal serrations are reduced in these three species. Within this group,
E. moerens and E. latro are considered to be sister species on the basis of similarities in wing
color and maculation.
Several derived character states are shared by E. dodi and E. infracta. Both have lost the
sclerotized flange-like projection from the ovipositor valve, the second sub-basal diverticulum of
the vesica, and both have pubescent harpes.
The phylogeny of lineage 12 has been depicted as a trichotomy but four taxa must be
considered in the discussion because E. detersa consists of two subspecies. No characters could
be found which would suggest that any two of the species are more closely related to each other
than to the third. In fact, the two subspecies of E. detersa are more dissimilar in wing pattern
than are any of the three species.
Evolution of the genitalia
Species of Euxoa are not well suited for studies of evolution of genitalia. Structural
uniformity among species makes it difficult to determine the significance of character
transformations.
Character transformations in genitalia may be arranged in three categories: those of
membranous structures (bursa copulatrix, vesica), those of male valves and unci, those of
ovipositor valves.
There is apparently a general correlation between transformations in the male vesica and
those in the bursa copulatrix. Position (e.g. dorsal, lateral) and angle of the portion of the
corpus bursae that leads to the ductus seminalis, with respect to position of the ductus bursae, is
related to direction and angle of projection of the vesica. The possible importance of this, and of
shape and position of accessory diverticula of the vesica, to spermatophore placement and
insemination is discussed in the following section. Such transformations are. probably not
related directly to origin of species since many closely related species do not differ in vesica or
bursa shape. The sequence of evolutionary changes in the male vesica is shown in diagram 1 .
Character transformations in sclerotized structures of male genitalia are, in most instances,
not lost or acquired structures but are changes in proportion. Most of these changes involve size
of sacculus, length of harpe and length of sacculus extension. Although these structures are all
components of the “claspers”, the effects of transformations do not necessarily affect clasping
ability in the sense of the “lock and key theory”. It is possible that these structures have a
sensory or stimulatory funcation during copulation. The same could be said of transformations
of shape and setal arrangement of the uncus. A correlation may exist between sacculus
The Euxoa detersa group (Lepidoptera: Noctuidae)
67
extension length and uncus shape; species groups that have apically enlarged unci in males,
have very short sacculus extensions. It is possible that in males of these groups there has been a
transfer of a sensory or clasping role from sacculus extensions to unci.
Character transformations in ovipositor valves are related to oviposition habits rather than
to copulation. Frequently, ovipositor valve shape or structure differs among females of closely
related species (e.g. those of lineages 1, 3, 7, 8, 11 in Fig. 187). It is probable that females that
differ in characters of ovipositor valve lay their eggs in different soil types; thus, there is a
partitioning of resources among closely related species. The functional significance of a
sclerotized projection, or of stout setae, on ovipositor valves is unknown. The sequence of
evolutionary changes in the ovipositor valves are shown in diagram 2.
In summary, functional and evolutionary significance of character transformations in
genitalia of Euxoa species will not be known until behavioral studies have been made; this must
be done with living material, not by comparison of structures using genitalic preparations.
Diagram 1. Sequence of evolutionary changes of male vesica of Euxoa detersa group. a-Ancestral condition (e.g. E.
agema ); b-Vesica projected laterally (e.g. E. auripennis)\ c-Median diverticulum mesial (e.g. E. idahoensis); d-Vesica
with basal twist, vesica projected dorsally (e.g. E. brevipennis)\ e-Extra sub-basal diverticulum present (e.g. E. teleboa)\
f-Median diverticulum with basal bulge (e.g. E. moerens); g-Extra sub-basal diverticulum absent (e.g. E. infracta);
h-Basal bulge on median diverticulum absent (e.g. E. melura)\ i-Extra sub-basal diverticulum and basal bulge on median
diverticulum both absent (e.g. E. detersa ).
Quaest. Ent., 1981, 17 (1,2)
68
Lafontaine
Diagram 2. Sequence of evolutionary changes of ovipositor valves of Euxoa detersa group, a - Ancestral condition (e.g. E.
olivalis)', b - Enlarged ventrally (e.g. E. citricolor)\ c - Stout setae apically (e.g. E. servita)-, d - Sclerotized flange absent
(e.g. E. brevipennis)\ e - Conical setae apically (e.g. Euxoa eostata)\ f- Sclerotized flange absent (e.g. E. infraeta)\ g -
Sclerotized flange fin-like (e.g. E. teleboa)-, h - Stout setae dorsally and apically (hypothetical); i - Stout setae basally
(e.g. E. dargo ); j - Basal row of long setae absent (e.g. E. detersa).
The Euxoa detersa group (Lepidoptera: Noctuidae)
69
The genitalia as an isolating mechanism
In any study involving evolution of male and female genitalia structures, the question arises
as to whether or not the genitalia function as an isolating mechanism (the lock and key theory).
This has been the subject of much debate (see review by Rentz, 1972). It is reasonable to
assume that both situations exist; genitalia structure may not be an isolating mechanism among
species that have elaborate behavioral or olfactory premating cues but may be an important
isolating mechanism among species that lack these cues (Rentz, 1972).
Although mechanical aspects of copulation and insemination have been described for
numerous species of Lepidoptera (e.g. see Arnold and Fischer, 1977; Callahan and Chapin,
1960; Ferro and Akre, 1975; Hardwick, 1965b; Proshold et al., 1975; Scott, 1978; StekoFnikov,
1965), problems in mating between members of species that have dissimilar genitalia have been
described only for Helicoverpa (Hardwick, 1965b, p. 37).
In general, the spermatophore must be placed in the bursa copulatrix so that its opening is
adjacent to the opening of the ductus seminalis, although successful insemination without a
spermatophore has been observed (see George and Howard, 1968). While members of two
species that have dissimilar male valves may be able to mate successfully (e.g. Euxoa
campestris (Grote) X E. declarata (Walker), Byers and Hinks, 1978), those of two species in
which male vesicae or female bursae are dissimilar may not be able to cross successfully.
Combination of vesica shape and bursa shape is critical to positioning of the spermatophore in
the bursa (Callahan and Chapin, 1960; Hardwick, 1965b). Shape of the bursa is also important
in sperm transfer since the bursa contracts and holds the spermatophore in place so that sperm
is ejected into the ductus seminalis (see Proshold et al. , 1975).
Intergroup matings have been made in Euxoa (e.g. E. campestris (Grote) ( E. declarata
group) X E. basalis (Grote) (E. basalis group) and E. niveilinea (Grote) ( E. detersa group) X
E. plagigera (Morrison) ( E. tessellata group), J.R. Byers, pers. comm.), however, no fertile
eggs were obtained. In each mating a spermatophore was passed to the bursa, this indicating
that valve differences between the male used and a male conspecific with the female did not
prevent mating; it was not determined if the spermatophore was positioned correctly in the
bursa, or if sperm reached the spermatheca.
In genera in which vesica and bursa shape differ among species, the shape of these
structures may be a more important isolating mechanism than is shape of male valves. Further
work on the importance of spermatophore placement to insemination may prove to be very
rewarding.
Conclusions
In reconstructing the phylogeny of both the species groups of Euxoa and the species of the
E. detersa group, an attempt was made to rely on discrete, shared, derived character states
whenever possible. Unfortunately, the paucity of suitable characters, together with
susceptibility of many characters to parallel and convergent evolution, made it necessary to
group taxa according to overall similarity in several instances. The Hennigian method of
grouping taxa on the basis of shared, derived character states has limited applicability when
dealing with morphologically similar species such as those of the E. detersa group.
Nevertheless, the resulting phylogenies, while admittedly tenuous, do represent hypotheses with
which new data can be compared and thereby provide predictive, testable models.
Quaest. Ent., 1981, 17 (1,2)
70
Lafontaine
ACKNOWLEDGEMENTS
I extend my sincere appreciation to the many individuals who have contributed to this study.
I thank my supervisor G.E. Ball of The University of Alberta for direction and
encouragement during the course of this study and for his painstakingly detailed review of the
manuscript.
I am indebted to D.F. Hardwick of the Biosystematics Research Institute for
encouragement and guidance in this and other studies of Euxoa . Dr. Hardwick initiated
revisionary work on Euxoa checked and photographed most of the type material, and collected
the bulk of the specimens used in this study.
I express my gratitude to W.G. Evans, J.G. Packer and C.E. Schweger of The University of
Alberta, and C.L. Hogue of the Los Angeles County Museum for reading and criticizing the
manuscript.
I thank H. Goulet, D.F. Hardwick, and E.G. Munroe for reviewing portions of the
manuscript and providing numerous useful suggestions.
I am grateful to J.R. Byers for making available data from his studies on immature stages,
life histories, and hybridization experiments.
I express my appreciation to J.S. Ashe, A. Borkent, H. Goulet, and R.E. Roughley, fellow
students with whom I have had numerous discussions that have greatly influenced the course of
this study.
I thank my associate, E.W. Rockburne, for invaluable technical assistance, T.H. Stovell who
photographed adults and genitalia, and the Bio-graphic Unit, Research Branch, Canada
Department of Agriculture, for preparing the plates.
I am indebted to B.E. Bowen and G. E. Lewis for assistance on field trips.
I thank all individuals and institutions listed in Section 2.1 for loan of material. The
hospitality of J.G. Franclemont and R.W. Hodges and his wife Elaine, made trips to Cornell
University and the United States National Museum possible and a pleasure.
I thank E.L. Quinter, who provided invaluable assistance to me during a visit to the
American Museum of Natural History.
I express my sincere appreciation to my wife Herma, who accompanied me on numerous
field trips, endured the tribulations and frustrations of being the wife of an entomologist and
student, and was a constant source of encouragement.
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Figs. 1-8. Adults of Euxoa spp. 1, Euxoa costata (Grote), male, Golden, B.C., 2 mi E., 3000 ft; 2, Euxoa costata
(Grote), female, Radium Hot Springs, B.C., 16 mi N., 2900 ft; 3, E. castanea Lafontaine, new species, male holotype,
Golden, B.C., 2 mi E., 3000 ft; 4, E. castanea Lafontaine, new species, female paratype, Radium Hot Springs, B.C., 16
mi N., 2900 ft; 5, E. foeminalis (Smith), male, Prescott, Ariz., 5 mi N., 5450 ft; Yavapai Co.; 6, E. foeminalis
(Smith), female, Prescott, Ariz., 5 mi N., 5450 ft, Yavapai Co.; 7, E. idahoensis (Grote), male, Exshaw, Alta., 5 mi
NE., 4225 ft; 8, E. idahoensis (Grote), female, Exshaw, Alta., 5 mi NE„ 4225 ft. (Reproduced at about 1.3X)
The Euxoa detersa group (Lepidoptera: Noctuidae)
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Figs. 9-16. Adults of Euxoa spp. 9, E. idahoensis (Grote), male, Lethbridge, Alta.; 10, E. idahoensis (Grote), male.
Radium Hot Springs, B.C., 16 mi N., 2900 ft; 11, E. idahoensis (Grote), male. Eureka, Utah; 12, E. idahoensis
(Grote), male, Hill City, Pennington Co., S. Dak.; 13, E. clausa McDunnough, male, Lethbridge, Alta.; 14, E. clausa
McDunnough, female, Lethbridge, Alta.; 15, E. brevipennis (Smith), male, McDermitt, Nev., 6 mi W., 4600 ft; 16, E.
brevipennis (Smith), female, Warm Springs, Nev., 15 mi ESE., 5300 ft. (Reproduced at about 1.3X).
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Figs. 17-24. Adults of Euxoa spp. 17, E. servita (Smith), male, Beverley, Sask., 1 mi S., 2400 ft; 18, E. servita
(Smith), female, Lloydminster, Alta.; 19, E. servita (Smith), male, Sault St. Marie, Mich.; 20, E. redimicula
(Morrison), male, Perth Road, Frontenac Co., Ont.; 21, E. redimicula (Morrison), female, Perth Road, Frontenac Co.,
Ont.; 22, E. auripennis Lafontaine, male paratype, Choteau, Mont., 10 mi NW., 4050 ft; 23, E. auripennis Lafontaine,
female, Leavenworth, Wash., 9 mi NW., 2200 ft; 24, E. arizonensis Lafontaine, female paratype, Greer, White Mts.,
Apache Co., Ariz., 8500 ft. (Reproduced at about 1.3X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
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Figs. 25-32. Adults of Euxoa spp. 25, E. olivalis (Grote), male, Mt. Evans, Colo., 9800 ft; 26, E. olivalis (Grote),
female, McGaffey, Zuni Mts., McKinley Co., N. Mex., 7500 ft; 27, E. olivalis (Grote), female, Lee Vining, Calif., 7
mi WSW., 9600 ft; 28, E. agema (Strecker), male, Gallatin Gateway, Mont., 43 mi S., 6700 ft; 29, E. agema
(Strecker), male, Seneca, Ore., 4.5 mi NW., 4800 ft; 30, E. oblongistigma (Smith), male, Eastend, Sask., 2 mi WSW.,
3200 ft; 3.1, E. oblongistigma (Smith), male, Laramie, Wyo., 8 mi SE., 8400 ft; 32, E. oblongistigma (Smith), female,
Ovando, Mont., 5 mi W., 4000 ft. (Reproduced at about 1.3X).
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Figs. 33-40. Adults of Euxoa spp. 33, E. citricolor (Grote), male, Fallon, Nev., 11 mi W., 4100 ft; 34, E. citricolor
(Grote), male, McDermitt, Nev., 6 mi W., 4600 ft; 35, E. citricolor (Grote), female, Walsenberg, Colo., 2 mi W„ 6500
ft; 36, E. tronella (Smith), Tempiute, Nev., 6 mi SSW., 5600 ft; 37, E. tronella (Smith), male. Badlands Nat. Mon., 6
mi NW. Interior, S. Dak., 2300 ft; 38, E. tronella (Smith), female, Eureka, Utah, 1 1 mi NW„ 5800 ft; 39, E. teleboa
(Smith), male, Sand Springs, Mont., 3100 ft; 40, E. teleboa (Smith), female, Capulin Nat. Mon., 6 mi SW. Folsom, N.
Mex., 7300 ft. (Reproduced at about 1.3X)
The Euxoa detersa group (Lepidoptera: Noctuidae)
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Figs. 41-48. Adults of Euxoa spp. 41, E. moerens (Grote), male, Austin, Nev., 2 mi SE., 7600 ft; 42, E. moerens
(Grote), male, Lukachukai, Ariz., 13 mi SE., 7300 ft; 43, E. moerens (Grote), male, Ely, Nev., 5 mi SW., 7400 ft; 44,
E. latro (Barnes and Benjamin), male, Chiloquin, Ore., 21 mi N., 4500 ft; 45, E. latro (Barnes and Benjamin), female.
Big Bear City, Calif., 6 mi S., 6600 ft; 46, E. murdocki (Smith), male, Yakima, Wash., 20 mi NW„ 2100 ft; 47, E.
dodi McDunnough, male, Sand Springs, Mont., 1 mi S., 3100 ft; 48, E. infracta (Morrison), female. Radium Hot
Springs, B.C., 16 mi N., 2900 ft. (Reproduced at about 1.3X).
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Figs. 49-56. Adults of Euxoa spp. 49, E. laetificans (Smith), male. Wolf Creek, Mont., 13 mi N., 3800 ft; 50, E.
laetificans (Smith), female, Wolf Creek, Mont., 13 mi N., 3800 ft; 51, E. quadridentata (Grote and Robinson), male.
Burns, Ore., 10 mi W., 4300 ft; 52, E. quadridentata (Grote and Robinson), female, Follyfarm, Ore., 5 mi SE., 4250 ft;
53, E. inscripta Lafontaine, new species, male holotype, Craig, Colo., 22 mi N., 7100 ft; 54, E. inscripta Lafontaine,
new species, female paratype, Kemmerer, Wyo., 8 mi SW., 6800 ft; 55, E. niveilinea (Grote), male, Hugo, Colo., 6 mi
NW„ 5200 ft; 56, E. niveilinea (Grote), male, Capulin Nat. Mon., 6 mi SW., Folsom, N. Mex., 7300 ft. (Reproduced
at about 1.3X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
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Figs. 57-64. Adults of Euxoa spp. 57, E. unica McDunnough, male holotype. Saskatoon, Sask.; 58, E. dargo (Strecker),
male? Swift Current, Sask.; 59, E. melura McDunnough, female, Satus Creek, Yakima Co., Wash.; 60, E. d. detersa
(Walker), male, E. Wareham, Mass.; 61, E. detersa personata (Morrison), male, Simcoe, Ont.; 62, E. cicatricosa
(Grote and Robinson), male, Austin, Nev., 17 mi. E., 6600 ft; 63, E. cicatricosa (Grote and Robinson), male, Tonasket,
Wash., 8 mi S., 1100 ft; 64, E. recula (Harvey); male, Trona, Calif., 12 mi NNE., 2700 ft. (Reproduced at about
1.3X).
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(Grt.); 66, E. castanea Lafontaine, new species; 67, E. foeminalis (Sm.). (Reproduced at about 9X). Legend: unc.,
uncus; jux., juxta; sacc., sacculus; cue., cucullus; har., harpe; sacc. ext., sacculus extension; s.b. div., sub-basal
diverticulum; med. div., median diverticulum; ves., vesica; ap. div., apical diverticulum.
The Euxoa detersa group (Lepidoptera: Noctuidae)
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Figs. 68-70. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 68, E.
idahoensis (Grt.); 69, E. clausa McD.; 70, E. brevipennis (Sm.). (Reproduced at about 9.6X).
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Figs. 71-73. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 71, E. servita
(Sm.); 72, E. redimicula (Morr.); 73, E. auripennis Lafontaine. (Reproduced at about 9.6X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
89
Figs. 74-76. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 74, E.
arizonensis Lafontaine; 75, E. olivalis (Grt.); 76, E. agema (Stkr.). (Reproduced at about 10. 8X).
Quaest. Ent. , 17 (1,2)
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Figs. 77-79. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 77, E.
oblongistigma (Sm.); 78, E. citricolor (Grt.); 79, E. tronella (Sm.). (Reproduced at about 10. 2X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
91
Figs. 80-82. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 80, E. teleboa
(Sm.); 81, E. moerens (Grt.); 82, E. latro (B. & Benj.). (Reproduced at about 10. 8X).
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Figs. 83-85. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 83, E. murdocki
(Sm.); 84, E. dodi McD.; 85, E. infracta (Morr.). (Reproduced at about 10. 8X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
Figs. 86-88. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 86, E.
laetificans (Sm.); 87, E. quadridentata (Grt. & Rob.); 88, E. inscripta Lafontaine, new species. (Reproduced at about
9.6X).
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89
90
91
Figs. 89-91. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 89, E. unica
McD.; 90, E. niveilinea (Grt.); 91, E. dargo (Stkr.). (Reproduced at about 10. 8X).
The Euxoa detersa group (Lepidoptera: Noctuidae)
95
Figs. 92-95. Male genitalia of Euxoa spp. (Aedeagus removed and shown at right with vesica everted). 92, E. ntelura
McD.; 93, E. detersa (Wlk.); 94, E. cicatricosa (Grt. & Rob.); 95, E. reeula (Harv.). (Reproduced at about 9.6X).
Quaest. Ent. , 17 (1,2)
96
Lafontaine
Figs. 96-101. Female genitalia of Euxoa spp. 96, E. costata (Grt.); 97, E. castanea Lafontaine, new species; 98, E.
foeminalis (Sm.); 99, E. idahoensis (Grt.); 100, E. clausa McD.; 101, E. brevipennis (Sm.). (Reproduced at about
4.5X).
97
The Euxoa detersa group (Lepidoptera: Noctuidae)
105
Figs. 102-109. Female genitalia of Euxoa spp. 102, E. servita (Sm.); 103, E. redimicula (Morr.); 104, E. auripennis
Lafontaine; 105, E. arizonensis Lafontaine; 106, E. olivalis (Grt.); 107, E. agema (Stkr.); 108, E. oblongistigma (Sm.);
109, E. citricolor (Grt.). (Reproduced at about 4.5X).
Quaest. Ent. , 17 (1,2)
98
Lafontaine
114 115 117
Figs. 110-117. Female genitalia of Euxoa spp. 110, E. tronella (Sm.); Ill, E. teleboa (Sm.); 112, E. moerens (Grt.);
1 13, E. latro (B. & Benj.); 114, E. murdocki (Sm.); 115, E. dodi McD.; 116, E. infracta (Morr.); E. infracta (Morr.);
1 17, E. laetificans (Sm.). (Reproduced at about 4.5X).
Figs. 1 18-125. Female genitalia of Euxoa spp. 118, E. quadridentata (Grt. & Rob.); 119, E. inscripta Lafontaine, new
species; 120, E. niveilinea (Grt.); 121, E. dargo (Stkr.); 122, E. melura McD.; 123, E. detersa (Wlk.); 124, E.
cicatricosa (Grt. & Rob.); 125, E. recula (Harv.). (Reproduced at about 4.5X).
Quaest. Ent. , 17 (1,2)
Figs. 126-133. Ovipositor valves of Euxoa spp., dorsal aspect. 126, E. costata (Grt.); 127, E. castanea Lafontaine, new
species; 128, E. foeminalis (Sm.); 129, E. idahoensis (Grt.); 130, E. idahoensis (Grt ); 131, E. clausa McD.; 132, E.
brevipennis (Sm.); 133, E. servita (Sm.). (Reproduced at about 21.6X).
The Euxoa deters a group (Lepidoptera: Noctuidae)
101
Figs. 134-141. Ovipositor valves of Euxoa spp., dorsal aspect. 134, E. redimicula (Morr.); 135, E. auripennis
Lafontaine; 136, E. arizonensis Lafontaine; 137, E. olivalis (Grt.); 138, E. agema (Stkr.); 139, E. oblongistigma (Sm.);
140, E. citricolor (Grt.); 141, E. tronella (Sm.). (Reproduced at about 20. 8X).
Quaest. Ent. , 17 (1,2)
102
Lafontaine
Figs. 142-149. Ovipositor valves of Euxoa spp., dorsal aspect. 142, E. teleboa (Sm.); 143, E. moerens (Grt.); 144, E.
latro (B. & Benj.); 145, E. murdocki (Sm.); 146, E. dodi McD.; 147, E. infracta (Morr.); 148, E. q. quadridentata
(Grt. & Rob.); 149, E. quadridentata flutea Sm. (Reproduced at about 20. 8X).
The Euxoa deter sa group (Lepidoptera: Noctuidae)
103
Figs. 150-157. Ovipositor valves of Euxoa spp., dorsal aspect. 150, E. laetificans (Sm.); 151, E. inscripta Lafontaine,
new species; 152, E. niveilinea (Grt.); 153, E. dargo (Stkr.); 154, E. melura McD.; 155, E. detersa (Wlk.); 156, E.
cicatricosa (Grt. & Rob.); 157, E. recula (Harv.). (Reproduced at a about 20. 8X).
Quaest. Ent. , 17 (1,2)
104
Lafontaine
Figs. 158-161. Distribution of Euxoa spp. 158, E. costata (Grt.); 159, E. castanea Lafontaine, new species; 160, E.
idahoensis (Grt.); 161, E. foeminalis (Sm.) (▼), E. clausa McD. (■).
The Euxoa detersa group (Lepidoptera: Noctuidae)
Figs. 162-163. Distribution of Euxoa spp. 162, E. servita (Sm.); 163, E. redimicula (Morr.).
Quaest. Ent. , 17 (1,2)
106
Lafontaine
Figs. 164-167. Distribution of Euxoa spp. 164, E. auripennis Lafontaine (0), E. arizonensis Lafontaine (T); 165, E.
olivalis (Grt.); 166, E. agema (Stkr.); 167, E. oblongistigma (Sm.).
The Euxoa detersa group (Lepidoptera: Noctuidae)
107
Figs. 168-171. Distribution of Euxoa spp. 168, E. brevipennis (Sm.); 169, E. citricolor (Grt.); 170, E. tronella (Sm.);
171, E. teleboa (Sm.).
Quaest. Ent. , 17 (1,2)
108
Lafontaine
Figs. 172-175. Distribution of Euxoa spp. 172, E. moerens (Grt.); 173, E. latro (B. & Benj.); 174, E. murdocki (Sm.);
175, E. dodi McD.
The Euxoa detersa group (Lepidoptera: Noctuidae)
109
Figs. 176-179. Distribution of Euxoa spp. 176, E. infracta (Morr.); 177, E. laetificans (Sm.); 178, E. quadridentata
(Grt. & Rob.); 179, E. inscripta Lafontaine, (0), new species E. unica McD. (■).
Quaest. Ent. , 17 (1,2)
110
Lafontaine
Figs. 180-182. Distribution of Euxoa spp. 180, E. niveilinea (Grt.); 181, E. dargo (Stkr.); 182, E. melura McD.
The Euxoa detersa group (Lepidoptera: Noctuidae)
111
Figs. 183-185. Distribution of Euxoa spp. 183, E. detersa (Wlk.); 184, E. cicatricosa (Grt. & Rob.); 185, E. recula
(Harv.).
Quaest. Ent. , 17 (1,2)
112
Lafontaine
CHORIZAGROTIS
ol i vi<
septentrional is
vernal is
mums
2— misturata
atri strigata
cinereopallida
mitis
luctuosa
Fig. 186. Reconstructed phylogeny of subgenera and species groups of Euxoa
OROSAGROTIS
The Euxoa detersa group (Lepidoptera: Noctuidae)
113
costata
Fig. 187. Reconstructed phylogeny and habitat of species of Euxoa detersa group. Circled numbers are lineages
described in text. Small numbers represent characters listed in tables 2 and 3 (negative numbers represent the
secondary loss of derived character states). Habitat symbols: □ open aridlands; A pinon-juniper woodland; ■ conifer
forest.
Quaest. Ent. , 17 (1,2)
The Euxoa detersa group (Lepidoptera: Noctuidae)
115
Index to Names of Taxa
FAMILY GROUP TAXA
Agrotinae, 48
Antilocapridae, 3
Bovidae, 3
Heliothidinae, 48
Incurvariidae, 3
Lepidoptera, 3
Micropterygidae, 3
Noctuidae, 3, 7, 47, 48
Noctuinae, 48
Plusiinae, 48
Sterrhini, 48
GENERA AND SUBGENERA
Agrotis, 49, 63
Chorizagrotis, 4
Euxoa, 2, 3, 4, 5, 7, 9, 10, 35, 37, 40, 48,
49, 50, 54, 62, 66, 67, 69, 70
Euxootera, 49
Feltia, 49, 63
Helicoverpa, 69
Longivesica, 62
Loxjagrotis, 48, 49
Orosagrotis, 4, 50, 54, 62, 63
Peridroma, 48
Pleonectopoda, 50, 54, 62
Protexarnis, 48, 49
Pseudorthosia, 48
Rhinochenus, 48
SPECIES AND SUBSPECIES
&XINDX, 69
agema (Strecker), Euxoa, 11, 16, 27, 28,
29, 65, 66
agema Barnes and
McDunnough, Euxoa, 28
agema Dyar, Paragrotis, 28
agema Hampson, Euxoa, 28
agema Strecker, Agrotis, 28
albipennis group, Euxoa, 62
angulirena Rindge, Euxoa, 22
angulirena Smith, Euxoa, 22, 23
arizonensis Lafontaine,
Euxoa, 1 1, 14, 16, 24, 25, 26, 27, 65
auripennis Lafontaine,
Euxoa, 10,11,14,16,24,25,26,27,
65
azif Dyar, Paragrotis, 44
azif Lafontaine, Euxoa, 44
azif Strecker, Agrotis, 43, 44, 45
basalis (Grote), Euxoa, 69
basalis group, Euxoa, 62, 69
brevipennis (Smith), Euxoa, 11, 13, 18,
23,63
brevipennis Barnes and
McDunnough, Euxoa, 21
brevipennis brevistriga
Barnes and
McDunnough, Euxoa, 22
brevipennis Draudt, Euxoa, 22
brevipennis Dyar,
Paragrotis, 22
brevipennis Hampson,
Euxoa, 22
brevipennis Smith, Agrotis, 22, 23
brevipennis Smith,
Carneades, 22
brevistriga Rindge, Euxoa, 22
brevistriga Smith, Euxoa, 22, 23
camalpa Dyar, Euxoa, 54
campestris (Grote), Euxoa, 66, 69
castanea Lafontaine, new
series, Euxoa, 10, 11, 14, 17, 19, 20,
21,65
choris group, Euxoa, 54
cicatricosa (Grote and
Robinson), Euxoa, 9, 10, 11, 12, 15,
41,42, 44, 45,46, 47,48
cicatricosa Draudt, Euxoa, 45
cicatricosa Dyar,
Paragrotis, 45
cicatricosa Grote and
Robinson, Agrotis, 45, 46
cicatricosa Hampson,
Euxoa, 45
cicatricosa McDunnough,
Euxoa, 45
cicatricosa neomexicana
Quaest. Ent., 1981, 17 (1,2)
V
116 Lafontaine
Barnes and
McDunnough, Euxoa, 45
cicatricosa neomexicana
Draudt, Euxoa , 45
cicatricosa recula Barnes
and McDunnough,
Euxoa , 47
cicatricosa recula Draudt,
Euxoa , 47
cicatricosa Smith,
Carneades , 45
cinereopallida (Smith),
Euxoa, 44
cinereopallida group,
Euxoa, 50, 62, 63
citricolor (Grote), Euxoa, 10, 11, 12, 16,
29, 30,31
citricolor Draudt, Euxoa , 29
citricolor Dyar, Paragrotis, 29
citricolor Grote, Agrotis, 29, 30
citricolor Grote, Carneades , 29
citricolor postmedialis
Draudt, Euxoa, 29
citricolor postmedialis
McDunnough, Euxoa, 29
citricolor postmedialis
Strand, Euxoa, 29
citricolor Smith, Carneades, 29
clausa McDunnough,
Euxoa, 11, 14, 17, 21, 22, 23, 35, 65
costata (Grote), Euxoa, 10, 11, 14, 17, 18,
19, 20, 22, 23,65
costata Barnes and
McDunnough, Euxoa, 18, 21
costata Draudt, Euxoa, 18, 22
costata Grote, Agrotis, 18,19
costata Hampson,
Triphaena, 18
Costata idahoensis Barnes
and McDunnough,
Euxoa, 21
costata idahoensis Draudt,
Euxoa, 21
costata idahoensis
McDunnough, Euxoa, 21
costata Smith,
Rhynchagrotis , 18
dargo (Strecker), Euxoa, 11, 13, 15, 40,
41,42, 43
dargo Barnes and
McDunnough, Euxoa, 42
dargo Draudt, Euxoa, 42
dargo Dyar, Paragrotis, 42
dargo Forbes, Agrotis, 42
dargo Strecker, Agrotis, 42
declarata (Walker), Euxoa, 69
declarata group, Euxoa, 62
detersa (Walker), Euxoa, 1,9, 10, 11, 15,
43, 44, 45, 46, 47, 48, 66
detersa azif Barnes and
McDunnough, Euxoa, 44
detersa azif McDunnough,
Euxoa, 44
detersa detersa (Walker),
Euxoa, 9, 11, 12, 45
detersa Draudt, Euxoa, 43
detersa Dyar, Paragrotis, 43
detersa Forbes, Agrotis, 43
detersa Grote, Agrotis, 43
detersa Hampson, Euxoa, 43
detersa McDunnough,
Euxoa, 43
detersa personata
(Morrison), Euxoa, 9, 11, 12, 44, 45,
46
detersa personata Barnes
and McDunnough,
Euxoa, 43
detersa personata Draudt,
Euxoa, 43, 44
detersa personata
Lafontaine, Euxoa, 43
detersa Rings and Johnson,
Euxoa, 43
detersa Smith, Carneades, 43
detersa Walker, Charaeas, 43, 44, 45
didi Draudt, Euxoa, 34
divergens group, Euxoa, 62
dodi McDunnough, Euxoa, 11, 12, 18, 23,
34,35,36,66
ducalis Barnes and
McDunnough, Euxoa , 45
The Euxoa detersa group (Lepidoptera: Noctuidae)
117
ducalis Draudt, Euxoa, 45
ducalis Lafontaine, Euxoa, 45
ducalis McDunnough,
Euxoa, 45
ducalis Smith, Setagrotis, 45, 46
edictalis group, Euxoa, 50
flutea Rindge, Euxoa, 37
flutea Smith, Euxoa, 37, 38
foeminalis (Smith), Euxoa, 11, 14, 17, 20,
22, 23, 65
foeminalis Draudt, Euxoa, 20
foeminalis Dyar,
Paragrotis, 20
foeminalis Hampson,
Euxoa, 20
foeminalis Rindge,
Carneades, 20
foeminalis Smith,
Carneades, 20
furtiva Barnes and
McDunnough, Euxoa, 21
furtiva Draudt, Euxoa, 21
furtiva Hampson, Euxoa, 21
furtivus Dyar, Paragrotis, 21
furtivus Smith, Agrotis, 21
furtivus Smith, Carneades, 21
idahoensis (Grote), Euxoa, 10, 15, 18, 20,
21,22, 65
idahoensis Dyar,
Paragrotis, 21
idahoensis Grote, Agrotis, 21
idahoensis Hampson,
Euxoa, 21
idahoensis Smith,
Carneades, 21
infracta (Morrison), Euxoa, 12, 18, 23,
35, 36, 40, 66
infracta Draudt, Euxoa, 35
infracta Dyar, Paragrotis, 35
infracta Hampson, Euxoa, 35
infracta Morrison, Agrotis , 35
infracta Morrison,
Carneades , 35
infracta Smith, Carneades, 35
inscripta Lafontaine, new
species, Euxoa, 11, 13, 14, 16, 38, 39,
40
intrita group, Euxoa, 54, 62
laetificans (Smith), Euxoa, 11, 12, 18, 36,
37,38,64
laetificans Draudt, Euxoa, 36
laetificans Dyar,
Paragrotis, 36
laetificans Hampson,
Euxoa, 36
laetificans Smith,
Carneades, 36
latro (Barnes and
Benjamin), Euxoa, 11, 13, 17, 33, 34,
35.66
latro Barnes and Benjamin,
Agrotis, 33
luctuosa group, Euxoa, 50, 63
luteola Barnes and
McDunnough, Euxoa, 32
luteola Dyar, Paragrotis, 32
luteola Hampson, Euxoa, 32
luteola Smith, Agrotis , 32, 33
luteola Smith, Carneades, 32
masculinus Barnes and
McDunnough, Euxoa, 36
masculinus Rindge,
Carneades, 36
masculinus Smith,
Carneades, 36
mcdunnoughi Cook, Euxoa, 27
mcdunnoughi
McDunnough, Euxoa, 27
melura McDunnough,
Euxoa, 10, 11, 15,42,43,64
mitis group, Euxoa, 50
moerens (Grote), Euxoa, 11, 13, 17, 32,
33.34.35.66
moerens Draudt, Euxoa, 32
moerens Dyar, Paragrotis, 32
moerens Grote, Carneades, 32, 33
moerens Hampson, Euxoa, 32
moerens Smith, Carneades, 32
murdocki (Smith), Euxoa, 11, 12, 17, 34,
35.66
murdocki Draudt, Euxoa, 34
murdocki Dyar, Paragrotis, 34
Quaest. Ent., 1981, 17 (1,2)
Lafontaine
118
murdocki Hampson,
Euxoa, 34
murdocki Smith, Agrotis , 34
murdocki Smith,
Carneades, 34
murdocki Smith, Carneaes, 34
neomexicana Hampson,
Euxoa , 45
neomexicana Lafontaine,
Euxoa , 45
neomexicana Rindge,
Carneades , 45
neomexicana Smith,
Carneades , 45, 46
neomexicanus Dyar,
Par agrotis, 45
niveiline Hampson, Euxoa , 40
niveilinea (Grote), Euxoa, 10, 11, 13, 16,
40,41,42, 46, 69
niveilinea Barnes and
McDunnough, Euxoa, 41
niveilinea Draudt, Euxoa, 41
niveilinea Dyar, Paragrotis, 40
niveilinea Grote, Agrotis, 40, 41
niveilinea rabiata Draudt,
Euxoa, 41
niveilinea rabiata
McDunnough, Euxoa, 41
niveilinea Smith,
Carneades, 40
numatana Smith,
Carneades, 42
obeliscoides group, Euxoa, 62
oblongistigma (Smith),
Euxoa, 11,16,27,28,29,66
oblongistigma Draudt,
Euxoa, 29
oblongistigma Dyar,
Paragrotis, 28
oblongistigma Hampson,
Euxoa, 28
oblongistigma Smith,
Agrotis, 28, 29
oblongistigma Smith,
Carneades, 28
olivalis (Grote), Euxoa, 11, 16, 21, 27, 28,
29, 39, 42, 43, 66
olivalis Draudt, Euxoa, 27
olivalis Dyar, Paragrotis, 27
olivalis Grote, Agrotis, 27
olivalis Hampson, Euxoa, 27
olivalis Smith, Carneades, 27
olivia group, Euxoa, 54,61
orosagrotis group, Euxoa, 50
orthogonia Morrison,
Agrotis, 5
pallipennis group, Euxoa, 54, terrena
group, Euxoa
pedalis Draudt, Euxoa, 31
pedalis Dyar, Paragrotis, 3 1
pedalis Hampson, Euxoa, 31
pedalis Lafontaine, Euxoa, 31
pedalis Smith, Carneades, 31, 32
personata Dyar, Paragrotis, 43
personata Forbes, Agrotis, 43
personata Grote, Agrotis, 43
personata Hampson,
Euxoa, 43
personata Morrison,
Agrotis, 43, 44, 45
personata Smith,
Carneades, 43
personata Smith, Euxoa, 43
pity chorus Grote, Agrotis, 43, 44
pity chorus Smith,
Carneades, 43
pitychrous Grote, Agrotis, 45
pitychrous Smith,
Carneades, 43
plagigera (Morrison),
Euxoa, 69
plagigerra (Morrison),
Euxoa, 27
postmedialis Draudt,
Euxoa, 30
pugionis Barnes and
McDunnough, Euxoa, 37
pugionis Dyar, Paragrotis, 37
pugionis Hampson, Euxoa, 37
pugionis Rindge,
Carneades, 37
pugionis Smith, Carneades, 37, 38
The Euxoa detersa group (Lepidoptera: Noctuidae)
119
punctigera, 10
quadridentata (Grote and
Robinson), Euxoa, 10, 11, 13, 15, 37,
38
quadridentata Draudt,
Euxoa , 37
quadridentata Dyar,
Para grot is , 37
quadridentata flutea Barnes
and McDunnough,
Euxoa , 37
quadridentata flutea
Draudt, Euxoa , 37
quadridentata flutea Smith,
Euxoa, 1 1, 38
quadridentata Grote and
Robinson, Agrotis, 37, 38
quadridentata Hampson,
Euxoa, 37
quadridentata
quadridentata (Grote
and Robinson), Euxoa, 1 1, 38
quadridentata Smith,
Carneades , 37
rabiata Forbes, Agrotis, 41
rabiata Lafontaine, Euxoa, 41
rabiata Rindge, Euxoa, 41
rabiata Smith, Euxoa, 41
rabita Smith, Euxoa, 41
recticincta Draudt, Euxoa, 3 1
recticincta Dyar,
Paragrotis, 31
recticincta Lafontaine,
Euxoa, 31
recticincta Smith,
Carneades, 31,32
recticinta Hampson, Euxoa, 3 1
recula (Harvey), Euxoa, 11, 12, 15, 46,
47,48
recula Dyar, Paragrotis, 47
recula Hampson, Euxoa, 47
recula Harvey, Agrotis, 47
recula Lafontaine, Euxoa, 47
recula Smith, Carneades, 47
redimacula Morrison,
Agrotis, 24
redimicula (Morrison),
Euxoa, 11, 13, 16,24,25,26,65
redimicula Barnes and
McDunnough, Euxoa, 23
redimicula Dyar,
Paragrotis, 24
redimicula Forbes, Agrotis, 25
redimicula Hampson,
Euxoa, 24
redimicula Lafontaine,
Agrotis, 25
redimicula Lafontaine,
Euxoa, 25
redimicula McDunnough,
Euxoa, 25
redimicula Morrison,
Agrotis, 24, 25
redimicula Smith,
Carneades, 24
refula group, Euxoa, 62
ridingsiana group, Euxoa, 63
rufula group, Euxoa, 54, 62
rumatana Barnes and
McDunnough, Euxoa, 42
rumatana Rindge,
Carneades, 42
rumatana Smith,
Carneades, 42
septentrionalis group,
Euxoa, 61
serotina Lafontaine, Euxoa, 54
serricornis group, Euxoa, 54, 62
servita (Smith), Euxoa, 10, 11, 14, 15, 23,
24, 25, 26, 65
servita Dod, Euxoa, 23
servita Forbes, Agrotis, 23
servita Hampson, Euxoa, 23
servita Lafontaine, Euxoa, 23
servita McDunnough,
Euxoa, 23
servita novangliae
McDunnough, Euxoa, 23, 24
servitus Dyar, Paragrotis, 23
servitus Rindge, Carneades, 23
servitus Smith, Carneades, 23, 24
setonia group, Euxoa, 62
Quaest. Ent., 1981, 17(1,2)
120
Lafontaine
simulata group, Euxoa, 62
teleboa (Smith), Euxoa, 11, 12, 16, 21,
31.32
teleboa Draudt, Euxoa, 31
teleboa Dyar, Paragrotis, 3 1
teleboa Hampson, Euxoa, 31
teleboa Smith, Carneades , 31, 32
teplia Barnes and
McDunnough, Euxoa, 45
teplia Rindge, Euxoa, 45
teplia Smith , Euxoa, 46
teplia Smith, Euxoa, 45
terrena group, Euxoa, 54
tessellata group, Euxoa, 54, 62
tronella (Smith), Euxoa, 11, 13, 16, 29,
30.31.32
tronella Hampson, Euxoa, 30
tronellus Barnes and
McDunnough, Euxoa, 30
tronellus Draudt, Euxoa, 30
tronellus Rindge,
Carneades, 30
tronellus Smith, Carneades, 30, 31
unica McDounnough,
Euxoa, 11, 16, 39, 40, 64
westermanni group, Euxoa, 54
wilsoni group, Euxoa, 63
REDESCRIPTION AND SYSTEMATIC PLACEMENT OF OREADOMYIA ALBERTAE
KEVAN AND CUTTEN-ALI-KHAN (DIPTERA: CHIRONOMIDAE)
D. R. Oliver
Biosystematics Research Institute
Agriculture Canada
Ottawa, Ontario, Canada K1A 0C6 Quaestiones Entomologicae
17:121-128 1981
ABSTRACT
The female of a small micropterous fly, Oreadomyia albertae Kevan and Cutten-Ali-Khan,
is redescribed. Based on attributes of the female terminalia it is transferred from the
Nymphomyiidae to the Chironomidae. Division of gonapophysis VIII into three lobes, absence
of an anal point from sternum VIII, flagellum with less than 14 flagellomeres and absence of
setae from the postnotum places it in the subfamily group Chironominae + Orthocladiinae +
Prodiamesinae + Diamesinae. Although no synapomorphies could be found Oreadomyia is
tentatively placed in the Orthocladiinae.
RESUME
On redecrit la femelle de la petite mouche microptere Oreadomyia albertae Kevan and Cutten-Ali-Khan. Les
caracteristiques des terminalia de la femelle font passer cette mouche des Nymphomyiidae aux Chironomidae. La
division de la gonapophyse Vlll en trois lobes, I’absense d’une pointe anale sur le sternite VIII, la presence d un
flagellum comportant moins de 14 flagellomeres et V absences des soies sur le postnotum suggerent de classer cet insecte
dans le groupe de sous-familles siuvantes: Chironominae + Orthocladiinae + Prodiamesinae + Diamesinae. Bien qu’on
ait trouve aueune afinite synapomorphique pour un groupe determine, on range provisoirement Oreadomyia parmi les
Orthocladiinae.
122
Oliver
1
Fig. I . Oreadoniyia albert cie, lateral view of female.
Oreadomyia albertae Kevan and Cutten-Ali-Khan
123
Kevan and Cutten-Ali-Khan (1975) described a small micropterous fly, Oreadomyia
albertae , and placed it in the family Nymphomyiidae. Only a single female is known. This
insect is obviously incapable of flight and exhibits many modifications in head and thoracic
structures generally associated with flightless behaviour. Many characters, such as wing
venation, normally used for systematic placement, are unavailable. The terminalia appear
relatively unmodified and Saether (1977) has recently demonstrated the usefulness of this
structure in determining systematic position. The structure of the terminalia shows that
Oreadomyia belongs in the Chironomidae. This systematic placement, and its position within
the Chironomidae, is disscussed here. A redescription of the holotype, augmenting the original
description of Kevan and Cutten-Ali-Khan (1975), is given also.
The type specimen, originally preserved in ethanol, has been mounted in Canada Balsam on
a microscope slide. Terminology of the female terminalia follows that of Saether (1977).
Redescription of Oreadomyia albertae Kevan and Cutten-Ali-Khan
Female (Fig. 1)
Antenna. Three-segmented. Scape large, saucer-shaped, diameter subequal to that of pedicel; without setae. Pedicel
sub-globular with an apical cup-shaped depression; without setae. Flagellum (Fig. 2) undivided, clavate with a narrow,
short base inserted into cup-shaped depression of pedicle; with two longitudinal rows of five sensilla basiconica each; setae
short, basally in two irregular rows; apex without strong seta.
Head. Rounded with dome-shaped vertex, small elipsoidal eyes and short mouthparts. Vertex with six or seven setae
per side; coronal suture incomplete; coronal triangle absent. Individual ommatidia of eye enlarged. One darkly pigmented
larval eye (stemmatic bulla), without ommatidia, lying posteroventral to eye. Clypeus (Fig. 3) subtriangular, without
setae. Maxillary palpus (Fig. 3) with three discernable segments.
Thorax. Broad antepronotal lobes completely fused medially; one ventrolateral seta present. Limits of dorsally
flattened scutum, scutellum, and postnotum obscure, but transverse row of four setae may be scutellar setae; acrostichal
and supra-alar setae absent; one seta present on each side of scutum midway between antepronotum and transverse row of
setae, may be dorsocentral setae. Anepisternum and katepisternum compressed by enlarged coxae. Anapleural suture not
evident. Anterior spiracle appearing to lie in anterior part of anepisternum not in anepisternal cleft. Halter absent, but
metanotum with low protuberance in usual position of halter base.
Wing. Narrow, strap-like, and shorter than one-half of length of thorax. Venation not discernable except for traces of
radial vein evident in basal third.
Legs. Long, relatively thick with enlarged coxae. Fore L.R. ca. 0.58. One short, straight tarsal spur on each leg; basal
one-third to one-half of each spur encircled with fine spines; hind tibial comb absent. Fourth tarsomere slightly widened
apically, shorter than length of fifth tarsomere. Claws apically pointed; empodium plumose; pulvilli absent.
Abdomen. Segments I-V somewhat dorsoventrally flattened, remaining segments cylindrical and progressively larger
posteriorly.
Quaest. Ent., 1981, 17(1,2)
124
Oliver
Figs. 2-6. Oreadomyia albertae , female. Fig. 2, flagellum. Fig. 3, maxillary palpus and clypeus. Fig. 4, ventral views
of terminalia,- csa: coxosternapodeme IX, no: notum, r: ramus, spt: spermatheca. Fig. 5, dorsal view of tergum, IX.
Fig. 6, ventral view of gonapophysis VIII,- apl: apodeme lobe, dml: dorsomedial lobe; VII: ventrolateral lobe.
Oreadomyia albertae Kevan and Cutten-Ali-Khan
125
Terminalia (Fig. 4). Gonapophysis VIII divided; ventrolateral lobe large, rounded, with moderately long setae;
dorsomedial lobe narrow; apodeme lobe distinct, without microtrichia (Fig. 6). Coxosternapodeme IX curved.
Notum extending anteriorly into abdominal segment VII; rami diverging around anterior part of genital cavity.
Tergum IX undivided with caudomesal projection; setae in two groups, one on either side of narrow bare median
area (Fig. 5); about one and one-half times as long as gonocoxite IX. Gonocoxite IX setiferous. Segment X without
setae. Postgenital plate weak. Cercus one-segmented; pediform. Two ovoid spermathecae; without microtrichia.
Holotype female
Bald Hills, Jasper National Park, Alberta (52° 43' N, 117° 41’ W; elev. 7200’), 19-31 Aug.
1970, P. Kuchar; Type No. 16041, deposited in Canadian National Collection, Ottawa.
Remarks. - Oreadomyia albertae resembles the antarctic midge, Belgica antarctica
(Jacobs) - compare Fig. 1 with fig. IE in Wirth and Gressitt (1967). Both have narrow
strap-like wings, no halteres, dorsally flattened thorax, enlarged coxae and strong legs, and a
rounded head with small ellipsoidal eyes and reduced ‘mouthparts. They differ in antennal
structure. The female of Belgica has four flagellomeres whereas Oreadomyia has only one.
They also differ in structure of the terminalia, notably in that Belgica lacks a well defined
apodeme lobe.
A double row of five sensilla basiconica suggests that the flagellum of Oreadomyia
originated by fusion of six flagellomeres. In other female chronomids the apex of each
flagellomere, except the terminal one, bears a pair of sensilla basiconica. The terminal
flagellomere also bears sensilla basiconica, but these are arranged in one row or are scattered.
The apical spine-like seta described by Kevan and Cutten-Ali-Khan (1975; figs. 1C and ID) is
not present. They also considered the flagellum to be two-segmented. The second being a “very
small, apparently undivided, finger-like, subapical terminal segment.” This could be a
misinterpretation of a sensillum basiconicum. Kevan and Cutten-Ali-Khan (1975; figs. 1C and
ID) described the eye as being composed of two parts, each with ommatidia. The
posteroventral part is composed only of pigment and is clearly without ommatidia. It does not
appear to be part of the adult eye but a remnant of the larval eye spot, and therefore not a
vestige of the ventral eye-bridge found in the Nymphomyiidae.
SYSTEMATIC POSITION
Kevan and Cutten-Ali-Khan placed Oreadomyia in the Nymphomyiidae “principally on the
basis of the characteristic form of the antennae”. The shape of the ommatidia and general
habitus were also considered. The antenna of Oreadomyia and those of some Nymphomyiidae,
e.g., Felicitomyia and Palaeodipteron, are similar in that the flagellum is club-shaped.
However, the flagellum of Nymphomyiidae has an apical spine-like seta, a subterminal group
of sensilla, and an annulated basal part. Oreadomyia lacks the spine-like seta, the sensilla
basiconica are arranged in two rows, and the flagellum consists of a single undifferentiated
flagellomere. Many other differences in detail between the Nymphomyiidae and Oreadomyia
also occur in other head and thoracic structures (see comparison given in Kevan and
Cutten-Ali-Khan 1975). Also the female terminalia differ fundamentally. Cerci are
two-segmented and gonapophysis VIII is long and undivided in the known Nymphomyiidae
(Saether 1977). In Oreadomyia the cerci are one-segmented and gonapophysis VIII is
relatively short and tri-lobed.
Quaest. Ent., 1981, 17 (1,2)
126
Oliver
Attainment or transition to flightlessness in chironomids and other nematocerous Diptera
frequently is accompanied by modifications of head and thoracic structures (Brundin 1966;
Downes 1969; Hackman 1964; Hansen and Cook 1976; Hashimoto 1962). These structural
modifications are usually reductions, e.g., male antenna becoming female-like, reduced eyes,
brachyptery and flattening of the dorsum of the thorax. Legs are often strengthened and the
male terminalia may be enlarged and heavily sclerotized, but the female terminalia are
relatively unmodified. In cases of extreme reduction many characters normally used for
systematic placement are unavailable.
The female of Oreadomyia is flightless. Newly emerged nymphomyiid adults are fully
winged (Kevan and Cutten-Ali-Khan 1975) but the wings are apparently deciduous (Cutten
and Kevan 1970) and some activity occurs in a flightless state. Similarities between some of the
apomorphies occurring in the two groups are probably due to convergence resulting from loss or
reduction in flight activity. Differences in detail of these apomorphies suggest that either the
ancestral condition was different or the reduction process followed different pathways.
Apomorphies that arise by loss or reduction have limited value in determining phyletic
relationshp because of difficulty in recognizing convergence (Brundin 1976), especially if the
selection of the apomorphies correlates with a particular mode of behaviour such as attainment
or transition to flightlessness. Evidence of relationships should be sought in complex structures,
such as female terminalia. Division of gonapophysis VIII into three lobes is a synapomorphy
which places Oreadomyia within the Chironomidae. This apomorphy does not occur in any
other family of Diptera and within the Chironomidae it only occurs in the Buchonomyiinae and
its sister group, Chironomidae + Orthocladiinae +Prodiamesinae + Diamesinae (Saether
1977; Brundin and Saether 1978).
The presence of several apomorphies shows that Oreadomyia cannot belong to the
Buchonomyiinae. The flagellum of the Buchonomyiinae has 14 flagellomeres and less than 14
flagellomeres is synapomorphic for its sister group (Brundin and Saether 1978). Use of this
synaphomorphy is admittedly tenuous because of the highly reduced state of the antenna of
Oreadomyia. However, there is evidence that the extreme apomorphic state of the antenna
arose by fusion of six flagellomeres (see Remarks). The postnotum of Oreadomyia lacks setae
which is a synapomorphy for the subfamily group Chironomidae + Orthocladiinae +
Prodiamesinae + Diamesinae (Brundin and Saether 1978). These setae are present, the
plesiomorphic character state, in the Buchonomyiinae. Also the presence of an anal point on
sternum VIII is synapomorphic for the Buchonomyiinae. The plesiomorphic character state,
lack of this anal point, exists in Oreadomyia , as well as in the subfamily group Chironomidae
+ Orthocladiinae + Prodiamesinae + Diamesinae.
Placement within the subfamily group Chironomidae + Orthocladiinae + Prodiamesinae
+ Diamesinae is difficult because of the extreme reduction of the female of Oreadomyia and
lack of information about male and immature stages. It keys to couplet 17, containing the
Orthocladiinae and Chironomidae, in the key to subfamilies and tribes of Chironomidae by
Saether (1977). Of the three characters given in couplet 17 only the third, the tarsal spurs, is of
use in placing Oreadomyia. They are clearly of the type occuring in the Orthocladiinae,
eliminating the Chironomidae from contention. The structure of tergum IX does not fit any of
the alternatives given in couplet 17 (Saether 1977) and a bare segment X occurs in both
subfamilies. Interpretation of tergum IX character is ambiguous. However, it should be noted
that although the structure of tergum IX is not clearly either Orthocladiinae or Chironomidae
it resembles that of Diplocladius, a member of the Orthocladiinae.
Oreadomyia albertae Kevan and Cutten-Ali-Khan
127
It is not possible to present phyletic evidence in support of the placement of Oreadomyia in
the Orthocladiinae, as no synapomorphies were found to conclusively eliminate the
Prodiamesinae and Diamesinae. The Diamesinae is regarded as the sister group of the
Chironomidae + Orthocladiinae + Prodiamesinae, and the Prodiamesinae the sister group of
the Chironomidae + Orthocladiinae (Saether 1977). The female of Oreadomyia has a long
notum. This is regarded as synapomorphic for Chironomidae + Orthocladiinae +
Prodiamesinae (Saether 1977:31), which would suggest elimination of the Diamesinae.
However, the dividing point between a long and a short notum is arbitrary and Saether
(1977:60) later states that the notum in the Diamesinae is “short to long”. Equally
unsatisfactory is the use of the rami to distinguish the Prodiamesinae. According to Saether
(1977) the presence of parallel-sided rami is synapomorphic for the Prodiamesinae. But this
character is not shared by all Prodiamesinae as the rami of Odontomesa are not parallel-sided
(see Fig. 35C in Saether 1977). Saether (1977) has provided a basis for the study of female
chironomids, however, more detailed studies are required before the phyletic relationships of
reduced forms such as Oreadomyia can be deduced solely from female terminalia.
ACKNOWLEDGEMENTS
I wish to thank A. Borkent, J.F. McAlpine and D.M. Wood for discussion and critical
comments on the manuscript. Fig. 1 was drawn by R. Idema.
REFERENCES
Brundin, L. 1966. Transantarctic relationships and their significance as evidenced by
chironomid midges. With a monograph of the subfamilies Podonominae and
Aphroteniinae and the austral Heptagyiae. Kungliga Svenska Vetenskapsakademiens
Handlingar. (4) 11: 1-472.
Brundin, L. 1976. A Neocomian chironomid and Podonominae-Aphroteniinae (Diptera) in the
light of phylogenetics and biogeography. Zoologica Scripta. 5: 139-160.
Brundin, L., and O.A. Saether. 1978. Buchonomyia burmanica sp. n. and Buchonomyiinae, a
new subfamily among the Chironomidae (Diptera). Zoologica Scripta. 7: 269-275.
Cutten, E.A., and D.K.McE. Kevan. 1970. The Nymphomyiidae (Diptera), with special
reference to Palaeodipteron walkeri Ide and its larva in Quebec and a description of a
new genus and species from India. Canadian Journal of Zoology 48: 1-24.
Downes, J.A. 1969. The swarming and mating flight of Diptera. Annual Review of
Entomology. 14: 271-298.
Hackman, W. 1964. On reduction and loss of wings in Diptera. Notulae Entomologicae. 44:
73-93.
Hansen, D.C., and E.F. Cook. 1976. The systematics and morphology of the Nearctic species of
Diamesa Meigen, 1835 (Diptera: Chironomidae). Memoirs of the American
Entomological Society. 30: 1-203.
Hashimoto, E. 1962. Ecological significance of the sexual dimorphism in marine chironomids.
Science Reports of the Tokyo Kyoiku Daigaku. 10: 221-252.
Kevan, D.K.McE., and F.E.A. Cutten-Ali-Khan. 1975. Canadian Nymphomyiidae (Diptera).
Quaest. Ent., 1981, 17(1,2)
I
128 Oliver
Canadian Journal of Zoology. 53: 853-866.
Saether, O.A. 1977. Female genitalia in Chironomidae and other Nematocera: morphology,
phylogenies, keys. Bulletin of the Fisheries Research Board of Canada. 197: 1-209.
Wirth, W.W., and J.L. Gressitt. 1967. Diptera: Chironomidae (Midges) pp. 197-203. In J.L.
Gressitt (ed.) Entomology of Antarctica. American Geophysical Union. Antarctic
Research Series. 10.
FUNCTION OF CAUDAL FILAMENTS AND CORRELATED STRUCTURES IN
MAYFLY NYMPHS, WITH SPECIAL REFERENCE TO BAETIS (EPHEMEROPTERA)
Lynda D. Corkum'
Department of Zoology
The University of Alberta
Edmonton, Alberta, Canada
T6G 2E9
Quaestiones Entomologicae
17:129-146 1981
Hugh F. Clifford
Department of Zoology
The University of Alberta
Edmonton, Alberta, Canada
T6G 2E9
ABSTRACT
Laboratory and field experiments demonstrate that caudal filaments of Baetis tricaudatus
and Rhithrogena hageni nymphs do not effectively deter stonefly predators or aid in molting.
A field study along a section of a Rocky Mountain stream shows that length of the terminal
filament in Baetis nymphs is correlated with a combination of factors and not specifically to
current velocity.
Subapical setae on the claws of Baetis bicaudatus and B. tricaudatus nymphs are probably
mechanoreceptors which may be used to detect subtle changes in flow and thus assist in
orienting nymphs into the current. All species of Baetis nymphs with subapical setae on claws
also have a reduced terminal filament and inhabit fast water areas.
RESUME
Des experiences en laboratoire et en nature demontrent que les filaments caudaux des nymphes de Baetis tricaudatus
et de Rhithrogena hageni ne facilitent pas la mue de ces Plecopteres, ni ne decouragent leurs predateurs. Une etude en
nature, realisee le long d’une section d’un ruisseau des Montagues Rocheuses, revele que la longueur du filament
terminal des nymphes de Baetis est correlee a un ensemble de facteurs, et non specifiquement a la vitesse du courant.
Les soies subapicales portees par les griffes des nymphes de Baetis Bicaudatus et de B. tricaudatus sont probablement
des mecanorecepteurs qui peuvent etre utilises pour detecter de legers cjamgements de debit du courant, et ainsi aider les
nymphes a s’orienter dans le scourant. Toutes les especes de Baetis dont Is nymphes possedent des soies subapicales sur
les griffes, presentent aussi un filament terminal reduit, et frequentent les zones a courant rapide.
‘Present address: Aquatic Environments Ltd., 1235A - 40th Avenue N.E., Calgary,
Alberta, Canada T2E 6M9
130
Corkum and Clifford
TABLE OF CONTENTS
INTRODUCTION
Our objective is to study numerous functions that have been attributed to mayfly caudal
filaments by examining: (1) success of a stonefly predator feeding on two species of mayfly
nymphs with and without (excised) caudal filaments; (2) molting rate of these two species with
and without caudal filaments and time required to generate filaments; (3) longitudinal
distribution of two- and three-tailed Baetis nymphs in a Rocky Mountain stream of Alberta,
Canada; (4) whether there is any difference between two and three-tailed Baetis nymphs in
avoiding a stonefly predator in this stream; and (5) relationships between reduction of the
middle (terminal) caudal filament in Baetis nymphs and corresponding changes in claw
structure.
Most mayfly nymphs possess three caudal filaments. However, of 60 North American
mayfly genera, seven (Baetidae: Baetodes Needham and Murphy, Heterocloeon McDunnough,
Pseudocloeon Klapalek; Oligoneuriidae: Lachlania Hagen; Heptageniidae: Epeorus Eaton,
Ironodes Traver, Spinadis Edmunds and Jensen) possess two lateral filaments and lack the
terminal filaments as nymphs (Edmunds et al., 1976).
Within the genus Baetis , relative length of the terminal filament to that of the lateral
filaments varies greatly in nymphs. There is only one representative, B. hudsonicus , in which
the terminal filament is similar in length to the lateral filaments. In all other species, nymphs
possess a terminal filament that is shorter than the lateral filaments. Of the 24 described North
American species of Baetis nymphs, seven ( B . lapponicus gp -B. amplus, B. insignificans, B.
lapponicus', B. rhodani gp -B. bicaudatus , B. foemina ) possess a terminal filament that has
been reduced to five segments or less (Morihara and McCafferty 1979).
Baetid nymphs possessing three well-developed caudal filaments with overlapping setae or
hairs are well suited for swimming away from and settling onto the substrate (Percival and
Whitehead 1929, Hughes 1966, Corkum and Clifford 1980). Although Baetis nymphs with a
reduced terminal filament occur in fast flowing water, the adaptive significance of two lateral
filaments and a reduced terminal filament is unclear.
Dodds and Hisaw (1924) and Hora (1930) related degree of reduction of the terminal
filament in mayfly nymphs with increased current velocity in headwater regions of streams. We
anticipated that if Baetis nymphs with two rather than three caudal filaments were better
adapted to fast flowing water, then these nymphs might also have some morphological
Caudal filaments in Mayfly nymphs
131
modification of claws enabling them to maintain their position on substrates.
Hynes (1970) was not convinced that reduction of the terminal filament was necessarily a
morphological adaptation to fast water. Nymphs of one African genus, Dicercomyzon
(Tricorythidae) (flattened body with an adhesive pad and no terminal filament), are atypical as
they inhabit slow water (Venter 1961). Hynes suggests that these features of Dicercomyzon
nymphs enable them to adhere to vegetation and avoid predation.
We have collected many mayfly nymphs with caudal filaments broken off. These
appendages may aid nymphs in avoiding predators, i.e. the predator seizes a caudal filament,
the prey escapes and is able to regenerate the damaged filaments (e.g. Schmidt 1951).
MATERIALS AND METHODS
Laboratory Studies
Caudal filaments and vulnerability in predation.
Laboratory studies were conducted to determine feeding success of nymphs of a stonefly
predator, Isogenoides elongatus (Hagen), on Rhithrogena hageni Eaton and Baetis
tricaudatus Dodds nymphs with caudal filaments and with filaments excised. Nymphs were
collected from the Pembina River, near Entwistle, Alberta (53°36’N, 115°00’W). River water
temperature was 5°C; this temperature was maintained in an environmental chamber in the
laboratory. The mayfly nymphs were of developmental stage II (Clifford 1970) and were 2
( + /- caudal filaments) x 2 ( + /- predator) x 2 ( + /- light) factorial therefore relatively small.
Each of 40 experimental cells consisted of 200 ml of aerated dechlorinated water in a glass
bowl (10.5 cm diam. x 4 cm depth), containing a stone and 1 g of prepared instant oatmeal.
Experiments were conducted using a 2 ( + /- caudal filaments) x 2 ( + /- predator) x 2 ( + /-
light) factorial design for each mayfly species with five trials for each condition and a 12 hr
light-dark cycle. All three caudal filaments of experimental nymphs were cut to a length
equaling the width of the 10th abdominal segment. Stonefly nymphs were starved for 24 hr
before the experiment. There were five prey and one predator (when present) per bowl.
Numbers of mayfly nymphs present were recorded each day at the beginning and end of the
light phase and then, if necessary, prey were replenished. After five days, insects were
preserved and foreguts of predators were analyzed.
A student’s t-test was used to determine if there was a difference in the number of prey
eaten by predators during light and dark phases. Two-way analysis of variance, was used to
determine influence of the predator and existence of caudal filaments on two mayfly species in
light and dark.
Caudal filaments and molting rate
A second experiment compared molting rates between R. hageni and B. tricaudatus nymphs
with and without caudal filaments (n= 16-19 nymphs). Drenkelfort (1910) suggested that
“pulling” movement of filaments was mainly responsible for the nymph escaping from its old
exoskeleton at ecdysis. Each of the four experimental cells consisted of a glass bowl (20 cm
diam. x 6 cm depth) containing 1 liter of aerated dechlorinated water, 10 stones, TetraMin food
Quaest. Ent., 1981, 17 (1,2)
Table 1. Distribution of Baetis nymphs collected at sites along the Highwood River, 24 June 1979.
132
Corkum and Clifford
03
3
3
"3
3
3
3
<N <N
ov O CO
^ co — '
(N
Size categories (I, II, III, IV) are based on wing-pad development ( after Clifford 1 970).
Caudal filaments in Mayfly nymphs
133
and mayfly nymphs. Caudal filaments were cut as previously described. Temperature was
constant at 5°C and a 12 hr light-dark cycle was used. Numbers of mayfly exuviae were
recorded and preserved daily for 50 days. For nymphs with excised filaments, we predicted
regeneration time of the filaments by plotting increase in length of caudal filaments against
time.
FIELD STUDIES
Caudal filaments and longitudinal distribution in a stream.
A field study was conducted at six sites along a section of the Highwood River (Table 1) in
the Rocky Mountain Forest of Alberta to determine distribution of baetid nymphs having two
(B. bicaudatus) and three caudal filaments ( B . tricaudatus). Insects were collected by taking a
1 min. kick sample (mesh opening: 180 microns) from a riffle area at each site.
The following parameters were recorded at the time each site was sampled: land-use,
aquatic, overhanging and riparian vegetation, current velocity (Gurley Pigmy Meter) stream
width and maximum depth, bottom type, air and water temperature, pH and dissolved oxygen.
Altitude, distance from stream source and stream order were obtained from topographic maps.
Water samples were immediately refrigerated and additional chemical characteristics analyzed
1 week later using standard methods (American Public Health Association 1971). Gower’s
similarity coefficient (Sneath and Sokal 1973) was used to determine similarity between stream
sites based on 3 1 physical and chemical parameters.
Caudal filaments and vulnerability to predation.
An in situ experiment was conducted on the Highwood River using flow-through plastic
containers (28.5 cm x 17 cm x 13 cm) with and without the stonefly predator Hesperoperla
pacifica (Banks) and either two-tailed (B. bicaudatus) or three-tailed ( B . tricaudatus ) nymphs.
Baetis bicaudatus Dodds and B. tricaudatus nymphs were collected from site 2 (upstream) and
site 6 (downstream), respectively (Table 1). The experiment was conducted midway between
the two collecting sites. Twelve 2 cm holes were drilled into both ends of each container.
Netting (mesh opening: 180 microns) was secured over the holes to exclude extraneous
organisms and prevent experimental organisms from escaping. Each container was half filled
with clean stones from the stream bed. Predators had been starved for 4 hrs before the
experiment. Fifteen mayflies (developmental stage III) representing two- or three-tailed forms
and one predator were added to the containers (10 trials). There were two control replicates (no
predators) for each mayfly species (total number of experimental cells = 24). After 24 hrs, the
containers were retrieved and the remaining nymphs collected. Foreguts of the stonefly nymphs
were analyzed for prey remains. A one-way analysis of variance was used to test for differences
between species remaining in the experimental cells.
Morphological Studies
Claws of B. bicaudatus and B. tricaudatus nymphs were observed with a scanning electron
microscope (SEM). To prepare the dissected claws for observation, they were first sonicated in
70% ethanol to remove debris. The claws were dehydrated in a graded ethanol series,
transferred to absolute acetone and allowed to dry. After the claws were mounted on stubs with
Quaest. Ent., 1981,17(1,2)
(
134 Corkum and Clifford
Table 2. Number of B. tricaudatus and R. hageni nymphs (with and without caudal
filaments) eaten by Isogenoides elongatus nymphs in the light and dark
32
4
59
19
Caudal filaments in Mayfly nymphs
135
Table 3. Significance of effects from a 2-way ANOVA test of caudal filaments (C.F.)
and predators on number of mayfly nymphs taken of each species in the light
*** p < 0.001
Quaest. Ent., 1981, 17 (1,2)
CUM. EXUVIAE
136
Corkum and Clifford
Figure 1. Cumulative number of nymphal exuviae of Baetis tricaudatus with (■) and without (□) caudal filaments
and of Rhithrogena hageni with (•) and without (O) caudal filaments over time.
CAUDAL FILAMENT LENGTH (mm)
Caudal filaments in Mayfly nymphs
137
Figure 2. Total length of regenerated caudal filaments for Baetis trieaudatus (■) and Rhithrogena hageni (O)
nymphs versus time.
Quaest. Ent., 1981, 17 (1,2)
138
Corkum and Clifford
conducting tape, and a gold-palladium layer applied to the stub, they were ready for
observation.
RESULTS
Laboratory Studies
Caudal filaments and vulnerability to predation.
Isogenoides elongatus (Hagen) nymphs ate significantly more B. bicaudatus than R. hageni
nymphs in light and dark (Table 2). There were, however, significantly more prey taken by
predators in the dark (Student’s t: p < 0.01). Based on the replicated two-way analysis of
variance (ANOVA) tests, removal of caudal filaments had no effect on the number of R.
hageni nymphs captured (Tables 3 and 4). During the light only, predators caught significantly
more B. tricaudatus nymphs without caudal filaments than with filaments (p < 0.025).
Caudal filaments and molting rate
Removal of caudal filaments had no effect on the molting rate of either mayfly species (Fig.
1). Mortality (7.5 nymphs, S.D. = 3.5) was comparable in both species whether or not
filaments were excised.
To predict regeneration time of the excised filaments, regression equations were calculated
for the increasing caudal filament length (expressed as mm) versus time (days) for the two
species. Logarithmic transformation of the dependent variable gave the best regression fit for
both species (Fig. 2):
B. tricaudatus Y= 1 .056e0023X R2 = 0.865 (1)
R. hageni Y = 0.948eO-°28X R2=0.823 (2)
The longest natural filament length for B. tricaudatus amd R. hageni nymphs representing this
sample population was 4.16 mm and 6.78 mm, respectively. Using these values, the above
equations would predict that regeneration of caudal filaments of B. tricaudatus in 59.6 days
and those of R. hageni in 70.3 days.
FIELD STUDIES
Caudal filaments and longitudinal distribution.
On 24 June 1979, two-tailed Baetis (B. bicaudatus) nymphs were found at all six sites on
the Highwood River (Table 1). Three-tailed forms (B. tricaudatus ) occupied the downstream
three sites, below 1500 m in altitude. Nymphs of both species occupied fast-water areas. There
was no relationship between distance downstream from the headwaters with either mean
current velocity (r = 0.019) or flow 3 cm from the stream bed (r = 0.175). Clearly, the two
Caudal filaments in Mayfly nymphs
139
populations are not separated by water flow.
The dendrogram resulting from the Gower similarity analysis depicts the three upstream
sites as distinct from the downstream sites (Fig. 3). The distribution of B. tricaudatus nymphs
is associated with the downstream grouping. Between sites 3 and 4 (the two groupings),
Etherington Creek enters the Highwood River. Stream width, specific conductance, alkalinity
GOWER COEFFICI ENT
S
I
T
E
Figure 3. Dendrogram showing similarities between physical-chemical characteristics at sites along the Highwood
River. Location of sites are indicated in Table 1 .
Quaest. Ent., 1981, 17 (1,2)
140
Corkum and Clifford
Plate I. Scanning electron photomicrographs of claws and subapical setae of Baetis bicaudatus (A & B) and B
tricaudatus (C & D). Photographs of C & D are of different specimens.
Caudal filaments in Mayfly nymphs
141
Table 5. A summary of Baetis nymphs with a reduced terminal filament and the
existence of subapical setae on the claws. Data obtained from Morihara & McCafferty
(1979), Muller-libenau (1969, 1971, 1974a) and Keffermiiller (1980).
Western Palearctic:
B. alpinus gp:
B. lapponicus gp:
B. lapponicus t.f. = stub +
B. sinaicus t.f. = stub
B. lutheri gp:
(continued on next page)
Quaest. Ent., 1981, 17 (1,2)
142
Corkum and Clifford
Table 5 (continued)
1 stub = a terminal filament of 5 segments or less (Morihara and McCafferty 1979)
2 + = present, - = absent, ? — unknown.
3 (.n) = ratio of length of terminal filament to that of the lateral filaments.
Caudal filaments in Mayfly nymphs
143
!|
ii
and salinity all increase after the confluence of the two streams.
jj Caudal filaments and vulnerability to predation.
There was no mortality or loss of nymphs in control cells of the in situ predator-prey
experiment conducted on the Highwood River. Fewer B. bicaudatus and B. tricaudatus
nymphs were found in cells containing H. pacifica nymphs. Contents of predator foreguts
indicated that nymphs of both mayfly species were eaten without preference. We tentatively
conclude from this short-term study that neither two-tailed nor three-tailed Baetis nymphs had
an advantage in avoiding predators.
Morphological Studies
Morphological studies showed one pair of subapical setae on each B. tricaudatus claw and
as many as three pairs on the claws of B. bicaudatus (Platte 1).
Published records (Table 5) show that all species of Baetis nymphs with subapical setae on
claws possessed a terminal filament which was greatly reduced (in some species to a stub) and
also inhabited fast-flowing waters. In North America, subapical setae on claws has been
reported in two species groups, B. lapponicus and R. hageni (Morihara and McCafferty 1979).
In the western Palearctic, these setae are present in four groups: Baetis, B. lapponicus , Baetis
and B. rhodani Mtiller-Liebenau 1969, 1971, 1974a). Species within these groups live in
running waters at high altitudes and latitudes or on islands (McDunnough 1936,
Miiller-Liebenau 1971, Edmunds et al., 1976, Cobb and Flannagan 1980). Nymphs of the one
northern species, Baetis macani bundyae, that lack subapical setae on claws inhabit ponds
(Lehmkuhl 1973). Miiller-Liebenau (1974b) found subapical setae on nymphal claws of all
three species of Heterocloeon McDunnough ( = Rheobaetis)\ these nymphs also possess a
reduced (one segment) terminal filament and inhabit swift-flowing rivers. We know of no other
mayfly species in which nymphs possess subapical setae on claws.
DISCUSSION
Our laboratory study showed that stonefly predators ate more B. tricaudatus than R. hageni
nymphs. Evidently, the flattened, clinging, R. hageni nymphs were more difficult for the
stonefly nymphs to capture than the rounded, swimming, B. tricaudatus nymphs. This tends to
support the hypothesis of Elynes (1970) that flattened nymphs adhering to substrate surfaces
may offer protection against predation. Fewer Baetis nymphs with caudal filaments were eaten
by predators during light periods. This was a minor advantage since predators fed significantly
more at night.
In the field, there was no significant difference between two- and three-tailed nymphs. (B.
bicaudatus and B. tricaudatus) in avoiding the stonefly nymph, H. pacifica. Based on these
studies, we discard the idea that the major function of the caudal filaments of mayfly nymphs is
to deter predators.
We also concur with Schmidt’s (1951) view that caudal filaments do not assist in the
molting process. Apparently, movement of caudal filaments does not significantly aid nymphs
during molt as suggested by Drenkelfort (1910).
Species replacement along a stream channel is a common phenomenon (Ide 1935, Allan
1975, Hawkes 1975, Ward and Berner 1980). In headwater areas of mountain streams, Dodds
Quaest. Ent., 1981, 17 (1,2)
144
Corkum and Clifford
and Hisaw (1924) and Hora (1930) suggested that species of Baetis with progressively shorter
terminal filaments live in habitats with increasing current velocities. Although both B.
bicaudatus and B. tricaudatus nymphs occupied fast-water areas, we found no relationship
betwen current velocity and stream site inhabited by these two- and three-tailed species of
Baetis. The difference between no current and some current may be of major significance to an
organism, whereas large differences among current velocities may be of minor importance
(Philipson 1954).
Edmunds et al., (1976) and Ward and Berner (1980) have indicated that B. bicaudatus
nymphs live at high altitudes. Ward and Berner (1980) report the upstream limit of B.
tricaudatus nymphs on St. Vrain Creek Colorado ca. 2400 m, several degrees of latitude south
of our site. On the Highwood River, the upstream limit of B. tricaudatus at 1500 m ('$0o23’ N)
corresponds to the entrance of Etherton Creek, In our study, no single environmental factor
influenced the distribution of the two mayfly species along a stream section. Rather, a
combination of factors were correlated with the upstream limit of the three-tailed, B.
tricaudatus nymphs.
Typically, fast-flowing water is well aerated and carries food to aquatic organisms. The
organisms, however, must either avoid the force of the current or be able to maintain their
position. Baetis nymphs frequently occupy the upper substrate surfaces and face into the
current (Dodds and Hisaw 1924). Hora (1930) suggests that nymphs facing into torrential
water must hold the caudal filaments together to aid water flow over their body. He conceives
that the terminal filament may have been “pressed out of existence, metaphorically speaking,
by the pressure of the lateral [filaments].’’ Perhaps caudal filaments act to reduce drag on
nymphs exposed to current flow.
From morphological studies of B. bicaudatus and B. tricaudatus nymphs, we observed
subapical setae on claws of both of these fast-water species. All species of Baetis with subapical
setae on claws also have a reduced terminal filament. We suggest that these structures are
mechanoreceptors. If so, they should be able to detect water-borne vibrations (Mclver 1975).
Although elliptical shape of the socket may restrict the directional movement of setae, it may
be possible that these structures are used by nymphs to detect subtle changes in flow within the
boundary layer ( sensu Ambiihl 1959). The setae may also assist in orienting nymphs into the
current. These structures would be of particular importance to nymphs inhabiting upper
substrate surfaces exposed to flowing water.
Popovici-Baznosanu (1906) suggested that caudal filaments may function as a respiratory
organ. Since Baetis nymphs with a reduced terminal filament appear to be found in specific
habitats (high altitude and latitude), gradual reduction in an appendage may be a response to
lower respiratory demands in these cooler, fast-flowing waters.
ACKNOWLEDGEMENTS
We thank J. J. H. Ciborowski, D. A. Craig, V. Gotceitas, B. K. Mitchell and I.
Muller-Liebenau for their comments on this study. R. L. Baker provided helpful information on
the study site and J. J. H. Ciborowski assisted with field work. We are grateful to R. W.
Mandryk who prepared mayfly specimens from which SEM photographs were taken by G.
Braybrook. Funding was provided by grants from the Boreal Institute for Northern Studies
(Alberta Government Grant-In-Aid program) to L.D.C. and N.S.E.R.C. to H.F.C.
Caudal filaments in Mayfly nymphs
145
LITERATURE CITED
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AMBUHL, H. 1959. Die Bedeutung der Stromung als okologischer Faktor. Schweizerische
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AMERICAN PUBLIC HEALTH ASSOCIATION. 1971. Standard methods for the
examination of water and wastewater. 13th ed. A.P.H.A., New York.
CLIFFORD, H. F. 1970. Variability of linear measurements throughout the life cycle of the
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Entomologist 46: 98-106.
COBB, D. G., and J. F. FLANNAGAN. 1980. The distribution of Ephemeroptera Biology in
Northern Canada, pi 55-166. In Advances in Ephemeroptera. Eds. Flannagan, J. F., and
K. E. Marshall. Plenum Publ. New York.
CORKUM, L. D., and H. F. CLIFFORD. 1980. The importance of species associations and
substrate types to behavioural drift, p. 331-341. In Advances in Ephmeroptera Biology.
Eds. Flannagan, J. F., and K. E. Marshall. Plenum Publ. New York.
DRENKELFORT, H. 1910. Neue Beitrage zur Kenntnis der Biologie und Anatomie von
Siphlonurus lacustris Eaton. Zoologische Jahrbucher (Anatomie) 29: 527-617.
DODDS, G. S., and F. L. HISAW. 1924. Ecological studies of aquatic insects. 1. Adaptations
of mayfly nymphs to swift streams. Ecology 5: 137-148.
EDMUNDS, G. E., Jr., S. L. JENSEN, and L. BERNER. 1976. The mayflies of North and
Central America. University of Minnesota Press, Minneapolis.
HAWKES, H. A. 1975. River zonation and classification, p. 312-374. In River Ecology. Ed. B.
A. Whitton. Blackwell Sci. Publ., Oxford.
HORA, S. L. 1930. Ecology, bionomics and evolution of the torrential fauna, with special
reference to the organs of attachment. Philosophical Transactions of the Royal Society.
B218: 171-282.
HUGHES, D. A. 1966. On the dorsal light response in a mayfly nymph. Animal Behaviour. 14:
13-16.
HYNES, H. B. N. 1970. The Ecology of Running Waters. University of Toronto Press,
Toronto.
IDE, F. P. 1935. The effect of temperature on the distribution of the mayfly fauna of a stream.
Publications of Ontario Fisheries Research Laboratory 50: 1-76.
KEFFERMULLER, M. 1980. Un Changement dans la Taxonomie de la Famille Baetide. p.
115-121. In Advances in Ephemeroptera Biology, eds. Flannagan, J. F,., and K. E.
Marshall. Plenum Publ. New York.
LEHMKUHL, D. M. 1973. A new species of Baetis (Ephemeroptera) from ponds in the
Canadian arctic, with biological notes. The Canadian Entomologist 105: 343-346.
McDUNNOUGH, J. 1936. A new arctic baetid (Ephemeroptera). The Canadian Entomologist
68: 33-35.
McIVER, S. B. 1975. Structure of cuticular mechanoreceptors of arthropods. Annual Review
of Entomology 20: 381-397.
MORIHARA, D. K., and W. P. McCAFFERTY. 1979. The Baetis larvae of North America
(Ephemeroptera: Baetidae). Transactions of the American Entomological Society 105:
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139-221.
MULLER-LIEBENAU, I. 1969. Revision der Europaischen Arten der Gattung Baetis Leach,
1815 (Insecta, Ephemeroptera). Gewasser und Abwasser, 48/49. 214 S.,
Max-Planck-Gesellschaft. Dokumentationsstelle, Gottingen.
MULLER-LIEBENAU, I. 1971. Ephemeroptera (Insecta) von den Kanarischen Inseln.
Gewasser and Abwasser, 50/5 1 : 7-40.
MULLER-LIEBENAU, I. 1974a. Baetidae aus Sudfrankreich, Spanien und Portugal
(Insecta, Ephemeroptera). Gewasser und Abwasser, 53/54: 7-42.
MULLER-LIEBENAU, I. 1974b. Rheobaetis : a new genus from Georgia (Ephemeroptera:
Baetidae). Annals of the Entomological Society of America 67: 555-567.
PERCIVAL, E. and H. WHITEHEAD. 1929. A quantitative study of the fauna of some types
of stream-bed. Journal of Ecology. 17: 282-314.
PHILIPSON, G. N. 1954. The effect of water flow and oxygen concentration on six species of
caddis fly (Trichoptera) larvae. Proceedings of the Zoological Society of London. A. 124:
547-564.
POPOVICI-BAZNOSANU, A. 1906. Contributions a l’etude de Forganisation des larves des
Ephemerines. Archives de zoologie experimental et generate 5: 66-78.
SCHMIDT, H. 1951. Amputation und Regeneration von Schwanzfaden und
Abdominlsegmenten bei Larven der Ephemeridenart Cloeon dipterum L. und ihr
Einfluss auf die Hautungsintervalle. Zool. (Phys) 62: 395-428.
SNEATH, P. H. A. and R. R. SOKAL. 1973. Numerical Taxonomy. W. H. Freeman and Co.,
San Francisco.
VENTER, G. E. 1961. A new ephemeropteran record form Africa. Hydrobiologia 18:
327-331.
WARD, J. V. and L. Berner. 1980. Abundance and altitudinal distribution of Ephemeroptera
in a Rocky Mountain stream, p. 169-177. In Advances in Ephemeroptera Biology. Eds.
Flannagan, J. F., and K. E. Marshall. Plenum Publ. New York.
THE DISTRIBUTION OF MOSQUITO (DIPTERA, CULICIDAE) LARVAE IN
SOUTHERN ALBERTA, 1976-1978
P.J. Scholefield'
G. Pritchard 2
M.A. Enfield3
Department of Biology
University of Calgary
Calgary, Alberta
T2N1N4
Quaestiones Entomologicae
17:147-168 1981
ABSTRACT
Twenty-seven species of Aedes, four species of Culiseta , two species of Culex and one
species of Anopheles were collected during five surveys of water bodies in Alberta south of
52° N during 1976, 1977 and 1978. Identifications were made on 4th stage larvae and
confirmed after rearing to adult. The species of Aedes showed four distribution patterns:
Group 1 was largely restricted to the mountains and foothills; Group 2 was found in all parts
of the study area; Group 3 occurred everywhere but in the prairie zone; Group 4 was largely
restricted to the parkland and prairie. Because of low numbers, the distribution of seven
species of Aedes could not be determined. Of the species in other genera, only Culiseta
inornata (a Group 2 species) was common enough to determine distribution, although Cs.
alaskaensis appears to belong to Group 1. Apart from these broad patterns, the habitat
preferences of individual species could not be determined. Of the 20 commonest species, 18
were associated with 13-20 other species at one time or another, and two had nine associated
species. The number of pools that formed by snow-melt or rain-fall, the dates of appearance of
larvae, and the relative species abundance all changed markedly from year to year.
RESUME
Vingt-sept especes d’ Aedes, quatre especes de Culiseta, deux especes de Culex et une espece d’ Anopheles ont ete
collectionnees au cours de cinq inventaires des plans d’eau situes au sud du 52ie me parallele Nord en Alberta, en 1976,
1977 et 1978. Les especes ont ete identifiees a partir des larves du quatrieme stade, et les identifications ont ete verifiees
•Present address: Pesticide Chemicals Branch, Alberta Environment, Calgary
2Author to whom correspondence should be addressed
3Present address: A.D.A.S. Sub Centre, Wye, Kent, U.K.
148
Scholefield, Pritchard and Enfield
grace a Velevage de ces memes larvaes jusqu’au stade adulte. La distribution des especes d’ Aedes formait quatre patrons:
le groupe I etait principalement restreint aux montagnes et aux piedmonts; le groupe 2 se retrouvait dans toutes les
regions inventoriees; le groupe 3 se retrouvait partout, sauf dans la prairie; le groupe 4 etait en grande partie restreint au
“parkland” et a la prairie. La distribution de sept especes d’ Aedes n pu etre etablie a cause des faibles captures. Parmi
les especes appartenant aux genres autres qu’ Aedes, seule Culiseta inornata etait assez commune pour pouvoir etablir sa
repartition (el le se classe dans le groupe 2); Cs. alaskaensis semble appartenir au groupe I. Mis a part ces patrons de
distribution generaux, Vhabitat prefere de chaque espece n’a pu etre determine. Parmi les vingt especes les plus
communes, dix-huit etaient associees avec de treize a vingt autres especes a une occasion ou I’autre, et deux especes se
retrouvaient chacune en association avec neuf especes. Le nombre d’etangs formes par la fonte des neige ou la pluie, les
dates d’emergence des larvaes, et I’abondance relative des especes ont tous varies d’une annee a I’autre.
TABLE OF CONTENTS
Introduction 148
Methods . 149
Results 154
Group 1 - Mountains and Foothills Species 154
Group 2 - Ubiquitous Species . 156
Group 3 - Non-prairie Species . 157
Group 4 - Parkland and Prairie Species . 157
Species of Uncertain Distribution . 159
Discussion . 161
Acknowledgements . 166
References 166
INTRODUCTION
Proper identification of species is basic to any biological study. Furthermore, studies of
mosquito-carried diseases and mosquito abatement in Alberta cannot make efficient use of data
already accumulated and analysed unless the species, both in the current programs and in work
already done, are properly known. The problem of adult migration into an area in which a
larviciding program has been carried out, can hardly begin to be solved without thorough
knowledge of the species involved.
In the report of the “Symposium on Biting Fly Control and Environmental Quality” held in
Edmonton in 1972, the summary of the section on Behaviour and Ecology of Populations
(Mclver, 1973) contained the following passage:
“ When considering a control program, four superficially simple questions need to be asked about the
pest species: 1 . What is it? 2. Where is it? 2. What is it doing? 4. How does it accomplish its observed
behaviour?”
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
149
This report attempts to answer the first two questions for mosquitoes in Southern Alberta.
Such a study was necessary because, although several reports about mosquitoes of Alberta
have included species lists (e.g. Strickland, 1938; Happold, 1965; Pucat, 1965; Wada,
1965; Tawfik and Gooding, 1970), these studies represented a limited approach to species
identification and all contained some misidentifications through use of incomplete or
inaccurate keys. For example, Enfield (1977) reported that two species, Aedes euedes
Howard, Dyar and Knab 1917 and Aedes mercurator Dyar 1920 have long remained
unrecognized. Larvae of Aedes euedes key to Aedes riparius Dyar and Knab in Rempel
(1950) and Carpenter and LaCasse (1955), while adult males have genitalia that resemble
those of Aedes excrucians (Walker) . Aedes barri Rueger 1958 and Aedes beklemishevi
Denisova 1955 are synonyms for Aedes euedes (Wood, 1977). Aedes mercurator larvae
key to one of four other species in conventional keys. Until this species was recognized in
Calgary in 1975 (Enfield, 1977), it was recorded only from Dawson, Yukon Territory
(Dyar, 1920), and from the U.S.S.R. (Danilov, 1974). Many adults labelled Aedes
stimulans (Walker) in the Canadian National Insect Collection are in fact Aedes
mercurator , and its range is now known to extend across much of Northern Canada
(Wood, 1977).
These problems stimulated a survey of the mosquitoes in Southern Alberta. We felt
that the area of Alberta approximately south of latitude 52°N, within which Calgary is
conveniently central, was the maximum that could be covered in a study of this kind.
Further, we thought that only by collecting extensively in all ecological zones and by both
identifying larvae and rearing them to obtain associated adults, could familiarity with the
whole group be gained and problems such as those involving Aedes mercurator and Aedes
euedes be exposed.
The data, which are summarized in this paper, may be found in a more extensive form,
together with a key to 4th stage Aedes larvae occurring in southern Alberta in an
unpublished report (Scholefield, Enfield and Pritchard, 1979).
METHODS
The study area was surveyed five times. Samples of larvae were collected along planned
routes so that the major ecological zones and different types of water bodies were included in
each trip. Distribution of sites that yielded larvae is shown in Figures 1 & 2. In the Spring
(April-May) of 1976, mosquito larvae were collected from 115 sites, from 45 sites in the Spring
of 1977, from 61 sites in the Summer (June- July) of 1977, from 124 sites in the Spring of 1978,
and from 41 sites in the Summer of 1978.
Most collections were obtained by driving along set routes and sampling water bodies which
were seen from the road. However, spring sampling in the Southeast of the province in 1977
and 1978 was done during a helicopter tour by Mr. J. McIntosh (Alberta Environment) to the
various abatement areas under his jurisdiction; one slough in or near each town was selected
from the air, then sampled. In 1977 and 1978, a record sheet containing information on each
site was filled out and a photograph of the site was attached.
Larvae were captured by means of a standard pint dipper, and, wherever possible, dipping
was continued at each site until about 50 larvae had been caught. The larvae were concentrated
Quaest. Ent., 1981, 17
150
Scholefield, Pritchard and Enfield
Fig. 1 . Sites which yielded mosquito larvae during spring (April-May) sampling. The 1 976 survey did not adequately cover
the mountain zone and northern foothills and this was corrected in later years. In 1978 the whole region was surveyed but
only those areas not visited in previous years are indicated. Also shown are the 4 ecological zones (heavy lines), the
Columbia Ice-fields (Cl) and the Cypress Hills (CH), the major rivers (thin lines), and some towns (B = Banff, BR =
Brooks, C = Calgary, CY = Cayley, CO = Coleman, D = Drumheller, F = Fort McLeod, J = Jasper, L =
Lethbridge, M = Medicine Hat, N = Nordegg, R = Rocky Mountain House
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
1
:|
Fig. 2. Sites which yielded mosquito larvae during summer (June-July sampling
Quaest. Ent., 1981, 17
152
Scholefield, Pritchard and Enfield
in a clear plastic cup by means of a cylindrical filter which was fitted into the top of the cup. As
water containing larvae was poured in, the larvae were retained in the cup while excess water
spilled out. Usually no more than about 25 larvae were kept in each cup, which was sealed with
a plastic lid, and returned to the laboratory in a cooler. When samples were being transported,
a stop of five minutes duration was made every 30 minutes to allow larvae to rise to the water
surface and breathe. When 4th stage larvae were collected about one-third of the sample was
killed in KAiAD 1 and preserved in 95% ethanol. These subsamples were potential
replacements in case a whole sample should die during transportation, since mosquito larvae
decay very rapidly after death, making identification impossible. Some pupae collected in the
field represented species of which there were no larvae in the same sample. Thus collection of
pupae for subsequent rearing to the adult stage was important, even though pupae are more
difficult to keep alive during transportation.
In the laboratory, larvae were kept in these same plastic cups at 20°C, in tap water which
had been conditioned with dead slough grass for one week. The water was aerated to prevent
formation of surface scum, and larvae fed on the micro-organisms on the surface of the
decaying grass. Larvae were identified when in stage IV and some were returned to the cups to
complete development; the remainder were killed in KAiAD and preserved in 95% ethanol.
Because examination of preserved specimens in Edmonton and Calgary showed that many
Aedes adults and larvae had earlier been misidentified, as many identified larvae as possible
were reared to adults and then reidentified to reduce the chance of further misidentification.
Pupae were placed individually or in groups of about 20 in cups, and the emerging adults were
trapped in inverted plastic vials on top of a funnel made of plastic fly screen. Several hours after
emergence, vials containing adults were placed in a freezer to kill the mosquitoes and within a
few days the specimens were mounted on a pin with shellac and returned to the freezer for a
few weeks to complete drying.
A draft of a key now published in Wood et al., (1979) was the main basis for identification
of larvae. Keys of Gjullin and Eddy (1972) and Carpenter and LaCasse (1955) (modified to
include Aedes euedes and Ae. mercurator) were used for identification of adults and for
identification of some larvae. Voucher specimens of stage IV larvae and adults of all species are
stored in the Department of Biology, University of Calgary.
1 KAAD= 1 part kerosene, 10 parts 95% ethanol, 2 parts glacial acetic acid and 1 part
dioxane. KAi AD = equal parts KAAD and 95% ethanol.
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
153
Fig. 3. Examples of the 4 types of distributions Group 1 - Mountains and Foothills: Ae. communis : Group 2 - Ubiquitous
Ae. euedes : Group 3 - Non-Prairie: Ae. mercurator ; Group 4 - Prairie and Parkland: Ae. flavescens.
Quaest. Ent., 1981, 17
154
Scholefield, Pritchard and Enfield
The Atlas of Alberta (Government of Alberta, 1969) was used to define ecological zones in
preference to the one used by Strickland (1938), because there appeared to be little evidence
that the prairie/parkland zone was divisible into as many different areas for Aedes species as
Strickland showed. Four ecological zones (Mountains, Foothills, Parkland, Prairie) were
included within the study area (Figs. 1, 2 & 3).
In 1977 and 1978, in addition to recording location, we attempted to categorize habitat type
for each species. Four habitat characteristics were noted at each site: 1). whether the pond was
open or shaded by trees; 2). whether the surface was shaded by slough grass or clear of
vegetation; 3). whether or not there were willows, conifers or poplars around the margin; and
4). whether the water body was stagnant or was continually fed by seepage or continuous
surface run-off. In 1978 water analysis was also made at each site containing larvae. The pH,
conductivity, temperature, and dissolved oxygen content at the time of sampling were recorded
with a view to further characterization of habitats of individual Aedes species.
RESULTS
Thirty-four species of mosquitoes in four genera were collected in this survey. Twenty-seven
of these species belong to Aedes , most larvae of which inhabit temporary bodies of water
formed by melting snow or rainfall. Although not important vectors of disease in Alberta, some
species develop huge populations with a high nuisance rating.
The species of Aedes show four distribution patterns (Fig. 3). Group 1 mosquitoes are
largely restricted to the mountains and foothills; Group 2 mosquitoes occur in all 4 zones;
Group 3 mosquitoes occur in 3 of the zones, but are missing from the prairie; and Group 4
mosquitoes are largely restricted to the parkland and prairie. Some overlap at the
foothills-parkland border is not uncommon in species in Groups 1 and 4. Because of rarity, the
distribution of seven species could not be ascertained.
Group 1 - Mountain and Foothills species
Aedes (Ochlerotatus) canadensis (Theobald) (15 collections)
This species is included in the mountains-foothills group although specimens were taken at
two sites west of Calgary at the western edge of the parkland zone. Most larvae were found in
spring, but were also collected in summer in the mountains in 1977. Most were found in
unshaded locations in forested areas, and in non-stagnant pools. The species was found on only
three occasions by itself; at other times it was associated with one or more of nine species of
Aedes.
Aedes ( Ochlerotatus ) communis (de Geer) (36 collections)
Classification of this species is discussed by Ellis and Brust (1973). It was one of the
commonest species in the mountains and foothills, where larvae were found only in spring in
snow-melt pools. Larvae were in forested areas, but in relatively open pools. The water in most
pools was clear and cool and laboratory rearing was much more successful when the water was
kept clear and well aerated. The species was found on 10 occasions by itself; at other times it
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
155
was associated with one or more of 20 other species.
Aedes (Ochlerotatus) hexodontus Dyar (34 collections)
Larvae of this species are also common inhabitants of open pools in the mountains, often
found with larvae of Ae. communis and Ae. punctor . Predominantly a spring species, larvae
were found at one location in the mountains in the summer of 1977. According to Carpenter
and LaCasse (1955), some hatching may be delayed until long after the pool has been filled by
snow-melt. It was rarely found by itself (four collections), occurring at one time or another with
19 other species.
Aedes ( Ochlerotatus ) pionips Dyar (11 collec
Not a common species, found predominantly in spring in open locations in forested areas of
the foothills, although larvae were collected during the summer on one occasion in the
mountains in 1977. It was found once by itself, on other occasions being associated with one or
more of 17 other species. Larvae were difficult to rear, even in clear, well aerated water.
Aedes ( Ochlerotatus ) pullatus (Coquillett) (47 collections)
One of the most common mountain species, larvae were in open pools in both spring and
summer. In 1977 it was found alone in a pool near the Columbia Icefields on 10th May (water
temperature 14°C) and again on 5th July (water temperature 20°C). On both occasions 4th
stage larvae and pupae were present, and the indication is therefore that eggs hatched on two
occasions in 1977. The species was not present in this pool in the spring of 1978; Ae. communis
was there instead. Aedes pullatus was found on 10 occasions by itself, and on other occasions
with 20 other species.
Aedes ( Ochlerotatus ) punctor (Kirby) (33 collections)
Another common mountain - foothills species, collected mainly in spring but also taken in a
few summer samples in 1977 and 1978. Larvae apparently prefer grass-covered, willow-fringed,
open pools. Found on three occasions by itself, it occurred at other times with 23 other species
of Aedes.
Aedes (Ochlerotatus) schizopinax Dyar ) (1 1 collections)
Like Aedes canadensis this species has been taken from the western edge of the parkland,
but is otherwise restricted to the mountains and foothills. It is rather local, being found at only
one location in 1978, but it did occur in fairly high numbers in certain locations in the southern
part of the study area during the summer of 1977, when its larvae were among the most
common in the Coleman area. Almost all samples were in open, grass-covered pools. Nielsen
(1961) reports that larvae prefer small to moderately sized pools containing considerable
vegetation and organic matter, often covered with scum. He also reports that adult females are
not known to take blood meals. Aedes schizopinax was found only once by itself; at other times
it occurred with one or more of 16 other species. During spring, larvae appeared later than
those of the other species, but developed quickly both in the field and laboratory.
Quaest. Ent., 1981, 17
156
Scholefield, Pritchard and Enfield
Group 2 - Ubiquitous Species
Aedes (Ochlerotatus) cataphylla Dyar (90 collections)
A common species in all zones, apparently having a wide tolerance of habitat conditions. It
is a spring species, and was the most frequently encountered species in the spring of 1976. Eggs
apparently hatch on the first flooding, whether warm or cold, and this could account for the two
records of larvae in the summer sampling in 1977. Larvae developed quickly and Carpenter and
LaCasse (1955) report that it is one of the first Aedes species to appear on the wing. Fourth
larval and pupal stages suffered heavy mortality when reared in the laboratory. Aedes
cataphylla larvae were found alone on 14 occasions, and with 22 other species at other times,
especially Ae. campestris, Ae.fitchii, and Ae. mercurator .
Aedes ( Ochlerotatus ) euedes Howard, Dyar and Knab (45 collections)
This species was fairly common in the 1976 survey, but was not recorded as frequently in
1977 or 1978. It is primarily a snow-melt pool species, but small numbers of larvae have been
found in rain-water pools during the summer in the mountains. Larvae were found in many
types of pools in all four ecological zones. It occurred only six times alone, at other times being
associated with 17 other Aedes species. This was the most difficult of the commoner species to
rear in the laboratory, partly because the larvae were very aggressive and apparently unable to
withstand even slight crowding. Rate of development was about average, but overall
development time was long because individuals grew to a larger size than most other species. In
the spring of 1975, this species required six to seven weeks to complete development from egg to
adult (Enfield, unpubl. obs.). Enfield (1977) and Wood (1977) discuss nomenclatural problems
surrounding this species.
Setae vary between western and eastern Canadian populations. In Alberta abdominal seta
7-II is multiple (more than four hairs), whereas eastern specimens have seta 7-II with three or
fewer hairs (Wood, pers. comm.).
Aedes ( Ochlerotatus ) fitchii (Felt and Young) (65 collections)
This common species in 1976 was less so in 1977 and 1978, but was widely distributed in
open habitats. As might be expected of a widely distributed species, it was rarely found alone
(three collections), but was found at one time or another with 22 other species, frequently with
Ae. cataphylla. It was found primarily in the spring, but also in the summer on two occasions in
1977. The larvae were among the slowest developers, many being in the second stage when
other species in the same water body were in the fourth. Larvae were easy to rear.
Aedes ( Ochlerotatus ) implicatus Vockeroth (44 collections)
Another early species which was, however, found in a few locations in the summer of 1977.
It has a wide tolerance of habitat conditions and is clearly not restricted to montane, forested
regions as stated by Carpenter and LaCasse (1955). Found on seven occasions alone, it
occurred at other times with 23 other species. The larvae were easy to rear in the laboratory.
Aedes ( Ochlerotatus ) increpitus Dyar
(17 collections)
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
157
Another species which was much commoner in 1976 than in the later years. Larvae prefer
open ponds in the foothills in the southern half of the region, and were found in enormous
numbers at Cayley in 1976. Although rare in mountains and prairie, this species is included in
the ubiquitous category rather than creating a separate category for it. All records were from
spring sampling except for a sole mountain record which was taken in the summer of 1977. It
was rarely (two collections) found in the absence of other species. The larvae, like those of Ae.
mercurator to which they are rather similar, are distinctly yellowish in life, unlike those of most
black-legged species.
Aedes ( Aedimorphus ) vexans (Meigen) (40 collections)
Aedes vexans is normally one of the commonest species in this area. However in the dry
years of this survey it was found on only two occasions in 1976 and 2 occasions in 1978.
Normally thought of as a typical summer species, all of the 1976 and 1978 records were from
spring sampling, and all of the records from the northern part of the region in 1977 were also
taken in the spring. Aedes vexans normally requires water temperatures above 8°C and a
period of prior conditioning for successful hatching (Horsfall, 1956). However in 1977 eggs
were hatching at much lower water temperatures. Our records show that larvae are found
usually in open, grass-covered ponds throughout the region. The larvae were found associated
with 21 other species, and on only seven occasions were they the only Aedes larvae taken at a
site. The larvae are quite easy to rear in the laboratory.
Group 3 - Non-prairie Species
Aedes (Ochlerotatus) excrucians (Walker) (40 collections)
This species was widely distributed through the mountains, foothills and parkland region.
Larvae were primarily in open, grass-covered ponds in forested areas at higher elevations.
Individuals develop slowly, like those of Ae. fitchii, to which they are very similar in many
respects. On only five occasions was Ae. excrucians the only species taken at a site; on other
occasions it was associated with one or more of 20 other species. It was found only during
spring, except for the single 1977 record.
Aedes ( Ochlerotatus ) mercurator Dyar (40 collections)
Except for two mountain samples in 1978 this species was taken during spring. Open,
snow-melt pools in non-forested areas are preferred. Like Ae. euedes, Ae. mercurator larvae
survive well in very cold water (Enfield and Pritchard, 1977). The confusion with Ae. stimulans
in Alberta is discussed by Enfield (1977). Many larvae were associated with those of Ae.
cataphylla.
Group 4 - Parkland and Prairie Species
Aedes ( Ochlerotatus ) campestris Dyar and Knab
(76 collections)
Quaest. Ent., 1981, 17
158
Scholefield, Pritchard and Enfield
One of the commonest species in the study area, Ae. campestris was found in all 4 zones but
was characteristic of open country and much commoner in the parkland and prairie than in the
mountains and foothills. Most larvae were in open pools, without marginal vegetation. Rempel
(1950) reports larvae from pools rich in organic matter with a pH on the alkaline side, but since
all of the locations from which larvae were taken in this study were alkaline the latter criterion
is not particularly helpful. Larvae were found during both spring and summer sampling. The
species was found alone on six occasions, but at other times with 17 other species, frequently
including Ae. dorsalis and Ae. cataphylla . The larvae were notably paler than those of other
species and were very easy to rear in the laboratory, being able to withstand considerable
crowding and fouling of the water. Development was rapid, except under crowded conditions
when the rate slowed, although larvae survived well for long periods.
Aedes (Ochlerotatus) dorsalis (Meigen) (66 collections)
This species was one of the commonest in the survey and was found in all four zones,
although it was taken at only one location in the mountains and at only a few in the foothills.
Most larvae were in open pools without marginal vegetation. It is normally thought of as a
summer species and was indeed uncommon in the spring of 1976 and was commoner during
summer of 1977 than in the spring of that year. However, in 1978 it was much commoner
during spring than in the summer. This was the only Aedes species present at 10 sampling sites,
and was found with one or more of 16 species on other occasions. Associated species were Ae.
vexans, Ae. cataphylla, and Ae. spencerii.
Aedes ( Ochlerotatus ) flavescens (Muller) (73 collections)
This was one of the most common species encountered in the survey. It was widely
distributed over the parkland and prairie regions and was also taken in the foothills zone,
although rarely. Like so many other spring species, larvae were taken during summer of 1977,
presumably reflecting lack of snow-melt pools in that year. Larvae were generally at an earlier
stage than those of other species in the same water body and this slow rate of development
could be a disadvantage in years when pools dry up quickly. Because of its geographical
distribution most larvae were in open habitats without marginal vegetation. Only twice found in
the absence of other species, Ae. flavescens often occurred with Ae. campestris. According to
Carpenter and LaCasse (1955), adult females may survive until late in the season, and in July
1977 females were biting during the day in the southern part of the region.
Aedes ( Ochlerotatus ) melanimon Dyar (13 collections)
This species was only found in the southern part of the parkland and prairie zones during the
summer of 1977. Most larvae were in open, grass-covered ponds, without marginal vegetation
and frequently fed by water from more permanent sources. First recorded in Canada from
Brooks, Alberta by Burgess (1957), this species was taken in only 13 sites in this survey. It
occurred with 9 other species, including Ae. vexans on 12 occasions and Ae. dorsalis on 11
occasions.
Aedes ( Ochlerotatus ) spencerii (Theobald) (71 collections)
Like Aedes flavescens, this species occurs occasionally in the foothills zone, but it is so
widespread over the parkland and prairie that it is best considered in this latter group. It was
one of the commoner species in this survey, even though it was only taken three times in 1977.
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
159
In that year, it presumably suffered, as did many other spring species, from the lack of suitable
snow-melt pools in the parkland and prairie zones. Aedes spencerii was found alone more than
any other species (27 times), and had a relatively low number of associated species (13) for
such a common mosquito. In addition, there were usually few larvae of any species in the pools
in which it was found.
Species of Uncertain Distribution
Aedes (Aedes) cinereus Meigen (11 collections)
The few records for this species are from the mountains, foothills and western edge of the
parkland, where larvae were found in pools fringed by poplars and willows. During this survey,
it was found only during spring, but during a population study in 1974 eggs hatched both in the
spring and later in the year along >yith Ae. vexans (Enfield and Pritchard, 1977).
Aedes (Ochlerotatus) diantaeus Howard, Dyar and Knab (4 collections)
This species is, according to Rempel (1950), rare and local and found only in forested
regions of Canada. Most larvae were collected from open, grass-covered pools with marginal
evergreens and willow, in water that was not completely stagnant. Carpenter and LaCasse
(1955) report the larvae from pools of melting snow, and both collections in 1978 were made
during spring. In 1977, however, larvae were taken in rainwater pools during the summer.
These are the first records for southern Alberta outside of the National Parks (recently
reported from Banff National Park; Wood, pers. comm.). Aedes diantaeus was reported by
Happold (1965) from Flatbush, 150 km north of Edmonton.
Aedes ( Ochlerotatus ) impiger (Walker)
(4 collections)
Larvae were found only three times: twice in the mountains in 1977 and once in the foothills
in 1978. Open, grass-covered, non-stagnant pools with marginal evergreens were the typical
habitat. A single adult female was taken in the parkland south of Lethbridge.
Aedes (Ochlerotatus) intrudens Dyar (3 collections)
A rare species whose larvae are found only in shaded, snow-melt pools with marginal willows
and poplars in the foothills and mountains.
Aedes (Ochlerotatus) nigromaculis (Ludlow) (1 collection)
The rarest species of Aedes encountered in the survey, Ae. nigromaculis was found only
once in an open, grass-covered pool formed from irrigation seepage into a grassy ditch,
north-west of Lethbridge during the summer collecting period in 1977. According to Rempel
(1950) it is a species of the open prairies and has been recorded previously from Lethbridge and
Medicine Hat. Perhaps the record for Beaverlodge (55° 13’N, 119° 26’W) (Strickland, 1938)
is in error.
Aedes ( Ochlerotatus ) sticticus (Meigen) (6 collections)
Quaest. Em.. 1981, 17
160
Scholefield, Pritchard and Enfield
Enfield (1977), having seen larvae of both Ae. implicatus and Ae. mercurator misidentified
as Ae. sticticus , suggested that the record by Pucat (1965) needed to be verified. Five samples
taken in 1977 and one in 1978, in late May or June, certified its presence, although it is
apparently not common. Larvae were taken from open or shaded grass-covered pools, generally
with marginal willows. The species was not collected from the mountain zone.
Aedes (Ochlerotatus) trichurus (Dyar) (3 collections)
This species was not common, with larvae being found only in a few willow-fringed pools in
the foothills and mountains. The larvae developed quickly, but were not easy to rear. They are
large, and like those of species which have banded-legged adults, have a distinctly yellowish
appearance in life.
Genera other than Aedes
The following species in the genera Anopheles , Culiseta and Culex overwinter as adults. In
the spring they are the first mosquitoes to be seen on the wing, especially the genus Culiseta ,
whose adults are quite large. Adults appear as early as April if the weather is warm, but the
larvae tend to appear later in the season (late springearly summer) after the appearance of the
larvae of aedine species. The females lay their eggs on the water surface and there is no
conditioning of the eggs, which hatch within a few days.
Anopheles ( Anopheles ) ear lei Vargas (2 collections)
There are no previous records of this species occurring in southern Alberta. Specimens were
taken from a site near Twin Butte in 1977 and another site in Waterton National Park on the
Red Rock Canyon Road in 1978. The abundance of this species is usually quite low and local
and no serious outbreaks have been recorded.
Culex (Culex) tarsalis Coquillett (10 collections)
This species is represented across the entire southern half of the province, and is especially
prevalent in the irrigated areas. Western equine encephalitis is associated with this species
(Shemanchuk 1969), and if weather and irrigation conditions are correct, it could prove to be a
problem. In the southern part of the study area it was often found in shallow pools in pasture
fields flooded by irrigation water as well as roadside ditches.
Culex (Neoculex) territans Walker (4 collections)
This is the first record of Culex territans south of Edmonton. We found it locally only in the
foothills. Most larvae were associated with bodies of water that contained a lot of vegetation.
Culiseta ( Culiseta ) alaskaensis (Ludlow) (17 collections)
Strickland (1938) reports this species from Banff National Park, the Nordegg area, and the
Calgary/Ft. Macleod area. It would seem to prefer higher elevations, being found in our survey
only in the mountains and foothills except for two records west of Calgary in the parkland. This
species was on the wing in early May in Jasper National Park in 1978, and was a persistent
biter.
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
161
Culiseta ( Culiseta ) incidens (Thomson) (2 collections)
This species was very rare in the study area having been found only once at Rocky Mountain
House in 1977 and once northeast of Banff in 1978.
Culiseta ( Culiseta ) inornata (Williston) (47 collections)
This is the most widespread Culiseta in southern Alberta and one of the most troublesome
species in the Lethbridge area during the summer, when, regardless of weather conditions,
irrigation produces ideal breeding grounds. Culiseta inornata is a troublesome biter of both
man and livestock as well as wild ungulates, and has been associated with western equine
encephalitis (Shemanchuk 1969).
Culiseta (Culicella) morsitans (Theobald) (1 collection)
Only one specimen of this species was taken during the study period, being taken south of
the Jasper townsite during the spring of 1977. According to Rempel (1950) it is very rare in
Western Canada, but has been previously reported from Alberta (Dyar, 1928; Wada, unpubl.).
DISCUSSION
During the three years of this study a number of changes have been made to published lists
(e.g. Pucat, 1965) of mosquitoes in southern Alberta. Enfield (1977) revealed the presence of
one species not previously recorded from Canada namely Aedes schizopinax . This species in
fact, has a wide distribution in the southern part of the mountains and foothills. It had
previously been found in Calgary by Robins (1972, unpubl.) and Scholefield (1973, unpubl.),
but not officially recorded.
Aedes euedes has a very wide distribution across the entire survey area (Fig. 3). Previously
identified as Ae. barri or Ae. riparius, its naming as Ae. euedes has now been properly
established (Wood, 1977). Aedes stimulans, which, according to records, had a wide
distribution in Alberta, apparently does not occur in the province. All specimens identified as
such, appear to be Aedes mercurator (Enfield, 1977; Wood, 1977), which is widely distributed
in the mountains, foothills, and parkland of the study area (Fig. 3).
Aedes increpitus is now recorded from southern Alberta, being first found in the Calgary
area by Scholefield (1973, unpubl.). It was not found north of Calgary, however. Enfield
(1977) indicated that the presence of this species in northern Alberta is suspect due to the fact
that all the records he had seen from that area were actually Ae. mercurator.
Another new record for southern Alberta is Aedes implicatus , being first recorded by
Enfield (1977). It ranges across the entire southern half of Alberta, although it is local in the
prairie.
Northern records for Aedes hexodontus have been published by Pucat (1965), Graham
(1969), and Tawfik and Gooding (1970). It too was first recorded in southern Alberta during
the first year of this present study (Enfield, 1977). Its distribution is restricted to the mountains
and foothills where it is quite common.
The ranges of six other species ( Ae. canadensis, Ae. pionips, Ae. punctor, Ae. communis,
Ae. trichurus and Ae. pullatus) have been somewhat extended. Previous reports in the
literature listed Banff as the only site in southern Alberta (Strickland, 1938; Rempel, 1950).
All are more or less restricted to the mountains and foothills, areas which have not been
Quaest. Ent., 1981, 17
162
Scholefield, Pritchard and Enfield
previously extensively collected.
impiger
diantaeus
ci nereus
i ntrudens
melanimon
f lavescens
spenceri i
campestri s
dorsalis
mercurator
excrucians
i ncrepi tus
vexans
impl icatus
fi tchi i
euedes
cataphylla
canadensi s
pionips
schizopinax
punctor
pullatus
hexodontus
communi s
"i — r
8
pH
10
fig. 4. The range of pH of the sites from which larvae of 24 species of Aedes were collected.
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
163
1 1 1 1 r 1 1 r — i 1
0 50 100
MICROMHOS X10
Fig. 5. The specific conductance (mean ± 95% confidence limits) of the sites from which larvae of 15 species of
Aedes were collected.
Quaest. Ent., 1981, 17
164
Scholefield, Pritchard and Enfield
Factors affecting mosquito distribution in southern Alberta are largely unknown. Most
species encountered in this study have wide distributions, although 15 species were collected
only from the mountain - foothills region and six were more or less restricted to the parkland -
prairie. On a finer scale, habitat distinctions between species are impossible to make on the
basis of information collected in this survey. Of the 20 species which were collected on at least
10 occasions, all but two were associated with 13-20 other species at one time or another, and
the other two species had nine associated species. Clearly, with such overlap there is no such
thing as a unique habitat for each species. This problem was further exposed when
characteristics of the individual sites were considered. There were few obvious departures from
the typical set of habitat conditions for the area, nor were there any obvious differences in pH
(Fig. 4) or specific conductance (Fig. 5) of the water bodies inhabited by the different species.
Water in the mountains and foothills has lower conductivity than water in the plains, and
species restricted to these areas are therefore found in water with lower conductance values.
This does not, of course, imply that conductance of the water is the reason for restriction of
these species to those zones. Similarly, water bodies in the mountains and foothills are more
likely to be shaded than are those in the parkland and prairies.
Maire and co-workers (e.g. Maire and Aubin, 1976; Maire et al., 1978; Mailhot and Maire,
1978) have related the distribution of mosquito larvae to vegetation types in Quebec. However,
the analysis is rarely clear-cut, even with only one half of the number of Aedes species that are
present in southern Alberta. For example, Maire (1977) proposes four associations based on
abundance of four species of Aedes and two of Culiseta. However, except for Ae. decticus,
which is found only in open sites, all species occur in all habitats, and three species are
abundant in more than one habitat type.
A further difficulty in judging habitats as suitable or not for particular species is that
relative species abundance can change quite markedly from year to year. There was a marked
decline in number of sites at which the 18 most common species were taken from 1976 (355
sites) to 1977 (226 sites), even though there was no summer sample nor were the mountains
well sampled in 1976. There was only slight recovery in 1978 (273 sites). Particular species
worth noting are Ae. campestris, Ae. cataphylla, Ae. euedes , Ae. fitchii , Ae. flavescens, and
Ae. mercurator which were all common in 1976, but much reduced in numbers in later years.
Aedes euedes and Aedes mercurator are especially noticeable since together they were so
abundant at a pond just northwest of Calgary in 1975 that they provided excellent material for
a population study (Enfield and Pritchard, 1977). Also noticeable are Ae. vexans , normally
thought of as the commonest species in this area, but which only achieved any level of relative
abundance in 1977, and Ae. spencerii , which was uncommon in 1977, but the commonest
species in 1978. Records were not kept of individual pools from year to year, except for a pool
near the Columbia Ice Fields which had two hatches of Ae. pullatus in 1977, but none in 1978.
In the latter year, Ae. communis was present instead.
Overall changes in abundance of mosquitoes are clearly related to number of pools formed
by snow-melt or rainfall. Availability of snow-melt pools depends upon amount of snow-fall,
pattern of Chinook winds during winter, and speed at which snow disappears during spring.
During this survey, there appeared to be fewer snow-melt pools in 1977, especially in the
parkland and prairie, than in other years. Rain-filled pools depend on the pattern of
precipitation during spring and summer, and on the level of the water table. There were fewer
such water bodies in 1978 than in 1977. Overall, the three years of this study coincided with a
period of relative drought which was inimical to development of mosquito populations. Our
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
165
experience with Aedes vexans exemplifies this trend. In 1972 (Robins, unpubl.) and 1973
(Scholefield, unpubl.), Ae. vexans was abundant in the Calgary area. In August of 1974 at an
experimental site outside Calgary, larvae of Ae. vexans were abundant, but the pond dried up
before larvae had completed development (Enfield and Pritchard, 1977). In 1975, the pond
flooded at snow-melt, was dry by early June and remained so for the rest of the year and no Ae.
vexans eggs hatched. At our experimental sites in N.W. Calgary it was necessary to artificially
flood pond beds in 1976 and 1977 in order to have populations with which to work. In 1978
these ponds flooded naturally in April, but dried up permanently for the year at the beginning
of July. No Ae. vexans larvae appeared. These trends applied generally over the whole study
area.
Related to the pattern of precipitation is the distinction between “spring” and “summer”
species, a concept which is by no means as tidy as many hold it to be. Spring species are those
whose eggs generally hatch on the first flooding of the year, this normally being the result of
snow-melt, and an egg diapause restricts the number of generations per year to one. Exceptions
to this latter statement may be Ae. pullatus and Ae. cinereus (see above), and perhaps other
species (see Magnarelli, 1977). Montane and foothills species generally belong to this group, as
might be expected since there will be an abundance of snow-melt pools in these regions.
So-called “spring species” apparently hatch in the first flooding whether warm or cold and this
accounts for their appearance in summer samples, especially in 1977 when there was a dearth
of snow-melt pools. However temperature requirements for hatching in these species appear not
to have been well studied.
Hatching of eggs of summer species such as Ae. vexans and Ae. dorsalis is complicated
(Horsfall, 1956). The basic pattern is that eggs that have completed embryogeny hatch only
after a period of drying, flooding with water above 8°C and a lowering of oxygen concentration.
Hovyever, there is variability between populations in response to these factors and even given
these conditions many eggs do not hatch. This erratic hatching has been viewed by some
authors as an inherent safeguard against unpredictable water conditions, but Gillett et al.,
(1977) have suggested that in Ae. aegypti this erratic hatching is due to the grazing of
oxygen-consuming bacteria from egg surfaces by the first larvae that hatch. Egg diapause in
Ae. vexans is also complicated (Taylor, pers. comm.).
Again, our experience with Ae. vexans is illustrative of the complexity of the problem. In
our population studies in 1974 (Enfield and Pritchard, 1977), eggs of Ae. vexans behaved
according to the above formula; i.e. no eggs hatched when flooded with water at 6°C and four
subsequent floodings at 20°C were required in order to hatch all the eggs. Few, if any eggs,
were laid in our experimental field sites in 1974 or 1975 and so there were few eggs available in
the ground in 1976 or 1977. When they were hatched artificially by flooding with water above
the threshold for hatching in the laboratory in these years, the hatch was complete with one
flooding, perhaps because of the low density and supporting the hypothesis of Gillett et al.,
(1977). However, more interestingly, in the survey, Ae. vexans larvae were generally taken in
spring sampling rather than summer sampling and the evidence suggests that eggs were
hatching in much colder water than the accepted threshold. Eggs of Ae. vexans collected from
soil cores in 1977 were in fact hatched in the laboratory at temperatures below 8°C. The same
remarks apply to Ae. dorsalis in 1978.
In view of the confused literature on this subject (e.g. Horsfall et al., 1973) and in light of
the above observations, a renewed study of the hatching requirements of Aedes eggs would
seem to be in order. Such a study might go a long way towards explaining the distribution and
Quaest. Ent., 1981, 17
166
Scholefield, Pritchard and Enfield
abundance of different species from year to year.
ACKNOWLEDGEMENTS
It is with much gratitude that we acknowledge the encouragement, advice and assistance
given to us by Dr. D.M. Wood of the Biosystematics Research Institute, Agriculture Canada,
Ottawa. Dr. R.H. Gooding, Mr. Will Johnson, and members of the Pesticide Chemicals
Branch, Pollution Control of Alberta Environment who assisted in the collection of larvae at
various times. We are especially grateful to Mr. Jock McIntosh of Alberta Environment in this
last regard. We also thank Dr. R.A. Ellis of the City of Winnipeg for his comments about a
draft of this report. Funds for this project were generously provided by the Pesticide Chemicals
Branch, Pollution Control Division Alberta Environment.
REFERENCES
Burgess, L. 1957. Note on Aedes melanimon Dyar, a mosquito new to Canada. Canadian
Entomologist. 89:352.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of North America (North of Mexico).
University of California Press.
Danilov, V.N. 1974. Restoration of the name Aedes (O.) mercurator Dyar to the mosquito
known in the U.S.S.R. as Aedes riparius ater Gutsevich (Diptera: Culicidae) (in
Russian). Parazitologiya 8(4):322-328.
Dyar, H.G. 1920. The mosquitoes of British Columbia and Yukon Territory, Canada (Diptera,
Culicidae). Insecutor Inscitiae Menstruus 8:1-27.
Dyar, H.G. 1928. The mosquitoes of the Americas. Carnegie Institute, Washington,
Publication No. 387:1-616.
Ellis, R.A. and R.A. Brust. 1973. Sibling species delimitation in the Aedes communis (DeGeer)
aggregate (Diptera: Culicidae). Canadian Journal of Zoology. 51:915-959.
Enfield, M.A. 1977. Additions and corrections to the records of Aedes mosquitoes in Alberta.
Mosquito News 37:82-85.
Enfield, M.A. and G. Pritchard. 1977. Estimates of population size and survival of immature
stages of four species of Aedes (Diptera: Culicidae) in a temporary pond. Canadian
Entomologist 109:1425-1434.
Gillett, J.D., E.A. Roman and V. Phillips. 1977. Erratic hatching in Aedes eggs: a new
interpretation. Proceedings of the Royal Society of London B 196:223-232.
Gjullin, C.M. and G.H. Eddy. 1972. The mosquitoes of the Northwestern United States. U.S.
Department of Agriculture Technical Bulletin No. 1447. Ill pp.
Government of Alberta and the University of Alberta. 1969. Atlas of Alberta. University of
Alberta Press.
Graham, P. 1969. Observations on the biology of the adult female mosquitoes (Diptera:
Culicidae) at George Lake, Alberta, Canada. Quaestiones Entomologicae 5:309-339.
Happold, D.C.D. 1965. Mosquito ecology in central Alberta. I. The environment, the species,
and studies of the larvae. Canadian Journal of Zoology 43:795-819.
Horsfall, W.R. 1956. Eggs of floodwater mosquitoes (Diptera: Culicidae). III. Conditioning
Mosquito (Diptera, Culicidae) larvae in Southern Alberta
167
and hatching of Aedes vexans . Annals of the Entomological Society of America
49:66-71.
Horsfall, W.R., H.W. Fowler Jr., L.J. Moretti and J.R. Larsen. 1973. Bionomics and
Embryology of the inland floodwater mosquito Aedes vexans . University of Illinois
Press. 211 pp.
Magnarelli, L.A. 1977. Seasonal occurrence and parity of Aedes canadensis (Diptera:
Culicidae) in New York State, U.S.A. Journal of Medical Entomology 13:741-745.
Mailhot, Y. and A. Maire. 1978. Caracterisation ecologique des milieux humides a larves de
moustiques (Culicides) de la region subarctique continental de’Opinaca (territoire de la
Baie de James. Quebec). Canadian Journal of Zoology 56:2377-2387.
Maire, A. 1977. Identification des biotopes a larves de moustiques des tourbieres de la
Basse-Mauricie (Quebec Meridional). Naturaliste Canadien 104:429-440.
Maire, A. and A. Aubin. 1976. Inventaire et classification ecologiques des biotopes a larves de
moustiques (Culicides) de la region de Radison (territoire de la Baie de James, Quebec).
Canadian Journal of Zoology 54:1979-1991.
Maire, A., C. Tessier and L.Picard. 1978. Analyse ecologique des populations larvaires de
moustiques (Diptera: Culicidae) des zones Riveraines du Fleuve SaintLaurent, Quebec.
Naturaliste Canadien 105:225-241.
Mclver, S. 1973. Summary - Behaviour and Ecology of populations. In : Hudson, A. (Editor).
Biting Fly Control and Environmental Quality. Defence Research Board, Ottawa. DR
217. pp. 150-152.
Nielsen, L.T. 1961. Aedes schizopinax Dyar in the Western United States. California
Mosquito Control Association 29:21-24.
Pucat, A. 1965. List of mosquito records from Alberta. Mosquito News 25:300-302.
Rempel, J.G. 1950. A guide to the mosquito larvae of Western Canada. Canadian Journal of
Research D 28:207-247.
Scholefield, P.J., M.A. Enfield, and G. Pritchard. 1979. Identification and distribution of the
aedine mosquitoes of southern Alberta. Alberta Environment Report. 1 14 pp.
Shemanchuk, J.A. 1969. Epidemiology of Western encephalitis in Alberta: response of natural
populations of mosquitoes to avian hosts. Journal of Medical Entomology 6:269-275.
Strickland, E.H. 1938. An annotated list of the Diptera (flies) of Alberta. Canadian Journal of
Research D 16:175-219.
Tawfik, M.S. and R.H. Gooding. 1970. Observations on mosquitoes during 1969 control
operations at Edmonton, Alberta. Quaestiones Entomologicae 6:307-310.
Wada, Y. 1965. Population studies on Edmonton mosquitoes. Quaestiones Entomologicae
1:187-222.
Wood, D.M. 1977. Notes on the identities of some common nearctic Aedes mosquitoes.
Mosquito News 37:71-81.
Wood, D.M., P.T. Dang, and R.A. Ellis. 1979. The Insects and Arachnids of Canada. Part 6.
The Mosquitoes of Canada. Agriculture Canada. 390 pp.
Quaest. Ent., 1981, 17
Patasson luna (Hymenoptera: Mymaridae)
NOTE
RECOVERY OF PATASSON LUNA (HYMENOPTERA: MYMARIDAE)
A PARASITE OF THE ALFALFA WEEVIL, HYPERA POSTICA (COLEOPTERA:
CURCULIONIDAE), IN ALBERTA
The alfalfa weevil, Hypera postica (Gyll.), was introduced into the United States near
Salt Lake City, Utah in 1904 and by 1907 had become a serious pest in Salt Lake County
(Titus, 1910). In southeastern Alberta, this pest was first located in alfalfa fields in the
valley of the Milk River in June 1954. No egg parasites were recorded in studies of the
biology and distrubution of H. postica from 1955 to 1958 (Hobbs et al., 1959). However, in
1978, during studies of the weevil in the Brooks area of Alberta, several specimens of a
parasite were reared from alfalfa weevil eggs. Adults sent to the Biosystematics Research
Institute, Agriculture Canada, Ottawa, were tentatively identified by Dr. Carl M.
Yoshimoto and later confirmed by Mr. M. Schauff of the University of Maryland, College
Park, Maryland, to be Patasson luna (Girault).
The characters used to separate this species from a very closely related species Anaphes
pratensis Foerster, can only be seen under a microscope and only in the females. The two
species can be easily mistaken for each other. This is apparently what happened in several
releases. During the years 1911-1913 and 1925-1928, P. luna that had been collected in
Italy were released in Utah in an attempt to establish a biological control agent for control
of the alfalfa weevil (Chamberlain, 1924a; Clausen, 1956). Patasson luna was probably
confused with A. pratensis in these releases, because, in 1926, A. pratensis was recovered
from several fields near Salt Lake City, Utah (Hamlin et al., 1949), but P. luna was not
recovered at this time. Patasson luna from France was introduced into alfalfa fields in
California around 1933 (Essig and Michelbacher, 1933) but has not been recovered since
its release (K. S. Hagen, per. comm.). However, P. luna was collected in Idaho in 1921 (M.
Schauff, per. comm. 1980; due to their condition, only a tentative identification of P. luna
has been given to these specimens), in Utah by D.W. Davies (Dysart and Day, 1976) in
1973, and in Alberta in 1978.
Patasson luna has been reared from alfalfa weevil eggs in the eastern United States
from Delaware, Indiana, Illinois, Maryland, Massachusetts, New Jersey, New York, Ohio,
Pennsylvania, and West Virginia. In eastern Canada, it has been collected in Ontario and
Quebec. The exact method of establishment in eastern North America has never been
documented. It seems probable that P. luna was introduced into this area with a shipment
of A. pratensis sent to Indiana for control of Hypera nigrirostris (F.) in 1928 (Christie,
1928, p. 43) and that it subsequently spread throughout most of the eastern United States.
At Brooks, Alberta, most seed alfalfa is grown on irrigated land surrounded by mixed
grains to the north and west, and by mixed grain or grasslands to the south and east. No
alfalfa hay is imported into these irrigated areas. Apparently, P. luna entered by migrating
over mountains from Idaho, or possibly from the southeast from Montana along the
Missouri and Milk River systems and then across the grainland and grassland.
In southern France, A. pratensis and Patasson spp. infest eggs of Sitona sp. and Hypera
sp. in alfalfa crops (Aechlimann, 1977). Chamberlain (1924 b) and Hamlin et al., (1949)
Quaest. Ent., 1981, 17
170
Schaber
showed that P. luna and A. pratensis parasitize eggs of the clover leaf weevil, Hypera
punctata (Fabr.) in the United States. When clover leaf weevils become scarce, both
parasites will use eggs of H. postica for oviposition. The egg of H. punctata is much larger
than H. postica and, according to Chamberlain (1924 b), P. luna places two eggs in each
egg of H. postica.
Patasson luna may also use a wide range of Sitona and other Hypera spp. as their
primary host and as these hosts become scarce they migrate and infest H. postica eggs as
they do in Europe.
The overwintering method or stage of this parasite is not known in Alberta, but, since a
few eggs of the clover leaf weevil persist through the winter (Clausen, 1956), it seems
possible that it is in this host that P. luna survives the winter.
During the spring of 1979, no specimens of P. luna were recovered from eggs of the
alfalfa weevil. Therefore, the species is considered at present to be of limited importance in
the control of the alfalfa weevil in Alberta.
REFERENCES
Aeschlimann, J.P. 1977. Notes on Patasson lameerei (Hym: Mymaridae), an egg parasite of
Sitona spp. (Col: Curculionidae) in the Mediterranean Region. Entomophaga
22:111-114.
Chamberlain, T.R. 1924a. Introduction of parasites of the alfalfa weevil into the United States.
United States Department of Agriculture Circular. No. 301. 9 pp.
Chamberlain, T.R. 1924b. Studies of the parasites of the alfalfa weevil in Europe. Journal of
Economic Entomology 17: 623-632.
Christie, G.I. 1928. Forty-first annual report of the Purdue University Agricultural Experiment
Station. Purdue University, Lafayette, Indiana.
Clausen, C.P. 1956. Biological control of insect pests in the continental United States. United
States Department of Agriculture Technical Bulletin. No. 1139: 116.
Dysart, R.J., and W.H. Day. 1976. Release and recovery of introduced parasites of the alfalfa
weevil in eastern North America. United States Department of Agriculture Research
Service, Production Research Report. No. 167: 2.
Essig, E.O., and A.E. Michelbacher. 1933. The alfalfa weevil. California Agriculture
Experiment Station Bulletin. No. 567. 99 pp.
Hamlin, J.C., F.V. Lieberman, R.W. Bunn, W.C. McDuffe, R.C. Newton, and L.J. Jones.
1949. Field studies of the alfalfa weevil and its enviironment. United States Department
of Agriculture Technical Bulletin. No. 975. 84 pp.
Hobbs, G.A., W.O. Nummi, and J. Virostek. 1959. History of the alfalfa weevil, Hypera
postica (Gyll.) (Coleoptera: Curculionidae), in Alberta. The Canadian Entomologist
91: 562-565.
Titus, E.G. 1910. The alfalfa weevil. Utah Agriculture Experiment Station Bulletin. No. 110:
19-72.
Burton D. Schaber
Research Station, Agriculture Canada
Lethbridge, Alberta
T1J 4B1
Book reviews
171
BOOK REVIEWS
Larsson, Sven Gisle. 1978. Baltic Amber - a Palaeobiological Study. Entomograph, vol.
1, 192 pp., 62 text-figures, 12 monochrome plates. Scandinavian Science Press Ltd.,
Klampenborg, Denmark. Price: 120 Danish Kroner, $25.32 Canadian.
One of the reasons that man is so attracted by amber is because it is the earth’s most
stable and therefore oldest natural biological product. As ambers go, European or Baltic
amber, the subject of Larsson’s study, is relatively young (35-40 m. yrs. as opposed to
75-80 m. yrs. for Canadian amber). Nevertheless it has been lying around the shores of
the present Baltic Sea at least 15 times longer than the fossils of the earliest known man
in Kenya, and it is still very much the same as it was when it was first produced by the
extinct pine, Pinites succinitera Conwentz.
Larsson’s attractive book is the first English-language synthesis of biological
information on Baltic amber. It is also the first volume in the Scandinavian Science
Press new series, Entomograph, designed especially for large important entomological
papers such as this. This study builds upon, and in some respects supersedes, Kjell
Ander’s (1942), Die Insekten Fauna des Baltischen Bernsteins nebst damit Verkniipten
Zoogeographischen Problemen, and Andolph Bachofen-Echt’s (1949), Der Bernstein
und sein Einschlusse.
The author’s theme (p. 7) is the rich content of fossils of terrestrial arthropods and
plants that are contained in Baltic amber, and the excellent opportunities these fossils
provide for learning the early history of plant and animal life. Although not stated, the
main purpose of the book appears to be to outline the problems involved in interpreting
the amber biota. It is also apparent throughout the book that a secondary purpose is to
publicize the relatively large, newly amassed collection of Baltic .amber (8000
specimens), derived almost exclusively from Danish coasts, in the Zoological Museum,
Copenhagen.
The book brings to fruition Larsson’s long standing interests in Baltic amber, and
culminates his efforts since 1949 to develop a permanent working collection of scientific
specimens from Denmark. Being a Dane, he is thoroughly familiar with the geography
of the parts of Europe where the amber was produced and deposited, and he brings
first-hand knowledge to bear on those problems. He is a professional entomologist with
close contacts with many other entomologicial specialists in the Zoological Museum,
and this is another essential qualification for fulfillment of his purposes. In developing
the geological and chemical framework provided for his presentation he was guided by
Danish experts in these fields. So the book has a solid foundation.
Larsson treated the subject in five main sections. Section 1 deals with the general
characteristics of amber and resin, its stability and production and its chemical and
physical properties. Section 2 is a treatise on the origins of the various ambers from the
Baltic region. Section 3 treats the original flora of the amber territory. These three
sections occupy 59 pages. Section 4 deals with the original fauna of the amber territory.
This, of course, makes up most of the book (116 pp.), and because of the great
predominance of insects in the Baltic amber inclusions, the book is primarily an
entomological work. The section is divided along ecological lines into seven chapters
entitled Plant Sucking Insects, Leaf-and Seed-Consumers, Gall Producers, Nectar
Quaest. Ent., 1981, 17
172
Seekers, Insects Trapped While Resting, The Fauna of Moss and Bark, and The
Hidden Fauna of Tree Trunks. The final section (Section 5) is a review describing the
nature and conditions of the original amber forest as envisaged by the author. It also
includes a list of the amber material in the Zoological Museum, Copenhagen. Each
section has its own separate list of references (645 in all). An index restricted to the
scientific names of genera and higher taxonomic categories is provided. The book
contains 12 plates of original halftone photgraphs of inclusions (mostly insects) in
amber and 62 line drawings, almost all of which are copied from the literature.
The author succeds to some degree in restating most of the age-old problems
involved in the study of the flora and fauna of Balltic amber, but few new facts or ideas
are introduced. The book has real value as a reference text and it is very useful as a
starting point for finding basic information on many aspects of amber and its inclusions.
On the other hand, it has some weaknesses and faults, and it should not be accepted as
‘the last word’ on the subject. Coverage of the literature is incomplete. Different ideas
and interpretations are frequently presented in such a way that the reader is left
uncertain as to which he favors. For example, I am still not very confident about the
biological status of Pinites succinifera or the extent of its role in the production of
Baltic amber. Much of the discussion on the age, geographical origin and host origin of
Baltic amber from diffetent parts of Europe is in the realm of speculation and the
reader ends up with a feeling of uncertainty and confusion on many points. What is said
in one place is sometimes contradicted in another. For example, it is stated on p. 103
that the only calyptrate fly known from amber belongs to Fabricius’ old and well known
species, Fannia scalaris, and on p. 108 attention is drawn to the fact that one of the
amber fossils of Cicindellidae is identical with the recent species Tetracha Carolina
(L.). However, the final sentence of the Review reads “But the original Baltic amber
fauna appears to be totally extinct.”
The decision to divide the main section on the fauna of Baltic amber into seven
chapters based on ecological considerations rather than on systematic grounds was
unfortunate. This has led to a considerable amount of overlap and repetition, and it has
created difficulty in locating certain data. For example, anyone seeking information on
specific groups such as Coleoptera or Diptera must search through seven chapters. It
would be better if all the arthropods were treated in a standard systematic sequence,
and the ecological correlation were presented in tabular form. Similarly, the decision to
break the bibliography into four sections was not a good one. In such a small book it
would be preferable to consolidate all the references in one composite list. As it is, the
reader must scan four lists if he wishes to locate all the papers cited for any given
author, and of course there is a considerable amount of duplication in the different lists.
The literary quality would have been improved by a good editing, but the book itself is
very well produced. The paper, the printing, and the binding are excellent.
In spite of its shortcomings it is the best single synthesis of paleobiological
information available on Baltic amber, and anyone interested in the subject should have
a copy.
J.F. McAlpine
Biosystematics Research Institute
Agriculture Canada
OTTAWA, Ontario, Canada K1A 0C6
Book reviews
173
GILLOTT, CEDRIC. 1980. Entomology. Plenum Press, New York and London,
xviii + 729 pp. Hard cover $49.50; soft cover $22.50.
Of the several books on general entomology published in recent years, this is one of
the best. Its coverage of basic entomology is extensive, making it an excellent reference
text. The author intends for the book to be used as a “text for senior undergraduates
taking their first course in entomology”, and, indeed, it would be good if such students
were given time to master the meterial in this text in one course. However, it covers too
much ground for a single semester but could be appropriate for a two semester course.
Section I (Chapters 1-10) on evolution and diversity , is perhaps a bit long (300
pages) for an introductory text. For example, Chapter 2 (Insect Diversity) contains 9
page discussion (a good one) on the origin and evolution of insect wings and a discussion
of the Heslop-Harrison theory of pupal origin which concludes with the criticism that
the theory lacks supporting evidence. The lengthy discussion of phylogenetic
relationships of pterygotes is also excellent, but would be difficult reading for those
unfamiliar with the terminology and having little experience with extant orders. Insect
Evolution may have been a better title for Chapter 2 than Insect Diversity.
Chapter 1 (Arthropod Evolution) discusses the three major theories of Arthropod
evolution and comes down firmly in favor of the polyphyletic school. In spite of this, the
major lineages suggested by this theory are not given equal coverage in the rest of the
chapter. Some comments on the relative abundance of arachnids would have been
helpful, in fact, arachnids and crustaceans could have been given equal space. Most of
the drawings in this chapter are unlabelled. When compared with other chapters in the
book, this one is somewhat sketchy, giving an unfortunate first impression.
Chapter 3 (External Structure) presents a well balanced overview of its topic, and
illustrated with relabelled drawings taken largely from Snodgrass’ works. Chapter 4
(Classification and Identification) wisely begins with clear definitions of terms such as
systematics, classification, identification and taxonomy and comments on
disagreements regarding these definitions. A short discussion of natural and artificial
classification schemes follows. The chapter also includes an essay on the history of
insect classification and a key to insect orders. All are well done.
The remainder of Section I (Chapters 5-10) consists of a review of the insect orders,
each covered in the following manner: order name, synonyms, common name and a
short description. These are arranged to stand out clearly from the rest of the text, and
are set in smaller type. Following each order’s description, are short sections on general
structure, life history, phylogeny and classification, and literature (including several
references for each order). Illustrations are appropriate and well done, and the
phylogenetic trees given for most major orders are a helpful addition. These 5 chapters
embody approximately one-third of the book.
Section II (Chapters 11-18) covers aspects of anatomy and physiology in 150 pages.
Chapter 11 is an excellent introduction to the insect integument, with highlights for this
reviewer being the discussions of cuticular structure and color.
Chapter 12 (Sensory Systems) is good as far as it goes. Some illustrations or
photographs and a discussion of sensillar ultrastructure would have enhanced the
presentation, as this area is by now well enough understood to make some
generalizations. The physiology of chemoreception is also better known than this
chapter suggests. Two helpful reviews in this regard, Kaissling (1971) and Hansen
Quaest. Ent., 1981, 17
174
(1978) are not mentioned. Chapter 13 (Nervous and Chemical Integration) presents a
concise and up to date overview of the endocrine system, but falls somewhat short in the
part on nervous integration. The author attempts to summarize the physiology of neural
integration, including a discussion of membrane physiology, in four pages with a single
diagram showing a simple reflex circuit. The result is probably too general to be very
useful. Extensive work has been done on insect walking which could have been used to
illustrate current concepts of nervous integration in insects. Pearson et al. (1973) and
Bowerman (1977) have written a reviews of this area. The discussion on pheromones
could have been improved by including something on the importance of ‘minor’
pheromone components. The review by Seabrook (1978) could have helped here.
Chapter 14 (Muscles and Locomotion), is generally well done, though a near classic
work on insect flight by Nachtigall (1974) seems to have been overlooked.
Chapter 15 presents an excellent overview of respiration in both terrestrial and
aquatic insects. Chapter 16 (Food Uptake and Utilization) includes gut morphology,
gut physiology and a discussion on metabolism, all are well done. Food selection and
feeding, however, are treated quite superficially superficially. Insect host-plant
relationships have been given considerable attention by ecologists, behaviourists and
physiologists for several decades, and recent advances in these areas would form an
excellent basis for a discussion of feeding behaviour. More serious is the lack of
reference to Dethier (1976). In this book, “The Hungry Fly” extensive studies on
feeding behaviour and related physiology covering more than 20 years are clearly
summarized.
Circulation, excretion and water balance are given good coverage in Chapters 17 and
18, while Chapters 19-21 present a thorough treatment of reproduction and
development. The discussion of environmental and endocrine factors affecting female
maturation and the figures comparing endocrine relationships in 4 insect species are
excellent.
Two chapters (22 and 23) on the abiotic and biotic environment give a good
introduction to classical insect ecology. The final chapter, entitled Insects and Man,
considers beneficial and harmful insects and stresses the importance of integrated pest
management. Major problems with chemical control, particularly resistance to
insecticides, are highlighted, and a good discussion of biological control with a table of
suitable examples is presented.
REFERENCES
Bowerman, R.F. (1977) Control of arthropod walking. Comp. Biochem. Physiol. 56A: 231-247.
Dethier, V.G. (1976) The Hungry Fly. Harvard University Press, Cambridge, London.
Hansen. K. (1978) Insect chemoreception, In: Receptors and Recognition, Ser. B. Vol. 5. Taxis
and Behaviour. G.L. Hazelbauer (ed.). Methuen.
Kaissling, K.E. (1971) Insect olfaction. In: Handbook of Sensory Physiology. IV, Chemical
Senses, Part 1, Olfaction. L.m. Beidler (ed.). Springer, Berlin, Hidelberg, New York.
Nachtigall, W. (1974) Insects in Flight. George Allen and Unwin Ltd., London.
Pearson, K.G., Fourtner, C.R. and Wong, R.K. (1973) Nervous control of walking in the
Book reviews
175
cockroach. In: Control of Posture and Locomotion. R.B. Stein, K.G. Pearson, R.S.
Smith, J.B. Redford (eds.). Plenum, New York.
Seabrook, W.D. (1978) Neurobiological contributions to understanding insect pheromone
systems. Ann. Rev. Entomol. 23: 471-485.
B.K. Mitchell
Department of Entomology
The University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Quaest. Ent., 1981, 17
NOTICE
PIFON, A NEW PERMANENT INTERNATIONAL FILE OF NATURALISTS
In 1980 a new institution, the Oxycopis Pond Research Station was established in a
wooded area in upstate New York. Among other projects the Institute compiles and
maintains a coded information file of naturalists of the world, PIFON (acronym for
Permanent International File of Naturalists). This international register includes well
over 10,000 contemporary naturalists, and in addition a file of all persons listed in The
Naturalist’s Directory ( International ) since the first edition in 1877. Thus PIFON is
the world’s most complete listing of naturalists, past and present. This wealth of data is
available for the use of any person registered in PIFON.
The 44th edition of The Naturalists’ Directory ( International ) is being compiled
from data in PIFON. All 10,000 persons were contacted in 1980 and all institutions in
1981 to urge registration in PIFON. Now the six 'parts of the new 44th edition (to be
called volume 44) are being published. The six parts are: Part I: The Naturalists’
Directory of Insect Collectors and Identifiers ( International ); Part II: The Naturalists’
Directory of Plant Collectors and Identifiers ( International ); Part III: The Naturalists’
Directory of Bird Watchers ( International '); Part IV: The Naturalists’ Directory of
Field Biologists ( International '); Part V: The Naturalists’ Directory of Rock, Mineral,
and Fossil Collectors ( International '); Part VI: The Naturalists’ Directory
( International '): Subject, Geographical, and Name Index. Each part is complete in
itself, but the last part is a general index and cross-reference for the other five parts for
individuals and libraries using two or more parts.
All persons registered in PIFON will receive a registration card showing their
unique registration number. Among the other advantages of being registered is access
to the unpublished data in the PIFON file. Registration is free. A charge of $1.00 to
cover cost of photocopying and mailing is required of those using the file. Requests may
be made at anytime by writing or by phone.
To be registered in PIFON send the following information (please use block letters
or type): 1) name (last or family name underlined; include Dr., Prof., Mr., Mrs., Ms.,
etc.); 2) mailing address (shortest form); 3) phone number (include area code); 4)
group of main interest, area of specialization (e.g., family group name, etc.) and kind of
interest (e.g., collecting, exchanging, systematics, ecology, etc.); 5) geographical area of
greatest interest; 6) short statement giving interest details (not over 25 words); 7) if a
taxonomist, list groups willing to identify for others; 8) signature and date. These data
will be coded and entered into the file. Those already listed should send changes as they
occur. No person is listed unless these data are supplied hy that person The right to edit
all listings published in the Directory is reserved.
The deadline for submitting entries for each edition is three months before
publication. Parts will be issued every four months, resulting in a two year cycle for the
entire volume. Regardless of the deadline dates, however, persons are urged to register
immediately upon reading this note, thus making available their data to others using the
PIFON file.
178
Notice
The parts of the volume are sold by the publisher, Flora and Fauna Publications, by
subscription only, either as single parts, or multiple parts, with a discount for
subscriptions to the entire volume.
Send registration and/or requests for further information to the compiler and
ediitor: Dr. Ross H. Arnett, Jr., Oxycopis Pond Research Station, 90 Wallace Road,
Kinderhook, NY 12106 USA. (Phone: (518) 758-7219).— R.H.A.
Ouaest. Ent. 1981. 17
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 Edition, 1964, except that titles of periodicals should be
given in full. For style of taxonomic papers, the Editor should be consulted.
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The Editor, Quaestiones Entomologicae
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Second Class Mail Registration Number 5222
He i
QI3
Ba^
Quaest
lones
Entomolog
icae
A periodical record of entomological investigations#
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 17
NUMBER 3-4
JULY-OCTOBER 1981
CONTENTS
Lafontaine-Classification and phylogeny of the Euxoa detersa group (Lepidoptera:
Noctuidae) 1
Oliver-Redescription and systematic placement of Oreadomyia albertae Kevan and
Cutten-Ali-Khan (Diptera: Chironomidae 121
Corkum and Clifford-Function of caudal filaments and correlated structures in mayfly
nymphs, with special reference to Baetis (Ephemeroptera) 129
Scholefield, Pritchard and Enfield-The distribution of mosquito (Diptera, Culicidae) larvae in
Southern Alberta, 1976-1978 147
Note-Schaber-Recovery of Patasson luna (Hymenoptera: Mymaridae) a parasite of the alfalfa
weevil, Hypera postica (Coleoptera: Curculionidae), in Alberta 169
Book Review-Larsson, Sven Gisle. 1978. Baltic Amber - a Palaeobiological Study.
Entomograph, vol. 1 171
Book Review-Gillott, Cedric. 1980. Entomology 173
Hudson, J.E.-Studies on flight activity and control of mosquitoes in the Edmonton area,
1971-73 179
Fredeen, F.J.H.-Keys to the black flies (Simuliidae) of the Saskatchewan River in
Saskatchewan 189
Doane, J.F.-Seasonal captures and diversity of ground beetles (Carabidae: Coleoptera) in a
wheat field and its grassy borders in central Saskatchewan 211
Carter, A.-Aspects of the comparative ecology of populations of four Patrobus species
(Coleoptera: Carabidae: Patrobini) at George Lake, Alberta 235
Roughley and Pengelly-Classification, phylogeny, and zoogeography of Hydaticus leach
(Coleoptera: Dytiscidae) of North America 249
Book Review-Tschirnhaus, M von. 1981. Die Halm- und Minierfliegen im Grenzbereich
Land-Meer der Nordsee 315
Editor’s Acknowledgements 317
Index 319
STUDIES ON FLIGHT ACTIVITY AND CONTROL OF MOSQUITOES IN THE
EDMONTON AREA, 1971-73
J.E. Hudson
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Quaestiones Entomologicae
17:179-188 1981
ABSTRACT
The effects on mosquito activity of barrier strips of dichlorvos and propoxur sprayed on
the vegetation in wooded ravines, were assessed by means of insect wind traps and by landing
counts on human bait. Dichlorvos at 1.12kg a.i./ha reduced trap catches by 70% for one day
after treatment; bite count data were inconclusive. Propoxur at 0.34kg a.i./ha reduced trap
catches by 44% and bite counts by 100% for one day after treatment. From these results a
continuation of the barrier strip method could not be recommended. Many other insects
( chiefly other Diptera) were caught in the traps, but there were no consistent reductions in
catches in the treated strips. Catches of mosquitoes were consistently higher on the
down-valley sides of the traps than on the up-valley sides, and catches were generally higher in
traps in the ravines than in traps on the plain above. The effectiveness of malathion sprayed by
helicopter was measured by kills of caged, wild-caught mosquitoes and by bite counts. Boom
and nozzle application at 0.7kg a.i./ha killed only 27 % of the mosquitoes (mostly Aedes
vexansj caged in the woods at Whitemud Creek. Ultra-Low-Volume application at 0.3kg
a.i./ha killed only 8% of the caged mosquitoes (mostly Aedes fitchiij at Whitemud Creek, but
94 % of the caged mosquitoes at Victoria Park golf course, a more open area.
RESUME
Le dichlorvos et le propoxur furent appliques sur des bandes de vegetation en ravin boise, et leurs effets sur iactivite
des maringouins furent etudies a I’aide de pieges eoliens, et en denombrant les individus atterrissant sur un appat
humain. Le dichlorvos, applique a raison de 1.12 kg i.a./ha, reduisit les captures de 70% pendant une journee suivant le
traitement; les resultats du denombrement des piqures ne furent pas concluants. Le propoxur en concentration de 0.34 kg
i.a./ha reduisit les captures de 44%, et le nombre de piqures de 100%, pendant une journee suivant I’application. Ces
resultats ne permettent cependant pas de recommander I’arrosage de bandes de vegetation pour controler les moustiques.
De nombreux autres insectes (surtout des Dipteres) furent captures dans les pieges, mais il n’y eut pas de diminution
uniforme des captures dans les bandes traitees. Les captures de moustiques furent toujours plus elevees pres du fond que
pres du rebord des ravins, et furent generalement plus abondantes dans les pieges places dans les ravins que dans ceux
situes sur la plaine environnante. L’efficacite du malathion repandu par helicoptere fut evaluee h partir de la mortalite
de moustiques sauvages encages, et du denombrement des piqures. L’epandage a Vatomiseur d’une solution concentree h
0.7 kg i.a/ha ne tua que 27%, des maringouins (principalement Aedes vexansj encages dans un bois a Whitemud Creek.
L’epandage a tres faible volume d’une solution a 0.3 kg i.a/ha ne tua que 8%> des maringouins ( surtout Aedes fitchiij
encages a Whitemud Creek, mais tua 94% de ceux encages au terrain de golf de Victoria Park, qui est un endroit plus
decouvert.
180
Hudson
INTRODUCTION
According to Klassen and Hocking (1964), the North Saskatchewan River valley and its
tributary ravines are a major route of entry of adult mosquitoes into the city of Edmonton,
because local air currents and the mosquitoes’ optomotor responses bring them down from the
plains into the valleys, and the prevailing winds at night are down-valley. To stop the mosquito
invasions, Klassen and Hocking applied lindane by aircraft, at 0.3kg/ha, to vegetation on the
sides of the river valley, in three 90m wide barrier strips at the eastern and three at the western
city limits. From bite counts and observations on male swarms, it was concluded that the
barrier strips were effective, (Klassen and Hocking, 1963), and the method was used, without
further evaluation, by Edmonton Parks and Recreation from 1959 to 1969, when lindane was
banned. This report describes attempts in 1971 and 1972 to reassess the barrier strip method of
control, and to test two substitutes for lindane, using traps designed to measure insect flight
activity and direction, (Hocking, 1970; Hocking and Hudson, 1974).
In the meantime, the City of Edmonton began using aerial sprays of Malathion against
adult mosquitoes, and this report also describes attempts to measure the effectiveness of these
sprays.
MATERIALS AND METHODS
The 1971 barrier strip experiment was performed in a wooded ravine which meets the North
Saskatchewan River Valley on the south side at a point 18 km southwest of the City of
Edmonton and 3 km east of Devon. The 1972 experiments were performed in the Whitemud
Creek Valley, which runs North to Edmonton, at 0.8 and 1.6 km south of the city limits. The
ravines at the two test sites were about 300m wide, and thickly wooded with aspen, poplar and
spruce, with small streams running through.
Two insecticides, dichlorvos (Vapona 20% e.c.,1 2,2 - dichlorovinyl dimethyl phosphate), and
propoxur (Baygon 70% w.d.p.,2 ortho-isopropoxyphenyl methyl carbamate) were chosen for the
tests because both are quick acting and have a fumigant action, properties they share with
lindane. For the dichlorvos trial, a rectangular plot 150 x 70m was marked out on the west slope
of the ravine near Devon, and on the 24 July 1971 all the vegetation in the outermost 10m of
this plot was treated with the dichlorvos emulsion from hand compression sprayers, at a dosage
of 1.11kg a.i.3 in 200 liter/ha. An insect trap was located in an unsprayed area 130 x 50m
within the plot. A similar plot on the eastern slope was sprayed with the insecticide solvent only
in water. For the propoxur trial, a 200m wide strip across the Whitemud Creek ravine, 0.8km
south of the city limits, was sprayed by a helicopter with boom-and-nozzle equipment, at a
dosage of 0.3kg a.i. in 39 liters of water per hectare, at 22:30 hrs on the 28 May 1972. A third
trial in June 1972 at Whitemud Creek to re-evaluate lindane was abandoned because of a
mistake by the helicopter pilot. A fourth trial in July at the same site was to test a new variant
of the trap, not an insecticide.
'Emusifiable Concentrate
2Wettable powder
3a.i. = active ingredient
Mosquitoes in the Edmonton area
181
Three variations of insect wind traps (Hocking, 1970) were used: the large (10m2) and
medium-sized (2m2) ones were set up with one side facing up-valley and the other side
down-valley, while the small ones (0.5m2, pivoted and wind-vaned) were set up in small
clearings in the forest so they could turn freely. For the dichlorvos trial six traps were used, one
large and two small on the west (treated) and the same numbers on the east (untreated) slope
of the ravine. For the propoxur trial, five traps were used: one small and two large traps on the
ravine bottom (one large inside the treated strip); and two small traps on the plain above, one
each side of the ravine. Traps were emptied once or twice daily, and mosquito biting counts
made near the traps when they were emptied. Mosquitoes from the traps were identified with
the aid of Carpenter and LaCasse (1955).
The effectiveness of the malathion sprays was assessed by putting wild-caught Aedes
females into cylindrical cages made from 1.4mm mesh galvanized mosquito screen and 10cm
plastic petri dishes and the cages were hung from trees and bushes in the areas to be sprayed.
One hour after spraying, the mosquitoes were transferred to clean plastic cups and held in the
laboratory for another 24h before the mortality count was made. Spray droplets were sampled
by white glossy cards (Kromekote Bristol 29 1M), pinned to leaves near the cages, and a few
cards were put into empty bioassay cages to measure the droplet penetration rate. The
helicopters were fitted in 1972 with boom-and-nozzle equipment to spray malathion emulsion
in drops of 500 microns at 0.7kg a.i. in 46 1 of water per hectare, in 1973 with Beecomist
ultralow volume (ULV) rotary atomizers to spray undiluted technical malathion in drops of 40
microns at 0.3kg/ha.
Reductions in trap catches and bite counts were calculated according to the formula:
% reduction = 100(1 - (Tn Co / To Cn))
where Tn and Cn are the numbers caught in the treated and untreated areas, respectively,
on day n after treatment, and To and Co are the corresponding values for the day before
treatment. Mortalities of caged mosquitoes in the malathion-sprayed areas (T) were
corrected for mortalities of mosquitoes caged in the unsprayed areas (C) by Abbott’s
formula, as follows:
Corrected mortality = 100(T - C/100 - C)
RESULTS
The wind traps caught many insects (Table I), and the mean catch per m2 at Devon was the
highest obtained anywhere in the world, except Tahiti (Hocking, 1970). Of the catch at Devon,
72.4% of the total were female mosquitoes, 97.4% of them Aedes vexans (Meigen), the worst
pest mosquito in western Canada. In the trials at Whitemud Creek, female mosquitoes, which
never formed more than 15% of the total catch, were mostly Aedes cataphylla in May, Aedes
fitchii in June and Aedes vexans again in July.
In the dichlorvos experiment there was a 69.8% reduction in trap catch in the treated plot
only on the day after treatment (Table II). There was no reduction in catch in the traps
up-valley and down-valley from the treated plot. The bite counts were too low to draw any
conclusions about the effectiveness of the treatment.
In the propoxur experiment there was a 43.8% and 100% reduction in the trap catches and
in the biting rates, respectively, in the treated strip the day after treatment (Table III). On the
second day after treatment, both flight and biting activity within the treated strip returned to
pre-treatment levels.
Quaest. Ent. 1981, 17 (3,4)
182
Hudson
The wind traps have the advantage that they can also detect changes in the activity for
non-target insects, but the results obtained in the propoxur trial were inconsistent. On the day
after treatment, there was a reduction in the catch of other Diptera in the treated strip, but no
reduction in the catch of Hymenoptera, and the catch of Lepidoptera actually increased.
According to Klassen and Hocking (1964) bigger catches could be expected in the traps in
the ravines than on the plain, and bigger catches on the up-valley sides of the traps (i.e.
mosquitoes flying down-valley) than on the down-valley sides. The 10m2 and 2m2 traps gave
consistently higher catches in the ravine than on the plain, but the 0.5m2 traps gave higher
catches on the plain, (Table IV). In three of four trials, catches were greater in the sides of the
traps facing down-valley, i.e. the direction of flight was predominantly up the valley, and
against the prevailing night winds, (Table V).
At Whitemud Creek in July 1972, malathion applied by boom-and-nozzle killed only 27.4%
of the caged mosquitoes, 84% of which were Aedes vexans, (Table VI), and four hours after
spraying the biting rate in the treated area (4.5/min.) was almost as high as in an untreated
area nearby (5.5/min). In the 1973 tests, 97% of the caged mosquitoes used were Aedes fitchii.
At Whitemud Creek, the average corrected mortality in treated area was only 8% (average
uncorrected 26%, average for untreated area 25%), and 0.09 droplets/cm2 were found on the
test cards. At Victoria Park, the average mortality of the caged mosquitoes was 94% and 0.26
droplets/cm2 were found on the cards. From the numbers of the droplets found on the cards
inside cages, it was estimated that the penetration rate was only 35.4%. In both 1973 tests,
mosquitoes bit us in the treated area less than one hour after treatment, and 23% of 48
mosquitoes caught in the treated area died after 24hr holding in the laboratory, compared with
2% mortality in 41 mosquiitoes from an untreated area.
DISCUSSION
A barrier strip of insecticide should kill all the mosquitoes attempting to cross it, for several
days after treatment, but the trap catches indicated reductions in mosquito activity for only one
day after treatment, and no measurable reduction in mosquito activity up-valley or down-valley
from the strips. The reasons for these poor results are uncertain and require further
investigation, but on this basis we could not recommend the resumption of barrier strip
spraying with dichlorvos and propoxur. Both dichlorvos and propoxur have very short residual
life. Lindane was more effective because of residual properties. Dichlorvos is unstable in the
presence of light and water. Propoxur is more stable but the volume of spray, 39 liters/ha, was
insufficient for adequate leaf coverage. Brown (1951) states that 50 gallons/acre
( = 5621itres/ha) are required for adequate leaf coverage in woodland. Another important
factor in the Edmonton region is the vulnerability of the spray deposits to the summer rains.
Two attempts were made in 1972 to bioassay the residual toxicity of propoxur and lindane
deposits on leaves, but on both occasions the toxicity of the deposits disappeared within 48h,
after heavy rains.
The insect wind traps were chosen for the trials because of their ability to sample a wide
selection of flying insects, and to measure both flight density and direction of flight. However,
in practice the traps proved to be less suitable for these studies than they had seemed. The ratio
of mosquitoes to other insects was low in 3 of the 4 trials. The pivoted 0.5m2 traps did not keep
facing properly into the light winds down in the ravines, though later a lighter, more efficient
wind vane was developed, (Hocking and Hudson, 1974). In studies elsewhere, Hocking found
that sometimes insects which had been flying upwind were caught in the upwind, not the
Mosquitoes in the Edmonton area
183
downwind, sides of the traps, (Hocking and Hudson, 1974). The three variants of the trap gave
different results in the same place for flight densities of mosquitoes, (see Table V), the 2m2
j fixed, pyramidal, traps giving the highest values per m2-hr. This was probably due to
differences in area distribution with height. For future studies of mosquito dispersal it would be
better to have a larger number of smaller traps, more specific for mosquitoes.
The boom-and-nozzle application of malathion was ineffective, probably because of the very
large droplets produced. The low temperature at the time of spraying, 10°C, was also
unfavourable. The ultralow volume equipment produced drops of around 40 microns in
diameter, which is much closer to the optimum of 25 microns for adult mosquitoes, (Weidhaas
[ et al. 1970). Malathion ULV sprayed at 0.3 kg/ha. gave satisfactory kills of caged mosquitoes
at Victoria Park golf course, a fairly open area, but not in the thickly wooded ravine at
Whitemud Creek. In both areas, mosquitoes were taken biting less than one hour after
treatment. These mosquitoes may have been protected by resting deep in the undergrowth,
below the level of the caged mosquitoes (about 1.5m). The ULV application of malathion was
i adopted as routine by Edmonton Parks and Recreation, with a higher dosage (0.5kg/ha) for
thickly wooded areas.
ACKNOWLEDGEMENTS
These studies were supported by grants from the Alberta Agricultural Research Trust and
City of Edmonton to the late Dr. Brian Hocking. Assistance of Hugh G. Philip, Brian M.
Rolseth, Bernard Boisvert and Jacqueline Marr is acknowledged. The propoxur wettable
powder was kindly donated by Chemagro.
REFERENCES
Brown, A.W.A.. 1951. Insect control by chemicals. New York, John Wiley. 817 pp.
Carpenter, S.J. and W.J. LaCasse. 1955. Mosquitoes of North America (North of Mexico).
Berkeley, University of California Press. 360 pp.
Hocking, B. 1970. Insect flight and entomologists’ inheritance. Ent. News 81:269-278.
Hocking, B., and J.E. Hudson. 1974. Insect wind traps: improvements and problems.
Quaestiones Entomologicae 10:275-284.
Klassen, W., and B. Hocking. 1963. Control of Aedes dispersing along a deep river valley.
Mosquito News 23:23-26.
Klassen, W., and B. Hocking. 1964. The influence of a deep river valley system on the dispersal
of Aedes mosquitoes. Bulletin of Entomological Research 55:289-304.
Weidhaas, D.E., M.C. Bowman, G.A. Mount, C.S. Lofgren and H.R. Ford. 1970. Relation of
minimum lethal dose to the optimum size of droplets of insecticides for use in mosquito
control. Mosquito News 30:195-200.
Quaest. Ent. 1981, 17 (3,4)
184
Hudson
Table I. Catches of mosquitoes and other arthropods in wind traps at Devon, 1971, and at
Whitemud Creek, 1972.
!Trial stopped because of pilot error
2In May 1972 mostly Ae. cataphylla
* Aedes fitchii plus Ae. mercurator
Mosquitoes in the Edmonton area
185
Table II. Catches of both sexes of mosquitioes in wind traps, and landing counts of females, in a
dichlorvos treated and an untreated plot near Devon, July 1971
!Sample not taken
Quaest. Ent. 1981,17(3,4)
186
Hudson
m
Table III. Catches of mosquitoes (both sexes) in wind traps and bite counts before and after
propoxur spraying, Whitemud Creek, May 1972.
Sample not taken.
Mosquitoes in the Edmonton area
187
‘Sample not taken.
Table V. Catches of mosquitoes of both sexes, per m2-hr, in the large insect wind traps (10m2
variant) sited in the ravines on the up-valley and down-valley sides of the traps
Quaest. Ent. 1981, 17(3,4)
188
Hudson
Table VI. Assessment of aerial sprays of malathion against mosquitoes in the Edmonton area,
1972-1973.
Corrected for control mortality by Abbott’s formula
KEYS TO THE BLACK FLIES (SIMULIIDAE) OF THE SASKATCHEWAN RIVER IN
SASKATCHEWAN1
F.J.H. Fredeen
Canada Agriculture Research Station
107 Science Crescent
Saskatoon, Saskatchewan, Canada Quaestiones Entomologicae
S7N0X2 17:189-210 1981
ABSTRACT
Illustrated keys to the larvae, pupae and adults of 15 species of black flies known to occur
in the North, South and Main Saskatchewan Rivers in Saskatchewan are presented. Keys are
based mainly upon external characteristics. Life histories, trends in abundance and economic
importance are discussed. Species treated include Ectemnia taeniatifrons (Enderlein),
Metacnephia saskatchewana Shewell and Fredeen, Simulium euryadminiculum Davies, S.
duplex Shewell and Fredeen, S. meridionale Riley, S. rugglesi Nicholson and Mickel, S.
bivittatum Malloch, S. griseum Coquillett, S. vittatum Zetterstedt, S. arcticum Malloch, S.
luggeri Nicholson and Mickel, S. decorum Walker, S. tuberosum (Lundstroem), S. venustum
Say, and S. verecundum Stone and Jamnback.
RESUME
Clefs illustrees pour les larves, les pupaes, et les adultes de chacune des quinze especes des mouches noires connues
d’exister dans les trois Rivieres de la Saskatchewan: la Nord, la Sud, et la Principale, sont presentees. Les clefs se basent
principalement sur des charateristiques externelles. Des histoires naturelles, des tendanes d’abondance et de la
signification economique sont discutees. Les especes sont Ectemnia taeniatifrons (Enderlein), Metacnephia saskatchewana
Shewell et Fredeen, Simulium euryadminiculum Davies, S. duplex Shewell et Fredeen, S. meridionale Riley, S. rugglesi
Nicholson et Mickel, S. bivittatum Malloch, S. griseum Coquillett, S. vittatum Zetterstedt, S. arcticum Malloch, S.
luggeri Nicholson et Mickel, S. decorum Walker, S. tuberosum ( Lundstroem ), S. venustum Say, et S. verecundum Stone
et Jamnback.
INTRODUCTION
For some 30 years the Agriculture Canada Research Station at Saskatoon has been involved
in research on black flies inhabiting the Saskatchewan River (Map 1), particularly those
species breeding in numbers sufficient to create economic problems. The river has changed
considerably during this period, partly because of construction of hydropower dams and has
trended toward relatively shallow, weedy conditions. Parallel with these changes, numbers of
black fly species have increased, and dominance has shifted from Simulium arcticum Malloch
to Simulium luggeri Nicholson and Mickel. This shift created new problems for residents in a
large area of Saskatchewan because Simulium luggeri females attack humans as well as other
animals and remain abundant all summer unlike Simulium arcticum. Direction of future
changes will depend upon climatic conditions and upon artificial manipulations of the river, but
'Contribution No. 803, of the Research Station, Saskatoon
190
Fredeen
Map 1. The North, South, and main Saskatchewan Rivers, in Saskatchewan.
Black flies (Simuliidae) of the Saskatchewan River
191
because of its large size the Saskatchewan River has potential for creating large black fly
outbreaks. Thus, surveillance, research and abatement may be required for many more years.
These keys to larvae, pupae, and adults of the 15 species known to inhabit the river were
prepared to aid future studies. External features are emphasized to facilitate sorting large
samples. Taxonomic concepts and nomenclature of structures follow those used by Davies et al.
(1962), Wood et al. (1963), Stone et al. (1965), Davies (1968), Stone and Snoddy (1969) and
McAlpine et al. (1981). The keys are based upon specimens stored at the Agriculture Canada
Research Station, Saskatoon, augmented by published descriptiions.
CHECKLIST OF SPECIES OF SIMULIIDAE FROM THE SASKATCHEWAN RIVER
SYSTEM, SASKATCHEWAN, CANADA.
1 . Ectemnia taeniatifrons (Enderlein)
2. Metacnephia saskatchewana Shewed and Fredeen
3. Simulium (Eusimulium) euryadminiculum Davies
4. Simulium (Eusimulium) duplex Shewed and Fredeen
5. Simulium (Byssodon) meridionale Riley
6. Simulium ( Byssodon ) rugglesi Nicholson and Mickel
7. Simulium (Psilopelmia) bivittatum Malloch
8. Simulium (Psilopelmia) griseum Coquillett
9. Simulium (Psilozia) vittatum Zetterstedt
10. Simulium (Gnus) arcticum Malloch
1 1 . Simulium (Phosterodoros) luggeri Nicholson and Mickel
12. Simulium (sensu stricto) decorum Walker
13. Simulium (sensu stricto) tuberosum (Lundstroem)
14. Simulium (sensu stricto) venustum Say
15. Simulium (sensu stricto) verecundum Stone and Jamnback
KEYS TO GENERA AND SPECIES
ADULTS, Males
1 Entire vein R of wing clothed with hairs (setae) dorsally (Fig. 1) 2
I' Basal section of vein R (Rb, Fig. 1) bare 5
2 (1) Ratio of length of basal section of radius (Rb, Fig. 1) to length of remainder of
wing 1:2.5-1:3.0; basal cell present (Fig. 1); calcipala on first hind tarsomere
narrow, pointed (Fig. 2); pedisulcus on second hind tarsomere insignificant 3
2' Ratio of length of basal section of radius to length of remainder of wing 1:2.9 -
1:3.9; basal cell absent; calcipala on first hind tarsomere broadly rounded (Fig.
3); pedisulcus deep 4
3 (2) Size smaller, wing 2.0-3.0 mm long; Sc setose ventrally; gonostylus with one
minute terminal tooth (Fig. 4)
Metacnephia saskatchewana Shewed and Fredeen
3' Size larger, wing 4.0 - 4.5 mm long; Sc bare ventrally; gonostylus with two
terminal teeth Ectemnia taeniatifrons (Enderlein)
Quaest. Ent., 1981, 17 (3,4)
192
Fredeen
4 (2') Mesepimeral tuft (Fig. 5) pale; ventral plate conical and about as wide as base
of gonostylus, markedly convex and with ventral surface clothed in long pale
bristles Simulium duplex Shewed and Fredeen
4' Mesepimeral tuft dark; ventral plate broadly rectangular, more than two times
wider than base of gonostylus, ventral surface bare, shining, concave
Simulium euryadminiculum Davies
5 (F) Gonostylus with three or more teeth and almost square; scutum with two silvery
pollinose vittae Simulium vittatum Zetterstedt
5' Gonostylus with one or no teeth and longer than wide in most species; scutum
generally without distinct vittae 6
6 (50 Scutellum yellow; lateral and anterior margins of scutum yellow; gonostylus
flat, quadrangular, shorter than gonocoxite; ventral plate broad and
membranous 7
6' Scutellum and margins of scutum not yellow; gonostylus more or less
cylindrical, longer than wide and as long as, or longer than gonocoxite; ventral
plate variously shaped 8
7 (6) Integument predominantly black; central area of scutum black with two
anterior, lance-shaped, silvery-pollinose spots or short vittae
Simulium bivittatum Malloch
T Integument predominantly greyish-yellow; scutum greenish-grey with two
anterior, rectangular, indistinctly pollinose spots
Simulium griseum Coquillett
8 (6') Ventral plate relatively broad with basal arms extended laterally, slightly
beyond margins of plate 9
8' Ventral plate spike or tooth-shaped with basal arms extended laterally well
beyond margins of plate 11
9 (8) Scutum velvety black suffused with ashen pollinosity especially on anterior half
(best seen with horizontal illumination); scutum without prominent shining,
white bars or spots, but with three indistinct dark vittae; scutum densely covered
with golden hairs; gonostylus without prominent medio-basal lobe
Simulium meridionale Riley
9' Scutum velvety black with prominent shining white bar near each anterior
corner; scutum sparsely clothed with golden or brown hairs; gonostylus with
prominent medio-basal lobe 10
10 (9') Medio-basal lobe on gonostylus covered with fine hairs only
Simulium rugglesi Nicholson and Mickel
10' Medio-basal lobe on gonostylus covered with short stout spines
Simulium tuberosum (Lundstroem)
1 1 (8') Ventral plate markedly compressed laterally, “Y”-shaped when seen in ventral
view 12
IF Ventral plate tooth-shaped in ventral view 13
12 (11) Ventral plate not prominently hairy; gonostylus about three times longer than
wide; scutum with single pair of silvery-pollinose spots extended from anterior
corners toward center of scutum
Simulium arcticum Malloch
Ventral plate prominently hairy; gonostylus about two times longer than wide;
12'
Black flies (Simuliidae) of the Saskatchewan River
193
scutum with two pairs silvery pollinose spots, each occupying more than one
third of disc Simulium decorum Walker
13 (IT) Posterior quarter of scutum with sparse, fine hairs; arms of ventral plate each
with postero-medial projection Simulium luggeri Nicholson and Mickel
13' Posterior quarter of scutum with strong, erect hairs; arms of ventral plate
without prominent lateral projections 14
14 (130 Ventral plate relatively broad with approximate outline of molar tooth; notched
margins of keel of ventral plate flared outward in “V”-shape
Simulium venustum Say
14" Ventral plate laterally compressed as narrow keel; notched margins of keel
turned inward and parallel with one another
Simulium verecundum Stone and Jamnback
ADULTS, Females
1 Entire vein R of wing clothed with hairs (setae) dorsally (Fig. 1) 2
Y Basal section of vein R bare 5
2 (1) Basal cell present (Fig. 1); ratio of length of basal section of radius (Rb, Fig. 1)
to length of remainder of wing 1:2.5-1:3.0; calcipala on first hind tarsomere
narrow and pointed; pedisulcus on second hind tarsomere shallow and indistinct
(Fig. 2) 3
2' Basal cell absent; ratio of length of basal section of radius to length of remainder
of wing 1:2.9— 1:3.9; calcipala on first hind tarsomere prominent except in S.
vittatum and rounded; pedisulcus on second hind tarsomere deep (Fig. 3) 4
3 (2) Frons with nearly parallel sides, narrowest near middle (Fig. 6); size larger
(wing 4.0-4. 5 mm long); R dorsally with setae, but lacking black spinules ....
Ectemnia taeniatifrons (Enderlein)
3' Frons wedge-shaped, narrowest anterio-ventrally (Fig. 7); size smaller (wing
2.0-3.0 mm long); R dorsally with setae plus black spinules
Metacnephia saskatchewana Shewed and Fredeen
4 (2') Medial and two lateral vittae on scutum narrow, black
Simulium duplex Shewed and Fredeen
4' Medial and two lateral vittae on scutum narrow, white
Simulium euryadminiculum Davies
5 (F) Claw with prominent, thumb-like basal projection (Fig. 8) 6
5' Claw simple or with minute sub-basal tooth (Figs. 9, 10, 1 1) 7
6 (5) Foreleg entirely dark; width of frons at narrowest point about one-quarter as
long as frons; frons and terminal abdominal terga grey-pollinose; scutum
densely grey, pollinose with three narrow, brown vittae
Simulium meridionale Riley
6' Fore coxa and about three-quarters of tibia pale yellow; width of frons at
narrowest point about one-half as long as frons; frons and terminal abdominal
terga shining dark brown; scutum thinly grey pollinose, subshining and without
three narrow brown vittae Simulium rugglesi Nicholson and Mickel
7 (57) Small yellowish flies with postnotum contrastingly dark and with dark dorsal
spots on abdomen and ends of some leg articles 8
Quaest. Ent., 1981,17(3,4)
194
Fredeen
Fig. 1. Wing of Ectemnia taeniatifrons (Enderlein), showing: basal cell (b); stem vein (Sv); basal section of radius (Rb);
subcosta (Sc); costa (C); radial sector (Rs); radius (R); media (M,, M2); submedial false vein (sf); cubitus, anterior
branches (Cu,, Cu2), and Anals (An,, An2). Fig. 2. Tarsomeres of E. taeniatifrons , showing: pointed calcipala (c), on first
tarsomere; and shallow pedisulcus (p), on second tarsomere. Fig. 3. Tarsomeres of Simulium duplex Shewed & Fredeen,
showing: rounded calcipala (c), and deep pedisulcus (p). Fig. 4. Terminalia of male Metacnephia saskatchewana Shewed
& Fredeen: parameral spines (p); ventral plate (v); ventral plate arm (a); gonocoxite (gc); gonostylus (gs); and terminal
spine (s).
Black flies (Simuliidae) of the Saskatchewan River
195
Quaest. Ent., 1981, 17(3,4)
196
Fredeen
V
Fig. 5. Male of Simulium duplex Shewell & Fredeen, left lateral aspect: mesepimeral tuft (t); scutum (s); scutellum (1);
postnotum (p); gonostylus of terminalia (d).
Figs. 6 and 7. Head, female, dorsal aspect, showing: antenna (a); eye (e); and frons (f), of : 6, Ectemnia taeniatifrons\ and
7, Metacnephia saskatchewana Shewell & Fredeen.
Figs. 8-1 1. Third leg of female Simulium : claw (cl); pedisulcus on second tarsomere (p); calcipala (ca); first tarsomere (t);
tibia (ti); femur (f); trochanter (tr); and coxa (c), of : 8. S. meridional Riley; 9, S. vittatum Zetterstedt; 10, S. luggeri
Nicholson & Mickel; and 1 1, S', arcticum Malloch.
Figs. 12 and 13. Terminalia of female Simulium , ventral view: cercus (c); anal lobe (a); hypogynial valve (h); and genital
fork (g), of : 12, S. luggeri ; and 13, S. arcticum.
Black flies (Simuliidae) of the Saskatchewan River
197
!
Quaest. Ent., 1981, 17 (3,4)
198
Fredeen
7
8 (7)
8'
9 (70
9'
10 (90
10'
11 (ioo
rr
12 (110
12'
13 (120
13'
14 (13')
14'
Greyish or brownish flies without contrastingly dark postnotum 9
Scutum with seven alternating stripes of yellowish-grey and brownish orange .
Simulium bivittatum Malloch
Scutum yellowish-grey without distinct striped pattern
Simulium griseum Coquillett
Scutum distinctly striped; color patterns of third leg segments as in Fig. 9 ...
Simulium vittatum Zetterstedt
Scutum not distinctly striped (indistinct vittae may be seen at certain angles of
view) 10
Subcosta without setae (rarely one or two) on ventral surface; terminalia with
hypogynial valves widely separated and not obscuring genital fork (Fig. 12);
color pattern of third leg as in Fig. 10
Simulium luggeri Nicholson and Mickel
Subcosta with row of setae on ventral surface (sockets visible if setae lost);
terminalia with hypogynial valves almost meeting medially, thus obscuring
genital fork (Fig. 13) 11
Hind tibia with distal half dark and proximal half pale (Fig. 11); claw with
minute subbasal tooth Simulium arcticum Malloch
Hind tibia dark except slanted “knee patch” (Fig. 10); claw without minute
subbasal tooth 12
Frons and terminal abdominal terga thinly grey pollinose; anal lobe twice as
long as cercus, almost square Simulium decorum Walker
Frons and terminal abdominal terga shiny black or brown; anal lobe scarcely
longer than cercus, wider than long 13
Fore coxa, trochanter and entire femora as dark as first tarsomere; setae on
stem vein and base of costa dark; entire antenna dark; a small dark species
(wing length often under 2.0 mm) Simulium tuberosum (Lundstroem)
Fore coxa, trochanter and proximal portion of femur paler than first tarsomere;
setae on stem vein and base of costa pale; pedicel of antenna relatively pale; size
various; abdomen pale ventrally 14
Medial margins of hypogynial valves straight and slightly divergent distally;
anterior margin of anal lobe not noticeably more sclerotized than rest of lobe;
width and length of dorsal sclerites on third and fifth abdominal segments
approximately equal Simulium venustum Say
Medial margins of hypogynial valves concave with oval space between them;
anterior margin of anal lobe noticeably more sclerotized than rest of lobe; dorsal
sclerites on third and fifth abdominal segments approximately two times wider
than long Simulium verecundum Stone and Jamnback
PUPAE
1 Cocoon loosely woven throughout 2
1' Cocoon tightly woven, at least in posterior portion 3
2 (1) Cocoon sock-shaped, attached or not to short stalk; ten filaments in respiratory
organ (Fig. 14) Ectemnia taeniatifrons (Enderlein)
2' Cocoon boot-shaped (elavated anterior collar), not attached to stalk; 17 to 19
Black flies (Simuliidae) of the Saskatchewan River
199
3 (10
3'
4 (3)
4'
5 (30
5'
6 (5)
6'
1 (60
r
8 (50
8'
9 (8)
9'
10 (9')
10'
11 (80
11'
12 (11)
12'
13 (110
13'
filaments in respiratory tuft, irregularly branched (Fig. 15)
Metacnephia saskatchewana Shewed and Fredeen
Pupal respiratory organ with four or fewer filaments 4
Pupal respiratory organ with more than four filaments 5
Pupal respiratory organ of two (rarely three or four) stout filaments (Fig. 16)
Simulium duplex Shewed and Fredeen
Pupal respiratory organ of four slender filaments (Fig. 17)
Simulium euryadminiculum Davies
Pupal respiratory organ of six filaments . 6
Pupal respiratory organ of more than six filaments 8
Pupa and cocoon relatively small (about 3.0 mm long) and dark; respiratory
filaments less than half as long as cocoon (Fig. 18)
Simulium tuberosum (Lundstroem)
Pupa and cocoon 4.0 mm or longer, pale; respiratory filaments more than half as
long as cocoon 7
Respiratory filaments relatively fine, spread about 90 degrees at base (Fig. 19)
Simulium venustum Say
Respiratory filaments relatively coarse, spread up to 180 degrees at base (Fig.
20) Simulium verecundum Stone and Jamnback
Respiratory organ with eight filaments 9
Respiratory organ with more than eight filaments 11
Respiratory filaments grouped as pairs on four long petioles (Fig. 21)
Simulium rugglesi Nicholson and Mickel
Respiratory filaments not paired 10
Cocoon more than four mm long, loosely woven (Fig. 22) . Simulium decorum
Walker
Cocoon about three mm long, tightly woven (Fig. 23)
Simulium bivittatum Malloch and Simulium griseum Coquillett1
Pupal respiratory organ with 12 filaments 12
Pupal respiratory organ with more thar 12 filaments 13
Filaments in respiratory organ paired; cocoon boot-shaped (with an elevated
anterior collar) (Fig. 24) Simulium arcticum Malloch
Filaments in respiratory organ in groups of three; cocoon generally
slipper-shaped (without anterior collar) (Fig. 25)
Simulium luggeri Nicholson and Mickel
Pupal respiratory organ with 16 filaments, in pairs; cocoon slipper-shaped (Fig.
26) Simulium vittatum Zetterstedt
Pupal respiratory organ with 22 to 26 irregularly branched filaments; many
cocoons boot-shaped (Fig. 27) Simulium meridionale Riley
lSimulium bivittatum Malloch and S. griseum Coquillett may be reliably separated as adults.
Quaest. Ent., 1981, 17 (3,4)
200
Fredeen
Figs. 14-20. Cocoon, and one set of respiratory filaments of pupa, of: 14, Ectemnia taeniatifrons (Enderlein); 15,
Metacnephia saskatchewana Shewed & Fredeen; 16, Simulium duplex Shewed & Fredeen; 17, S. euryadminiculum
Davies; 18, S. Tuberosum (Lundstroem); 19, S. venustum Say; 20, S. verecundum Stone & Jamnback.
Black flies (Simuliidae) of the Saskatchewan River
201
Figs. 21-27. Cocoon and one set of respiratory filaments of pupa, of: 21, Simulium rugglesi Nicholson & Mickel; 22, S.
decorum Walker; 23, S. griseum Coquillett, and S. bivittatum Malloch; 24, S. arcticum Malloch; 25, S. luggeri Nicholson
& Mickel; 26, S. vittatum Zetterstedt; and 27, S. meridionale Riley.
Quaest. Ent., 1981, 17 (3,4)
202
Fredeen
LARVAE
1
F
2 (1)
2'
3 (2')
3'
4 (30
4'
5 (20
5'
6 (5)
6'
7 (6)
T
8 (60
8'
9 (80
9'
10 (50
10'
11 (10)
IF
Three unbranched anal gills (Fig. 28) 2
Three compound anal gills 5
Lateral margins of abdominal segments 5 to 8 extended ventrally to about twice
depth of segments 1 to 4 (Fig. 28); prominent pair of ventral tubercles (Fig. 28);
ventral view of head capsule (Fig. 29) Ectemnia taeniatifrons (Enderlein)
Lateral margins of abdominal segments 5 to 8 not extended ventrally; anal
sclerites darkly sclerotized (Fig. 30); ventral tubercles insignificant 3
Arms of anal sclerite as in Fig. 31; postgenal cleft extended to base of
hypostomium (Fig. 32) Metacnephia saskatchewana Shewed and Fredeen
Arms of anal sclerite as in Fig. 33; postgenal cleft extended less than half
distance to base of hypostomium (Fig. 34) 4
Suboesophageal ganglion dark (Fig. 34); larva relatively large ( > 7 mm long
when mature) and with prominent dark, greenish transverse bands
Simulium vittatum Zetterstedt
Suboesophageal ganglion almost invisible (Fig. 35); larva relatively small (< 5
mm long when mature) and without prominent dark transverse bands
Simulium bivittatum Malloch and Simulium griseum Coquillett2
Eighth abdominal segment with two prominent ventral tubercles; abdominal
integument sparsely clothed in fine short bristles 6
Eighth abdominal segment without prominent ventral tubercles; abdominal
integument essentially bare 10
Postgenal cleft extended to base of hypostomium (Figs. 36, 37) 7
Postgenal cl€ft not extended to base of hypostomium (Fig. 38) 8
Prominent row of bristly tubercles in transverse reddish band encircling each of
segments 1 to 5, smaller tubercles on remaining segments; suboesophageal gland
colorless Simulium meridionale Riley
Abdomen without bristly tubercles; suboesophageal gland dark
Simulium rugglesi Nicholson and Mickel
Ventral tubercles broadly rounded; postgenal cleft broadly rounded, extended
about three quarters of distance to base of hypostomium (Fig. 38)
Simulium luggeri Nicholson and Mickel
Ventral tubercles distinctly conical; postgenal cleft extended no more than half
distance to base of hypostomium 9
Postgenal cleft extended about one-third of distance to base of hypostomium,
bilobed (Fig. 39) Simulium euryadminiculum Davies
Postgenal cleft extended about one-half of distance to base of hypostomium, not
bilobed (Fig. 40) Simulium duplex Shewed and Fredeen
Medial spots on dorsal surface of head capsule dark 11
Medial spots on dorsal surface of head capsule pale 12
Suboesophageal ganglion dark; postgenal cleft extended about half way to base
of hypostomium (Fig. 41) Simulium tuberosum (Lundstroem)
Suboesophageal ganglion almost colorless; postgenal cleft extended about
three-quarters of distance to base of hypostomium (Fig. 42)
S. bivittatum and S. griseum may be reliably separated as adults only.
Black flies (Simuliidae) of the Saskatchewan River
203
Fig. 28. Lateral view of posterior portion of abdomen of Ectemnia taeniatifrons (Enderlein) larva: anal gills; g); posterior
ring of hooklets (r); ventral tubercle (t). Not drawn to scale. Fig. 29. Ventral view of head Capsule of E. taeniatifrons
larva: hypostomium (h); suboesophageal ganglion (g); margin of postgenal cleft (p). Fig. 30. Lateral view of Metacnephia
saskatchewana Shewed & Fredeen larva: anal gills (g); portion of posterior ring of hooklets (r). Not drawn to scale. Fig.
31. Dorsal view of anal sclerites of M. saskatchewana larva: anal sclerite (s); portion of posterior ring of hooklets (r). Not
drawn to scale. Fig. 32. Ventral view of head capsule of M. saskatchewana larva: postgenal cleft (p). Fig. 33. Dorsal view
of anal sclerites of Simulium vittatum Zetterstedt larva: anal sclerite (s); portion of posterior ring of hooklets (r). Not
drawn to scale. Figs. 34-45. Ventral view of head capsule of Simulium larvae: suboesophageal ganglion (g); margin of
postgenal cleft (p), of: 34, S. vittatum Zetterstedt; 35, S. griseum Coquillett and S. bivittatum Malloch; 36, S.
meridionale Riley; 37, 5. rugglesi Nicholson & Mickel; 38, S. luggeri Nicholson & Mickel; 39, S. euryadminiculum
Davies; 40, 5. duplex Shewed & Fredeen; 41, S. tuberosum (Lundstroem); 42, S. articum Malloch; 43, S. decorum
Walker; 44, S', venustum Say; 45, S. verecundum Stone & Jamnback. (Scale applies to head capsules, only).
Quaest. Ent., 1981, 17 (3,4)
204
Fredeen
Simulium arcticum Malloch
12 (100 Mature larvae large (many 7 mm long); antenna extended as far as distal end of
stalk of cephalic fan; postgenal cleft slightly longer than wide (Fig. 43)
Simulium decorum Walker
12' Mature larvae smaller (many 6 to 6.5 mm long); tip of antenna extended beyond
distal end of stalk of cephalic fan; postgenal cleft about as long as wide (Figs.
44,45)
. . Simulium venustum Say and Simulium verecundum Stone and Jamnback3
NOTES ABOUT SPECIES
1. Ectemnia taeniatifrons (Enderlein) (Figs. 1, 2, 6, 14, 28, 29)
Thirty years ago larvae and pupae of this species were widely distributed throughout the
north, south, and main branches of the Saskatchewan River in Saskatchewan; none were found
in tributaries. In recent years fewer specimens have been collected and those mainly near the
confluence of the north and south branches near Prince Albert, the Red Deer River near
Estuary and the South Saskatchewan River near Pike Lake.
Ectemnia taeniatifrons is univoltine. Eggs laid in April and May hatch in late summer. At
fall freeze-up, some larvae are almost half grown. Larvae are notable in that many are found
attached to the tips of slender stalks up to 15 mm long, built from threads of salivary secretions
and debris and attached with broad bases to substrates. If undisturbed, a larva attaches its
cocoon on or near the tip of its stalk. Larvae continue growth under the ice and adults
commence emerging three to five weeks after ice break-up. Adults commonly feed on sap
seeping from the bruised bark of birch trees. Mating occurs in flight in thinly dispersed swarms
in clearings and thereafter, the females seek blood from humans and other animals. Because of
its rarity the species was not considered to be economically important.
2. Metacnephia saskat chew ana Shewed and Fredeen (Figs. 4, 7, 15, 30-32)
Larvae and pupae occur regularly on water weeds collected in the spring from the Shell and
Battle Rivers in Saskatchewan and occasionally from the North Saskatchewan River as far as
about 50 km downstream from the outlets of those small tributaries. I believe that the species is
not native to the Saskatchewan River, but that larvae appear there only after accidentally
drifting in from tributaries. The cocoons possess the same shape as those of S. arcticum , but are
loosely constructed and the pupa possesses 17 to 19 respiratory filaments instead of 12.
Only one generation per year is known. Adults emerge four to eight weeks after ice break-up
suggesting that the species overwinters as larvae under the ice of permanently flowing rivers.
Feeding habits of adults are not known.
Type specimens for this species were collected from the Shell River (Shewed and Fredeen,
1958) and since then it has been shown to be widely distributed in western Canada.
*S. venustum and S. verecundum may be reliably separated in the pupal and adult stages only.
Black flies (Simuliidae) of the Saskatchewan River
205
3. Simulium (Eusimulium) euryadminiculum Davies (Figs. 17 and 39)
This species breeds abundantly in the Battle and Shell Rivers. Larvae occasionally drift into
the North Saskatchewan River and have been collected as far as about 60 km downstream from
those tributaries. There, adults are sometimes produced judging by empty pupal cases, but
there is no evidence of permanent establishment in the Saskatchewan River. S.
euryadminiculum is univoltine with adults emerging in May about three to five weeks after ice
break-up. Blood-fed females have been taken from a dead common loon (Davies et al., 1962)
and have driven chickens indoors in Saskatchewan, without causing losses. The most severe
attack on chickens occurred May 16 to 18, 1981, near Prongua, Saskatchewan.
4. Simulium ( Eusimulium ) duplex Shewed and Fredeen (Figs. 3, 5, 16 and 40).
Larvae and pupae of this species occur regularly in the Battle and Shell Rivers, the source of
type specimens (Shewed and Fredeen, 1958) and in small numbers in the Saskatchewan River
below their outlets. They were collected once from the Red Deer River immediately above its
confluence with the South Saskatchewan River in southwestern Saskatchewan. S. duplex is
univoltine, with pupation usually occurring in late May. Feeding habits of adults are not
known.
5. Simulium (Byssodon) meridionale Riley (Figs. 8, 27 and 36).
S. meridionale is widely distributed throughout many rivers and streams in Saskatchewan.
Recently, larvae have become relatively abundant in the North Saskatchewan River, especially
in the vicinities of the Wingard and Cecil ferries. They also occur in the Red Deer and South
Saskatchewan Rivers in southwest Saskatchewan and occasionally downstream from Tobin
Lake on the main Saskatchewan.
Eggs overwinter in river bed sand and commence hatching within a month after ice
break-up in the spring. The species is multivoltine. Larval populations often peak in May and
June in the South Saskatchewan and in July in the north branch. It is sometimes abundant
enough to be a pest of humans and other animals. In some localities such as Melville,
Saskatchewan, it has been a pest of poultry and implicated indirectly in fatal proven
Leucocytozoon infections in flocks of domestic poultry. Those outbreaks originated in small
local streams not connected with the Saskatchewan River. Possibly sibling species of S.
meridionale possessing different host preferences are involved. Surprisingly, summer-long
emergences of relatively large numbers of S. meridionale in recent years from the North
Saskatchewan River did not result in poultry losses until early July 1981, when six farmers in a
100,000 ha area near Nipawin, Saskatchewan reported losses. Veterinarians considered those
fatalities due to exsanguination rather than disease. S. meridionale females were abundant
inside chicken house windows. Deaths occurred too rapidly to allow collection of blood smears.
Until more information is available poultry producers should avoid establishing new flocks
out-of-doors near the North Saskatchewan River. Swarms of S. meridionale have been
bothersome as far as 15 km from that river in recent years. Poultry producers already
established should report immediately any massive illnesses in their flocks to the Western
College of Veterinary Medicine at Saskatoon. Blood is required from living birds for diagnosis.
Quaest. Ent., 1981, 17 (3,4)
206
Fredeen
6. Simulium (Byssodon) rugglesi Nicholson and Mickel (Figs. 21 and 37)
Eggs, larvae and pupae of this species occur regularly and sometimes abundantly in small
clear rivers such as, the Battle, Shell, and Torch, and in small numbers in the North
Saskatchewan River below the outlets of those tributaries.
Eggs overwinter in river bed sand and commence hatching about one month after ice
break-up in spring. Larvae attain maximum abundance about mid-June, but may be found
until September, suggesting more than one generation per summer.
Shewed (1955) identified S. rugglesi as the species believed responsible for transmitting
fatal Leucocytozoon infections to ducklings and goslings in eastern Canada. However, there
have not been any reports of this black fly attacking poultry in Saskatchewan.
7. Simulium (Psilopelmiaj bivittatum Malloch and 8. S. (Psilopelmiaj griseum Coquillett
(Figs. 23 and 35)
I was not able to reliably separate immature stages of these two species. Adults reared from
pupae collected throughout the ice-free season in 1976 showed that the two species occurred
together that year in both the South Saskatchewan and Red Deer Rivers in southwest
Saskatchewan. In earlier years larvae and pupae of both species were found every summer in
the South Saskatchewan River. They were abundant in upper reaches, but rare in lower reaches
of that river and absent from the North Saskatchewan River. Populations in the lower South
Saskatchewan apparently were renewed annually by downstream drift of eggs and larvae from
breeding sites much further upstream because specimens have not been collected since
completion of a hydroelectric dam near Macrorie in 1968.
At Estuary, newly-hatched larvae appeared in May, a few weeks after ice break-up.
Development was rapid and by late May pupation commenced. Both species were multivoltine
and attained maximum abundance in July and August. Occasional severe outbreaks until about
1965 used to force people indoors at Saskatoon and Medicine Hat (Alberta). Adults of these
species also attacked horses, cattle, sheep and hogs, but apparently not poultry.
9. Simulium (Psilozia) vittatum Zetterstedt (Figs. 9, 26, 33, and 34).
Although S. vittatum is the most widely distributed black fly species in Saskatchewan it
apparently did not become established in the Saskatchewan River until the 1960’s, coincidental
with appearance of several species of water weeds. Now immature stages occur throughout this
river system and are particularly abundant in beds of weeds that provide attachment sites for
larvae and pupae, and substrates suitable for egg masses on emergent leaves.
S. vittatum overwinters as eggs in the river bed, and occasionally as larvae. It is univoltine
and attains greatest abundance in late July and early August. As a pest it is not considered too
bothersome, even when adults are abundant. These are relatively unobstrusive in attacks,
quietly entering the ears of animals where they feed undisturbed.
10. Simulium (Gnus) arcticum Malloch (Figs. 11, 13, 24, and 42)
S. arcticum was considered to be the only black fly species capable of killing livestock in
Saskatchewan until recently when S. luggeri became the dominant species in the Saskatchewan
River. Larvae and pupae used to occur throughout the entire Saskatchewan River in
Saskatchewan, but were particularly abundant in rocky rapids in the final 1 50 km or so of each
of the north and south branches and in the entire main river below their confluence. The most
destructive outbreaks occurred from 1944 to 1947, inclusive, when more than 1,100 farm
Black flies (Simuliidae) of the Saskatchewan River
207
animals, mainly cattle, were killed, including many expensive herd sires. Up to 3 million ha of
farm lands in Saskatchewan were affected at irregular intervals by these outbreaks until advent
of chemical larviciding in 1948 (Fredeen, 1958). Deaths were caused by direct toxemia. Native
born mature mammals were relatively immune to the toxins, but outbreaks still caused indirect
losses by interrupting grazing and breeding. Most outbreaks occurred in late May or early
June, rarely in August.
Since the early 1960’s numbers of larvae in these rivers gradually declined until this species
no longer threatens livestock. The last outbreak in which fatalities occurred was in 1972 near
Wingard. This decline seems not to have been caused by larviciding, as that affected only 150
km or shorter sections of these rivers, reinfested annually from untreated sections upstream.
Rather, the decline coincided with a combination of several ecological changes including
increases in amounts of dissolved plant nutrients, shallowing and clearing of water due to
summertime impoundment behind new hydroelectric dams, and development of massive beds of
aquatic plants. Reduced numbers of larvae of S. arcticum continue to inhabit rapids, however,
and the species might return in bothersome numbers if eutrophication declines. S. arcticum
overwinters as eggs in submerged river sand. First-instar larvae begin to appear 10 to 14 days
after ice break-up in the spring and pupation commences four or more weeks later. A second,
smaller generation often occurs in July or August.
Downstream drift of larvae undoubtedly played an important part in development of
outbreaks. When river levels declined as they usually did in May and early June, larvae
released and drifted downriver to more suitable sites in rocky rapids. It was from such
accumulation points that many destructive outbreaks originated. Accumulation of unfed
females awaiting weather conditions suitable for flight also was a factor. There was evidence,
however, that some females could lay the first batch of eggs autogenously, that is, from
nutrients stored during larval life (Fredeen, 1963).
Mammals within about 30 km of rapids were perhaps in greatest danger of attack. In 1947,
mammals were killed near Simpson, Saskatchewan, some 200 km south of the nearest point of
origin of black fly populations.
Three morphologically different forms of S. arcticum have been collected in Saskatchewan.
The pest species from the Saskatchewan River was identified as S. arcticum by Stone (1959)
after comparing it with Malloch’s types. A second relatively small, pale variety is widespread in
northern Saskatchewan and a third large, dark variety is in the Cypress Hills in southwest
Saskatchewan.
11. Simulium ( Phosterodoros ) luggeri Nicholson and Mickel (Figs. 10, 12, 25, and 38).
S. luggeri is the most northerly distributed species of Phosterodoros , and is the only one
known to occur in western Canada. Until the late 1960’s breeding of S. luggeri Nicholson and
Mickel in Saskatchewan was restricted to medium-sized clear rivers such as the Battle, Shell
and Torch Rivers where there was vegetation for attachment by larvae. From these rivers
larvae drifted in small numbers out into the North Saskatchewan River. However, they failed
to colonize either branch of the Saskatchewan until perhaps 1968 when larvae were detected
for the first time in the south branch near Fenton. By August, 1971, it was breeding so
abundantly in that river that emerging swarms of adults stampeded cattle in a pasture near St.
Louis. Also, for the first time in 1971 there was a distinct increase in numbers of larvae found
attached to artificial substrates anchored in the north branch near Prince Albert, indicating
that a breeding population had become established there. In 1976, larvae exceeded those of S.
Quaest. Ent., 1981, 17 (3,4)
208
Fredeen
arcticum in abundance and for the first time outbreaks of economic proportions originated
repeatedly from that river. Since then, outbreaks have continued to severely affect people and
livestock in some 18,000 km2 of farmlands and recreational areas in east central Saskatchewan.
Even though S. luggeri does not poison animals as severely as does S. arcticum , the habit of
swarming densely around heads of cattle causes stampeding. Grazing and breeding are
repeatedly interrupted throughout the summer and fences are broken down. Young animals
suffer broken bones and increased incidence of stress-related diseases such as pneumonia and
diarrhoea. Summer-long outbreaks reached a peak in 1978, causing losses including unrealized
weight gains that were estimated to have exceeded $1,400,000.00 Fredeen (in preparation).
Relatively intensive larviciding prevented re-occurrences of these outbreaks in 1979.
S. luggeri is capable of about five generations per summer in this region. Like S. arcticum it
overwinters as eggs in submerged sand. Hatching commences about two weeks after ice
break-up in the spring and adults emerge in late May or early June. Attacks on people and
livestock continue during warm days until late in the fall. Egg masses are attached to emergent
vegetation and also, single eggs, similar to those of S. arcticum may be dropped onto the water
surface .
12. Simulium (sensu stricto ) decorum Walker (Figs. 22, and 43).
This species is distributed sparsely throughout the province. Larvae occur most frequently in
the outfalls from small stream impoundments. Single larvae sometimes collected from the
North Saskatchewan River presumably originated in small streams. Not enough is known
about S. decorum to predict whether it could commence breeding in fast flowing sections of the
Saskatchewan River. It overwinters as eggs in wet river sand and apparently is capable of
several generations per summer. The females attack a wide vareity of animals. In northern
Saskatchewan adults are sometimes abundant enough to create severe local problems for'
humans and other animals.
13. Simulium (sensu stricto) tuberosum (Lundstroem) (Figs. 18 and 41).
Landau (1962) showed that S. tuberosum (Lundstroem) in southern Ontario consists of a
complex of at least four cytogenetically distinct species. Several morphological forms occur in
Saskatchewan indicating a complex here, also.
Larvae occur sparsely but regularly in the North Saskatchewan River, especially
downstream from tributaries that are normal habitats. Larvae were reliably reported from
South Saskatchewan River for the first time in 1979 and again in 1980, when a few were
collected from three sites, 20, 40 and 120 km upstream from the confluence with the north
branch. This widespread distribution and lack of stream tributaries which might have
contributed larvae, suggests that the species is now breeding in that river.
S. tuberosum (Lundstroem) may be multivoltine and adults have been collected along with
those of S. venustum Say from swarms attacking humans and other animals. However,
nowhere in the southern half of the province has it been abundant enough to create problems by
itself.
14. Simulium (sensu stricto) venustum Say (Figs. 19, and 44).
S. venustum Say is one of the most widely distributed black flies in Saskatchewan and
breeds in a wide variety of relatively unpolluted streams and rivers. Occasionally faster flowing
sections of small rivers such as the Assioniboine, Big and Beaver have produced brief outbreaks
Black flies (Simuliidae) of the Saskatchewan River
209
that have created problems for humans and other animals. In the early 1970’s S. venustum Say
commenced breeding sparsely in the last 50 km or more of both branches of the Saskatchewan
River. Although still uncommon there it should be watched because of the large productive
capacity of the Saskatchewan River.
S. venustum Say overwinters as eggs and is multivoltine, attaining greatest abundance in
June. Rothfels (1975) considers this to be a species complex.
15. Simulium (sensu stricto ) verecundum Stone and Jamnback (Figs. 20, and 45).
Larvae and pupae of this species occasionally have been colleted from the last 50 km or so of
each branch of the Saskatchewan River. Although it may become more abundant it probably
will not become economically important because it is not known to attack for blood.
PRACTICAL CONSIDERATIONS
The black fly fauna of the Saskatchewan River in Saskatchewan has changed considerably
during the past 15 years coinciding with changes in river conditions. Originally S. arcticum was
the dominant species. Sporadic outbreaks that killed livestock belonging to the earliest settlers
in 1885 were attributed to this species. As recently as 1972 an outbreak of S. arcticum
originating from an untreated portion of the North Saskatchewan River killed 18 farm animals
near Wingard. Since then, however, S. luggeri has gradually replaced S. arcticum as the
dominant species and the conversion is believed due to changes in river conditions that allowed
growth of large beds of aquatic plants on previously barren river beds. Losses estimated to have
exceeded $1,400,000.00 resulted from summer-long outbreaks of S. luggeri in 1978 (Fredeen,
in preparation).
Tendencies towards continuing changes in black fly populations were indicated by
appearances of larvae of S. tuberosum (Lundstroem) and S. venustum Say in several sites in
1979 and 1980. Troublesome outbreaks of those two blood sucking species have been reported
from several small prairie rivers. Thus their populations should be monitored annually along
with those of blood sucking species in the relatively large Saskatchewan River.
At present, chemical larviciding offers the only means of preventing outbreaks (Fredeen,
1977). Continued research may eventually provide other methods. Residents in areas recently
affected by outbreaks of S. arcticum Malloch and S. luggeri Nicholson and Mickel require
assurance that protection will be provided when required. It is hoped that this key to species
inhabiting the Saskatchewan River will prove useful for monitoring and research.
ACKNOWLEDGEMENTS
I am indebted to Dr. L. Burgess, Canada Agriculture Research Station, Saskatoon,
Saskatchewan, for reviewing this paper and Ralph Underwood for advice during preparation of
the figures.
REFERENCES
Davies, L. 1968. A key to the British species of Simuliidae (Diptera) in the larval, pupal and
adult stages. Freshwater Biological Association, Westmorland, England, scientific
publication number 24. 126 pp.
Davies, D.M., B.V. Peterson and D.M. Wood. 1962. The black flies (Diptera: Simuliidae) of
Quaest. Ent., 1981, 17(3,4)
210
Fredeen
Ontario. Part I. Adult identification and distribution with descriptions of six new species.
Proceedings of the Entomological Society of Ontario (1961). 92: 71-154.
Fredeen, F.J.H. 1958. Black flies (Diptera: Simuliidae) of the agricultural areas of Manitoba,
Saskatchewan and Alberta. Proceedings, Tenth International Congress of Entomology 3:
819-823.
Fredeen, F.J.H. 1963. Oviposition in relation to the accumulation of blood thirsty black flies
(Simulium (Gnus) arcticum Mall (Diptera)) prior to a damaging outbreak. Nature
(London) 200: 4910. p. 1024.
Fredeen, F.J.H. 1977. Black fly control and environmental quality with reference to chemical
larviciding in western Canada. Quaestiones Entomologicae 13: 321-325.
Fredeen, F. J.H. In preparation. Outbreaks of Simulium luggeri in Saskatchewan.
Landau, R. 1962. Four forms of Simulium tuberosum (Lundstr.) in southern Ontario: a
salivary gland chromosome study. Canadian Journal of Zoology 40: 921-939.
McAlpine, J.F., B.V. Peterson, G.E. Shewed, H.J. Teskey, J.R. Vockeroth and D.M. Wood.
1981. Manual of Nearctic Diptera. Volume 1, Monograph No. 27, Research Branch,
Agriculture Canada, Ottawa, Canada, 674 pp.
Rothfels, K.H. 1975. Personal communication. Department of Botany, University of Toronto,
Canada.
Shewed, G.E. 1955. Identity of the black fly that attacks ducklings and goslings in Canada
(Diptera: Simuliidae). The Canadian Entomologist 87: 345-349.
Shewed, G.E. and F.J.H. Fredeen. 1958. Two new black flies from Saskatchewan (Diptera:
Simuliidae). The Canadian Entomologist 90: 733-738.
Stone, A. 1959. Personal communication. United States Department of Agriculture,
Entomology Research Division, Washington, D.C.
Stone, A., C.W. Sabrosky, W.W. Worth, R.H. Foote and J.R. Coulson. 1965. A catalogue of
the Diptera of America north of Mexico, Agruculture Handbook No. 276, Agricultural
Research Service, United States of America, Washington, D.C. 1696 pp.
Stone, A. and E.L. Snoddy. 1969. The black flies of Alabama (Diptera: Simuliidae). Bulletin
390. Agricultural Experiment Station, Auburn University, Alabama. 93 pp.
Wood, D.M., B.V. Peterson, D.M. Davies and Helen Gyorkos. 1963. The black flies (Diptera:
Simuliidae) of Ontario. Part II. Larval identification, with descriptions and illustrations.
Proceedings of the Entomological Society of Ontario (1982) 19: 99-129.
SEASONAL CAPTURES AND DIVERSITY OF GROUND BEETLES (COLEOPTERA:
CARABIDAE) IN A WHEAT FIELD AND ITS GRASSY BORDERS IN CENTRAL
SASKATCHEWAN4
J.F. Doane
Research Station, Research Branch
Agriculture Canada
107 Science Crescent
Saskatoon, Saskatchewan, S7N 0X2
Canada
Quaestiones Entomologicae
17:211-233 1981
ABSTRACT
Pitfall trapping, at weekly intervals from spring to autumn during a 2-year period in a
wheat field and its grassy borders near Clavet, Saskatchewan, captured 87 species of carabids
in 25 genera. Seven percent of these species were captured only in the field, 30% only in the
field border and 63 % in both habitats. Bembidion obscurellum Mtsch., Bembidion
quadrimaculatum oppositum Say, Bembidion nitidum Kby., Amara lacustris Lee., Amara
carinata Lee., Amara farcta Lee., Harpalus herbivagus Say and Pterostichus lucublandus Say
were dominant in both habitats, while Bembidion timidum Lee. was dominant only in the field,
and Amara quenseli Schnh. and Amara obesa Say, were dominant only in the border. The
overall sex ratio in the four collections was about 1:1. However, data for some species differed
considerably from this ratio.
Species diversity and evenness were higher for the field border collections than for
collections from the field. An average of about 15 species per week was trapped in the border
compared to an average of 11 species per week in the field.
RESUME
Le Piegeage em fosses-collectionneuses dans un champs de ble sis pre de Clavet, Saskatchewan, et dans les bordures
herbeuses de de celui-ci, aux ecartements d’une semaine a partir du printemps jusqu’ a partir du printemps jusqu’ a /’
automne pendant deux ans, a pris 87 especes de carabides. reparties en 25 genres. En proportion, les especes prises ont ete
comme suit: seulement dans le champs, 7 p.c.; seulement dans les bordures, 30 p.c.; et a la fois dans le champs et dans les
bordures, 63 p.c. Bembidion obscurellum Mtsch., Bembidion quadrimaculatum oppositum Say, Bembidion nitidum Kby.,
Amara lacustris Lee., Amara carinata Lee., Amara farcta Lee., Harpalus herbivagus Say et Pterostichus lucublandus Say
etaient predominates parmi les especes qui se sont trouvees dans les deux habitats, tandis que Bembidion timidum Lee.
a predomine parmi celles qui se sont rencontrees seulement dans le champs et Amara quenseli Schnh. et Amara obesa
Say ont predomine parmi celles qui ont demeurees dans les bordures. La proportion sexuelle globale de quatre
collections a ete approximativement en rapport de 1:1 , cependant, cette proportion a souvent varie considerablement de ce
niveau dans le cas des especes particulieres.
La diversite et iuniformite des especes ont ete plus elevees dans les collections prises des bordures que dans celles des
champs. Une moyenne de 15 especes par semaine a ete captee dans les bordures a comparer dll especes par semaine en
moyenne dans le champs.
Contribution No. 793, from the Research Station, Saskatoon.
212
Doane
INTRODUCTION
Investigations of the way of life of carabids associated with agricultural lands have usually
been conducted to determine the potential importance of dominant species as predators and
their possible role in the population dynamics of crop pests (Rivard 1964; Frank 1971a; Kirk
1971a; Esau and Peters 1975; Finlayson and Campbell 1 976% Holliday and Hagley 1978). It is
also important to determine species composition, population structure and diversity of carabid
populations associated with various habitats, including arable lands. This information is
potentially useful for assessing impact of agricultural or other practices on stability of carabid
populations. Temporal changes or spatial differences in diversity could be a valuable biological
index for assessing effect of agronomic practices on carabid populations (Dritschilo and
Wanner 1980).
Carabids are known to attack wireworm larvae (Fox and MacLellan 1956) and are potential
predators of all stages of wireworms. This paper reports on pitfall trapping for carabids in a
wireworm infested field in Central Saskatchewan to determine: (1) dominant species that
might be important as predators of wireworms; (2) the species composition and diversity of
collections. For comparison, similar collections were made in the uncultivated field border
which also harboured a wireworm population. Observations on the sex ratio and seasonal
occurrence of some species are also included.
METHODS
The general methods, plot arrangement, plot size, and vegetation surrounding the study field
(52° 02' N., 106° 24' W.) near Clavet, Saskatchewan, were as described previously (Doane and
Dondale 1979). The field study, cultivated and seeded to spring wheat in June in 1975 and
1976, was harvested each year in late August and early September.
A pitfall trap was placed at the centre of each of 16 field plots and 15 field border plots.
Carabids were trapped in these two habitats for a 48-hr period each week, usually from
Monday until Wednesday, from April or May until mid-October, giving a total of 44 and 56
trap-days in 1975 and 1976 respectively. During the remainder of the week, traps remained in
place, but carabids captured were released in order to avoid possible over trapping of some
species. Sex of all carabids trapped was determined, and for species that comprised one per cent
or more of the total annual collection from each habitat, chi-square analysis was used to test
conformity to a 1:1 male-female ratio.
Abundance of different species within collections was expressed as a percentage of the total
collection for each year and habitat. To allow better comparison between years and habitats, in
which different numbers of trap-days and traps were used respectively, total seasonal catch per
species was also expressed as captures per metre of trap circumference per trap-day, since trap
circumference is one factor influencing number of captures (Turner 1962; Thomas and Sleeper
1977). The inside rim of the pitfall trap used in the present studies, described previously
(Doane 1961), had a circumference of 51.496 cm giving a total trapping circumference of 8.24
m (16 traps) and 7.72 m (15 traps) in the field and field border respectively.
Diversity and evenness of carabid collections made for the two years and sites was
determined using the Brillouin diversity index (H) and Simpson’s index of diversity (D) (Pielou
1969). Choice of the Brillouin index has been discussed previously for similar spider collections
(Doane and Dondale 1979). The Brillouin index was calculated using a table compiled by
Lloyd et al. (1968), while maximum diversity (Hmax) and evenness was determined by using
Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
213
equations given by Pielou (1969). Simpson’s measure of diversity (D) (Pielou 1969) was used
for comparison since it is worthwhile to have more than one measure of diversity.
RESULTS AND DISCUSSION
Abundance and seasonal diversity
Numbers of genera and species captured in each habitat for each year, were quite consistent
(Table 1). More individuals were captured in 1976 in both the field and its border, but on the
basis of numbers trapped per metre of trap circumference per trap-day the greatest numbers
were trapped in 1975 in the field (Table 2). Eighty-seven species were found in the four
collections (Table 2); of these 7% were collected only from the field, 30% only from the field
border and, 63% from both habitats.
Dominant species, those that comprised 3% or more of the total individuals in at least one of
the collections (Table 2), were the following: Bembidion obscurellum Mtsch., Bembidion
quadrimaculatum oppositum Say, Bembidion nitidum Kby., Bembidion timidum Lee., Amara
lacustris Lee., Amara carinata Lee., Amara fracta Lee., Amara quenseli Schnh., Amara obesa
Say, Harpalus herbivagus Say and Pterostichus lucublandus Say. Of these, B. timidum was
dominant only in the field and A. quenseli and A. obesa were dominant only in the border; the
remainder were dominant in both habitats.
For convenience, species abundance and seasonal occurrence are discussed under generic
groupings. Seasonal activity of dominant species that were trapped at Clavet, Saskatchewan,
agreed closely with that observed by Frank (1971a) for the same or congeneric species
occurring in Alberta.
Bembidion
Eighteen species were captured in the field and field border during the 2-year study period,
eight species in the field and 17 in the field border. B. obscurellum was by far the most
abundant species collected in the field in 1975, comprising about 40% of the total individuals in
the collection (Table 2), and about 85% of the individuals in that genus. Collection rate of
3.15/m trap circumference/trap-day was the greatest for any species during either year. In
1976, captures of B. q. oppositum were slightly in excess of B. obscurellum in the field. More
rare species of Bembidion were trapped in the field border than in the field.
Frank (1971a) captured 11 Bembidion species in a cultivated field at Calahoo, Alberta
(about 32 km northwest of Edmonton). There, B. q. oppositum was the most abundant species
followed by B. obscurellum , B. rupicola, B. nitidum and B. bimaculatum. Bembidion rupicola
was rarely trapped in Saskatchewan at Clavet.
At Clavet, Bembidion species were mainly active during mid-summer until about the first
week in August, when activity, as determined by pitfall captures, ceased abruptly. A small
amount of activity resumed in the late fall. Bembidion bimaculatum , collected in small
numbers at Clavet, was an exception in that it was active mainly from mid-summer on. The
main period of mating for the most abundant species B. obscurellum appeared to be in June,
assuming that the observed increased male activity during that period reflected mating activity.
Chi-square analysis showed that males were in excess of females (P<0.05) on June 4, June 18
and on July 2; on the latter date, 208 males and 1 13 females were captured. For the other 10
weekly collections so analyzed, the male-female ratio for B. obscurellum did not differ from a
Quaest. Ent. 1981,17(3,4)
214
Doane
1:1 expectation. Bembidion quadrimaculatum oppositum was the second most abundant
Bembidion species in the field collection in 1975, and in 1976 exceeded B. obscurellum in
numbers.
Amur a
The genus Amara was represented by 18 species in the two habitats; 15 were collected in the
field and 17 in the border. Amara lacustris and A. carinata were the most abundant species in
the field (Table 2). A. latior and A. fracta were slightly more abundant in the field than the
border while, A. quenseli and A. obesa were captured more often in the border. Species such as
A. apricaria and A. littoralis were captured regularly, but in low numbers, while others were
extremely rare (Table 2).
In Alberta, Frank (1971a) found many of the same species, but the most abundant in
Saskatchewan, A. lacustris , was rare at Calahoo, and A. carinata very abundant at Clavet, was
not found at the Alberta site. Amara torrida, common in Alberta, was extremely rare at
Clavet.
The seasonal activity peaks of most of the Amara species occurred in late summer and fall,
the same general pattern observed by Frank (1971a). However, there was some variation from
this predominant activity period exhibited particularly by A. fracta. In 1975, A. fracta was
active mainly from September to the close of the season, but in 1976, peak captures occurred in
April and again in midsummer, with relatively few captures in the fall.
Harpalus
Nine species of Harpalus were trapped, but of these, only H. herbivagus was abundant in
collections; it was taken regularly in both the field and border habitats. Lindroth (1968)
considered this species rare on the prairies where he considered Harpalus pleuriticus Kby..
more common. Frank (1971a) found H. pleuriticus second in abundance to H. amputatus but
did not capture H. herbivagus.
Harpalus herbivagus had a long period of activity during the season with several
pronounced peaks that varied somewhat in the two seasons. In 1975, this carabid was caught
throughout the early part of the season in low numbers, and after July, captures per week
steadily increased to a peak about the* third week in September. In 1976, however, it was
abundant in field traps from April to mid-July and was then captured in low numbers to the
end of the season. In the border, the same general pattern was observed, except that fewer
beetles were trapped, and the peak captures were observed about mid-June. Teneral H.
herbivagus adults were observed from mid-September until the end of the season.
Pterostichus
Pterostichus lucublandus was the most abundant of the five species trapped (Table 2). This
is an extremely widespread species described as transamerican (Lindroth 1966) and extending
south and east at least to North Carolina (Kirk 1971b). At Clavet it was found from early May
to October in both the field and field border. Peak activity occurred from about the end of May
until the first of July and then again from about the first week in August until late September.
In general, the peaks of activity agreed very closely with the pattern of activity described by
Kirk (1971b). Frank (1971a) concluded that this species was only active from May to July, but
noted a second peak one year. However, Kirk’s observations and those made at Clavet confirm
the existence of two activity peaks. The second peak is at least partially composed of young
adults, many of which were teneral in September at Clavet. Adults trapped in the fall may be a
mixture of newly emerged beetles and those of the previous generation (Kirk 1971b).
Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
215
Agonunt
Although five species of Agonum were trapped, few individuals per species were taken.
Frank (1971a) found Agonum cupreum Dej. and Agonum placidum Say, two of the same
species trapped at Clavet, to be quite common in Alberta.
Sex ratios
The male:female sex ratio for the 8741 carabids in the four collections, i.e., field and field
border 1975, and 1976, were 48:52, 48:52, 53:47 and 50:50% respectively. Rivard (1966), for a
collection of 10,000 carabids made over a 3-year period, observed a male:female ratio of 57:43
per cent. Although overall sex ratio of beetles trapped at Clavet was about 1:1, several species
did not conform to this ratio (Table 3). Usually males were in excess of females; exceptions
were A. carinata for which females were in excess of males in all four collections and P.
lucublandus for which females outnumbered males in three of the four collections (Table 3).
Disproportionate numbers of A. carinata males and females (overall about 3.5 times fewer
males than females) points to unknown characteristics in the life-cycle or activity that appear
to differ considerably from most of the species collected.
Diversity
Diversity and evenness were higher for the seasonal collections from the field in 1976 than in
1975 and, higher in the border collections in 1975 than in 1976 (Table 4). Both years, diversity
of the border collections was greater than that of the field collections. Although, total numbers
of species in the two habitats ranged from 45 to 66 (Table 1) during the two years, considerably
fewer species were captured in any one week (Table 5). More species per week were captured in
the border, but more individuals per week were trapped in the field.
Higher diversity of carabid species in the field border than in the field appears to be
consistent with the higher variety of plant and insect life in the uncultivated habitat. Lewis
(1969) observed that insect diversity was highest in hedgerows and that it declined in adjacent
crops as the distance from the hedgerow increased. Uncultivated lands therefore appear to be
important to the ecology of adjacent crops by providing niches and shelter both for beneficial
and pest species (Lewis 1965). Such areas may also provide alternate prey for predators when
pest species are not available as prey in the field (van Emden 1965). At Clavet, 63 per cent of
all carabid species were captured in both the field and its borders. Therefore, considerable
movement likely occurs between the two habitats. This suggests that the uncultivated border
may serve as shelter and a source of alternate prey for some species if the field becomes an
unsuitable habitat, as for example, during periods of summerfallowing.
Numbers of genera and species trapped each year in the field and its border (Table 1) and
the average numbers of species trapped in these habitats per week (Table 5) were remarkably
constant even for the field. Numbers of species in a habitat tend to reach an equilibrium
depending upon the habitat structure (MacArthur 1965). Thus, an approximately constant
number of species occur in the field and a few more in the field border as dictated by the
available resources and structure of the habitat.
Diversity indices of collections from the two habitats provide a convenient measure of
carabid species numbers and the relationship between the numbers of species and individuals
within species. Although these measures of diversity apply only to collections from the study
Quaest. Ent. 1981, 17 (3,4)
216
Doane
field and its borders, they will provide baseline information for studies in other fields and areas
of Saskatchewan when a similar collection method is used.
Potential of carabids as predators
Many species of carabids are primarily phytophagous (Johnson and Cameron 1969; Thiele
1977), so that food preferences of dominant species and their response to potential prey require
detailed study before the value of dominant species as predators can be assessed. Best and
Beegle (1977) investigated the feeding of five species of carabids on a variety of plant and
animal food in the laboratory and concluded that dead black cutworm larvae ( Agrotis ipsilon
(Hiifnagel)) were preferred by all the species tested. Frank (1971b) investigated the potential
of carabids from a cultivated field in Alberta as predators of eggs of the red-backed cutworm
( Euxoa ochrogaster Guenee) and reported that 21 species consumed eggs. These included
species of Bembidion (8), Amara (7), Agonum (2), Harpalus (2), P. lucublandus and
Trichocellus cognatus Gyll.. Serological tests showed that seven species fed on cutworm larvae
or pupae in the field. Natural predation by carabids on wireworm larvae in pastures has also
been demonstrated by the precipitin test (Fox and MacLellan 1956). Predatory species
included members of the genera Agonum , Amara , Harpalus and Pterostichus.
Perhaps the greatest chance for carabid predation in cultivated fields in Saskatchewan
would be upon wireworm eggs and newly hatched larvae. The main period of oviposition in the
field of the prairie grain wireworm, Ctenicera destructor (Brown), occurs from about the third
week in May until mid- June (Doane 1963). These beetles lay eggs under clods of soil in
summerfallow fields and in soil cracks (Doane 1967). Such eggs would be exposed to predation
when at least four species of Bembidion , i.e., B. obscurellum, B. q. oppositum , B. nitidum and
B. timidum are at peak populations and activity levels in the field. Bembidion species fed on
eggs of the red-backed cutworm (Frank 1971b) while, in Ontario, B. q. oppositum and B:
nitidum were shown to be predators of eggs of Hylemya brassicae (Bouche), the cabbage
maggot (Wishart, Doane and Maybee 1956). The role of Bembidion species as potential
predators of the egg and early larval stages of wireworms, therefore, is worthy of investigation.
ACKNOWLEDGEMENTS
Appreciation is expressed to George E. Ball, University of Alberta, for his kind assistance in
identification and to Drs. Ball and L. Burgess for helpful suggestions for improving the
manuscript. Thanks are extended also to R. K. Allan for valuable technical assistance, and to
summer students, Margaret McCrimmon and Rozanne Tennent, for much of the sorting and
pinning of collections.
REFERENCES
Best, L.R. and C.C. Beegle. 1977. Food preferences of five species of carabids commonly found
in Iowa cornfields. Environmental Entomology 6: 9-12.
Doane, J.F. 1961. Movement on the soil surface, of adult Ctenicera aeripennis destructor
(Brown) and Hypolithus bicolor Esch. (Coleoptera: Elateridae), as indicated by funnel
pitfall traps, with notes on captures of other arthropods. The Canadian Entomologist 93:
636-644.
Doane, J.F. 1963. Studies on ovipostion and fecundity of Ctenicera destructor (Brown)
(Coleoptera: Elateridae). The Canadian Entomologist 95: 1145-1 153.
Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
217
Doane, J.F. 1967. The influence of soil moisture and some soil physical factors on the
ovipositional behavior of the prairie grain wireworm, Ctenicera destructor. Entomologia
experimentalis et applicata 10: 275-286.
Doane, J.F. and C.D. Dondale. 1977. Seasonal captures of spiders (Araneae) in a wheat field
and its grassy borders in central Saskatchewan. The Canadian Entomologist 111:
439-445.
Dritschilo, W. and D. Wanner. 1980. Ground beetle abundance in organic and conventional
cornfields. Environmental Entomologist 9: 629-631.
van Emden, H.F. 1965. The role of uncultivated land in the biology of crop pests and beneficial
insects. Scientific Horticulture 17: 121-136.
Esau, K.L. and D.C. Peters. 1975. Carabidae collected in pitfall traps in Iowa cornfields,
fencerows, and prairies. Environmental Entomologist 4: 509-513.
Finlayson, D.G. and C.J. Campbell. 1976. Carabid and staphylinid beetles from agricultural
land in the lower Frazer Valley, British Columbia. Journal of the Entomological Society
of British Columbia 73: 10-20.
Fox, C.J.S. and C.R. MacLellan. 1956. Some Carabidae and Staphylinidae shown to feed on a
wireworm, Agriotes sputator (L.), by the precipitin test. The Canadian Entomologist 88:
228-231.
Frank, J.H. 1971a. Carabidae (Coleoptera) of an arable field in central Alberta. Quaestiones
Entomologicae 7: 237-252.
Frank, J.H. 1971b. Carabidae (Coleoptera) as predators of the redbacked cutworm
(Lepidoptera: Noctuidae) in central Alberta. The Canadian Entomologist 103:
1039-1044.
Holliday, N.J. and E.A.C. Haglev. 1978. Occurrence and activity of ground beetles
(Coleoptera: Carabidae) in a pest management apple orchard. The Canadian
Entomologist 110: 113-119.
Johnson, N.E. and R.S. Cameron. 1969. Phytophagous ground beetles. Annals of the
Entomological Society of America 62: 909-914.
Kirk, V.M. 1971a. Ground beetles in cropland in South Dakota. Annals of the Entomological
Society of America 64: 238-241.
Kirk, V. M. 1971b. Biological studies of a ground beetle Pterostichus lucublandus. Annals of
the Entomological Society of America 64: 540-544.
Lewis, T. 1965. The effect of shelter on the distribution of insect pests. Scientific Horticulture
17:74-84.
Lewis, T. 1969. The diversity of the insect fauna in a hedgerow and neighbouring fields.
Journal of Applied Ecology 6: 453-458.
Lindroth, C.H. 1966. The ground beetles of Canada and Alaska, 4. Opuscula Entomologica,
Supplementum 29, 240 pp.
Lindroth, C.H. 1968. The ground beetles of Canada and Alaska, 5. Opuscula Entomologica,
Supplementum 33, 295 pp.
Lloyd, M., J. H. Zar, and J.R. Karr. 1968. On the calculation of information theoretical
measures of diversity. American Midland Naturalist 79: 257-272.
MacArthur, R.H. 1965. Patterns of species diversity. Biological Review 40: 510-533.
Pielou, E.C. 1969. An introduction to mathematical ecology. Wiley-Interscience, New York.
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Belleville, Ontario. The Canadian Entomologist 96: 517-520.
Quaest. Ent. 1981, 17(3,4)
218
Doane
Rivard, I. 1966. Ground beetles (Coleoptera: Carabidae) in relation to agricultural crops. The
Canadian Entomologist 98: 189-195.
Thiele, H.U. 1977. Carabid beetles in their environments. Springer-Verlag, Berlin, New York.
Thomas, D.B. and E.L. Sleeper. 1977. The use of pitfall traps for estimating the abundance of
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Entomological Society of America 70: 242-248.
Turner, F.B. 1962. Some sampling characteristics of plants and arthropods of the Arizona
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Wishart, G., J.F. Doane, and G.E. Maybee. 1956. Notes on beetles as predators of eggs of
Hylemya brassicae (Bouche) (Diptera: Anthomyiidae). The Canadian Entomologist 88:
634-639.
Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
219
Table 1. Summary of individuals, species and genera of carabids captured in pitfall traps in a
wheat field and its borders near Clavet, Sask. from spring to autumn, 1975-76
aTotal captured in 44 and 56 days of trapping in 1975 and 1976 respectively.
b16 traps in the field and 15 traps in the field border.
Quaest. Ent. 1981, 17 (3,4)
Table 2. Species of carabids captured in pitfall traps expressed as number/meter trap circumference/trap day, and per cent of total captures, for
collections made in a wheat field and its borders in 1975 and 1976 at Clavet, Saskatchewan1
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan 221
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
225
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
227
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228
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(continued on next page)
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
229
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that comprised one percent or more of the collections made in a wheat field and its borders near Clavet, Sask.
230
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
231
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232
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Ground beetles (Coleoptera: Carabidae) in Central Saskatchewan
233
Table 5. Mean numbers of individuals and species of carabids captured in pitfall traps each
week in a wheat field and its borders near Clavet, Saskatchewan, 1975 and 1976.
234
ASPECTS OF THE COMPARATIVE ECOLOGY OF POPULATIONS OF FOUR
PATROBUS SPECIES (COLEOPTERA: CARABIDAE: PATROBINI) AT GEORGE LAKE,
ALBERTA
Alan Carter
Department of Soil Science
The University of British Columbia
Vancouver, Canada Quaestiones Entomologicae
V6T2A2 17:235-249 1981
ABSTRACT
Way of life and use of habitats by Patrobus stygicus Chaudoir, P. lecontei Chaudoir, P.
foveocollis Eschscholtz, and P. septentrionis Dejean were studied and compared with
previously hypothesized phylogenetic relationships of these species. Larvae and adults of P.
foveocollis were least hygrophilous and ranged from forest margins through sedge meadows to
Salix habitats adjoining the marshes. Larvae and adults ofP. stygicus were most hygrophilous
and ranged from forest margins through shallow marsh to the lakeside. Adults and larvae of
P. lecontei ranged from forest margins to shallow marsh. Most adults of P. septentrionis were
found in sedge meadows; their larvae were not recovered. Adults of P. lecontei were
characterized by rapid development of sexual maturity and a short life, whereas adults ofP.
stygicus had a longer prereproductive period and a longer life. Possible evolutionary
significance of the ecological characters specified above and their consequences are discussed
for the North American species o/Patrobus.
RESUME
La vie et I’usage de Vhabitat de Patrobus stygicus Chaudoir, P. lecontei Chaudoir, P. fovoecollis Eschscholtz, et P.
septentrionis Dejean ont ete etudies et compares quant aux relations phylogenetiques. Les larves et les adultes de P.
foveocollis etaient le moins hygrophiles et s’etendaient depuis des clairieres, a des Lisieres, des patis couverts de latches,
jusqu' aux habitats Salix contigus aux marecages. Les larves et les adultes de P. stygicus etaient le plus hygrophiles et
s’etendaient depuis des lisieres, a des marecages peu profonds jusqu’ aux bords d’un lac. Les Patrobus lecontei larves et
les adultes s’etendaient des lisieres aux marecages peu profonds. Les adultes de P. septentrionis se trouvaient dans la
plupart des cas dans des patis couverts de latches; leurs larves n’ont pas ete trouvees. Les adultes de P. lecontei etaient
caracterises par un developpement rapide de maturite sexuelle et d’une vie courte, tandis que les adultes de P. stygicus
avaient une periode adulte prereproductive plus longue et une vie plus longue. La signification evolutionnaire possible
des caracteristiques ecologiques mentionnees ci-dessus et leurs consequences ont ete traitees, en se limitant aux especes
Patrobus nord-americaines.
INTRODUCTION
Little is known of the natural history of the majority of North American carabid beetles.
However, the excellent taxonomic revision of Canadian, and Alaskan carabids by Lindroth
(1961-1969) and studies by Darlington (1938) and by Ball and associates (for example, Ball,
1966) have paved the way for field and laboratory investigations. Few North American workers
have studied the comparative ecology of closely related carabid species, for example from the
same subgenus, in an attempt to define ecological and behavioural characters which are useful
236
Carter
in taxonomy and phylogeny.
Populations of four Patrobus species ( P . stygicus, P. lecontei, P. foveocollis and P.
septentrionis ) which frequent wet boreal habitats are considered in this study. The objectives of
this study were to use ecological characters to test taxonomic and phylogenetic hypotheses
about closely related species and to compare their habitat use.
The following questions were asked:
(1) Are these Patrobus species separated in space during their period of main
activity?
(2) Are populations of different species, that occur in the same general habitats,
separated by seasonal activity?
(3) Does spatial distribution of these carabids vary according to time of year?
Life history features were considered as adaptations (Cole, 1954). Evolutionary
interpretation of these features and other ecological characters were tested against Darlington’s
(1938) hypotheses about relationships of these species, with the exception that P. fossifrons and
P. stygicus are regarded as distinct species rather than subspecies (see Lindroth, 1961).
THE FIELD STATION AND STUDY AREAS
The Field Station
The study area is at 53° 57'N and 114° 06'W at George Lake, about 64 km northwest of
Edmonton, Alberta. The area is at the southern margin of the boreal mixed forest subzone (La
Roi, 1968). Graham (1969) described the main vegetation zones.
Main Habitat Types and Study Areas
Habitats were classified by indicator plant species (Elton and Miller, 1954) and changes in
water level within and between years were noted. Wet habitats (Figure 1) were classified as
marsh or transition zone.
MARSH ZONE. - There were two main marsh habitats: (a) deep marsh was characterized
by a floating mat of Typha latifolia L. which extended from lakeside to shallow marsh; this
subzone corresponds with reed swamp as defined by Moss (1953,1955); (b) shallow marsh
which extended from deep marsh to Salix habitats in the transition zone. It consisted of
tussocks of Carex species among pools; leading dominants included Carex aquatilis Wahlenb.
and C. rostrata Stokes
TRANSITION ZONE. — Transition zones were delimited readily by shallow marsh on one
frontier bordered by Salix habitats, and forest margins on the other frontier (Figure 2). The
main habitat types were the following: (a) Salix habitats. These ringed the main marshes
adjoining the lakeside. Specimens of C. rostrata , decreased in abundance where the willow
growths were more dense. Moss and Carex associations occurred in lowlying parts, (b) Sedge
Meadows. These extended from Salix habitats to forest margin, but were isolated in other
areas away from the main marshes. Hollows and pools abounded in these meadows and growths
of C. rostrata or C. atherodes were dominant, (c) Forest margins normally were defined by the
limit of waterlogged soil. Clumps of willows extended into the margins from sedge meadows,
(d) Grassy forest clearings were isolated from the above habitats.
Ecology of four Patrobus species
237
METHODS
The sampling program differed between years because of periodic flooding of habitats in
1969. Larvae and adults of Patrobus species were rare in comparison with some other marsh
carabids and were mainly sampled by pitfall trapping. Various workers, particularly
Greenslade (1964), and Southwood (1966), also Carter (1980) have pinpointed problems
associated with data gathered by pitfall trapping of arthropods. In this study, I have restricted
analyses to presence and absence data and to determining whether individuals of each species
occurred at low or high levels of abundance in the same or different habitat types.
Patrobus populations were also sampled by hand collecting. These beetles were also
collected from under cryptozoan boards (Cole, 1946) and were extracted by Tullgren funnels
from quantitative soil samples brought back to the laboratory. For further details of the
sampling program, see Carter (1971).
RESULTS AND DISCUSSION
Physical Differences Between Years
There were major differences in water levels and extent of waterlogged soil between 1968,
on one hand, and 1969 and 1970, on the other hand. In 1968, melt waters from the winter
snows had largely gone by mid-June and many hollows waterlogged in spring had dried out.
Standing water had disappeared from transition habitats by mid-summer. By contrast, levels of
marsh water in May 1969 were higher than at any time in the previous year. Heavy rains in
July, August and September resulted in transition habitats becoming largely waterlogged and
most hollows were flooded. Water levels remained high throughout 1970.
General Natural History of Patrobus Species
Data about seasonal activity and reproductive patterns of the four Patrobus species in
various transition and marsh habitats are summarized in Figure 2 and Tables 1 and 2.
As noted previously by Lindroth (1961), North American Patrobus species overwinter as
larvae and adults or as larvae only. At George Lake, only P. lecontei overwintered solely as
larvae. Overwintered larvae of all species pupated in late June and July and gave rise to
summer adults. Adults which had overwintered to a second year were known as spring adults.
Both generations occurred together and could only be delimited during the period when
summer adults were teneral with soft elytra.
Habitat Occupation
Ranges of spatial distribution of both larvae and adults of each species differed according to
time of year. Ranges were greatest during the beetles4 main periods of activity between May
and September, and September and November for adults and larvae respectively (Carter,
1971). The ranges were least prior to and just after overwintering, the periods of low activity.
During main periods of activity, individuals of P. stygicus, P. lecontei and P. foveocollis
occurred mainly in different zones from forest clearings to lakeside, but there was overlap of all
species in general habitat occupation (Figure 2). Larvae and adults of P. lecontei occupied
mainly wetter parts of the transition zone, chiefly sedge meadows and Salix habitats, while
individuals of P. stygicus ranged into deep marsh. Adults of P. septentrionis appeared to be as
Quaest. Ent. 1981, 17(3,4)
238
Carter
hygrophilous as P. lecontei but were rare at George Lake and occurred chiefly in sedge
meadows. Individuals of all four species were found in sedge meadows, and Salix habitats, but
only P. lecontei and P. stygicus adults frequented large pools where there was little or no sedge
growth.
In the dry year of 1968, larvae and adults of P. foveocollis predominated in drier parts of
the transition zone, such as forest clearings, and were not associated with water. However,
ranges of both stages of this species expanded radically in the wet years of 1969 and 1970. Both
larvae and adults were widespread in all transition habitats, wet Salix habitats and pools in
sedge meadows, and also in dry aspen poplar habitats. High activity of larvae near the marshes
in 1969 indicated that females oviposited there.
Habitat occupation of adults differed prior to and during periods of reproductive activity.
Breeding spring adults of P. stygicus and summer adults of P. lecontei occupied the same
general habitats in late June and July, namely pools in sedge meadows. Salix habitats, and (to
a lesser extent) shallow marsh. In both years, summer adults and possibly spring adults of P.
stygicus dispersed from these habitats into deep marsh in late July and August (Figure 2). It
was not possible to compare data collected in 1968 with those in 1969 because of the different
sampling programs in each year.
Data from mid-summer of 1969 (Table 3) were analyzed in tests of independence for which
the G-test (Sokal and Rohlf, 1969) was employed, the distribution of which can be
approximated by the X2 distribution. Frequencies of adults of P. stygicus trapped in Salix
habitats, shallow marsh and deep marsh varied according to time (G = 30.1, p < 0.005). After
July 25, activity in Salix habitats and shallow marsh decreased while that in deep marsh
concomitantly increased. This was not so with P. lecontei adults (G = 5.9, p <0.1) which were
not trapped in deep marsh.
Range of spatial distribution of larvae and adults decreased as they dispersed to drier
habitats such as Salix, banks of pools, and forest margin, prior to overwintering (Figure 2). No
adults of P. lecontei were found to survive past their breeding season, but adults of the other
three species tended to occur together in fall and winter. There were differences among late
third instar larvae and pupae with respect to vertical distribution above and below soil surface.
Larvae and pupae of P. stygicus were mainly in hummocks and bark above soil, while those of
P. lecontei were found above and below soil. Despite intensive sampling, no larvae and pupae of
P. septentrionis or no late third instar larvae and pupae of P. foveocollis and P. septentrionis
were found. These stages of these two species probably occurred deeper in soil.
It is evident from the above data that different habitat associations of the Patrobus species
at George Lake, particularly during their main periods of activity, were reflected mainly in
horizontal distributions. Seasonal differences in habitat occupation have also been observed for
adults of some marsh carabids in Britain. Dawson (1965) and Murdoch (1966) found clear
differences in summer habitats between a number of congeneric marsh carabids, but observed
that they overwintered together in drier habitats. It may be that adults of hygrophilous
carabids normally aggregate in dry overwintering quarters, such as bark in forest margin, but
disperse to different spring and summer habitats according to their respective responses to
water. Thus congerneric carabids for example, such as those of the Patrobus species at George
Lake, may come to occupy different habitats during their main breeding seasons.
Ecology of four Patrobus species
239
Taxonomic Characters
A number of biologists (for example, Mayr, 1963 : 59ff.) have stressed the importance of
studying physiological, behavioural, and ecological characters together with morphology. A
number of ecological characters of the Patrobus species studied at George Lake (Tables 1 and
2) are here related to the classification and phylogeny of the group.
In a short-term study, such as this, it is difficult to delimit ecological characteristics of
relevance to classification of a group of organisms, particularly when the study is confined to
one part of the species4 range. It is important to consider characters which are more general in
nature and which are not likely to be affected markedly by environmental factors peculiar to
any one area. Of the characters included in Table 1, period of reproductive activity, seasonal
activity patterns of adults and larvae, and pupation sites are likely different in various parts of
the species4 range (for example, see Greenslade, 1965). Other characters such as overwintering
stages, habitat occupation of adults and larvae, maximum length of adult life, and maximum
age at which reproduction occurs are likely less affected. For instance, Lindroth‘s ecological
notes (1961) indicate that habitats occupied, by P. stygicus and P. lecontei, in relation to water
are uniform in much of their range.
In this study, observations were made over a wide range of weather conditions as differences
were noted in habitat occupation, for example, between the dry year of 1968 and the wet years
of 1969 and 1970. Thus, a better range of this ecological character was obtained. For instance,
it appears that populations of P. foveocollis at George Lake are adapted to a wide spectrum of
environmental conditions. This species is thought not to be associated with water (Lindroth,
1961), but the local distribution of populations in some other parts of the species range may
expand and contract as they did at George Lake, according to weather changes between years.
Adults of P. stygicus are readily distinguished from those of P. lecontei by leg colour,
features of prothorax, (see Figure 3), and male genitalia (Darlington, 1938; Lindroth, 1961).
Likewise, the two species are distinguished by a number of ecological characters, including
degree of hygrophily, age at which reproduction begins, maximum age at which reproduction
occurs, and length of adult life (Table 2).
Patrobus longicornis Say frequents temperate environments in meadows, deciduous forests
and arable land. It often occurs near the margins of lakes and rivers (Lindroth, 1961), but does
not occur at George Lake. Features of habitat occupation of P. stygicus and P. lecontei (Table
1) indicate that these two species are more closely related to one another than either is to P.
longicornis so also do morphological characters (Darlington, 1938).
The four Patrobus species at George Lake share a number of ecological characters. These
include overwintering as larvae only or as larvae and adults, and overlap in general habitat
preference (Tables 1 and 2). These and morphological characters, which Darlington (1938)
Kuhnelt (1941) and Lindroth (1961) described, indicate the close relationship of these species.
Regarding stage(s) of overwintering, the majority of carabids of marshes and surrounding wet
habitats of North America and Northern Europe overwinter as adults only (see Lindroth, 1949,
1963b, 1961-1969; Murdoch , 1967). The six North American Patrobus species (one of which,
P. septentrionis, is holarctic) are exceptions, overwintering mainly as larvae or as larvae and
adults (Lindroth 1961). At least two other species, the European P. atrorufus Stroem (see
Thiele 1977) and P. assimilis Chaudoir (see Larsson 1939, Forsskahl 1972) have similar
overwintering stages.
Thiele (1977) noted that certain overwintering carabid larvae, including Patrobus atrorufus
are active on the soil surface in late autumn and winter and are very cold hardy. The larvae of
Quaest. Ent. 1981, 17 (3,4)
240
Carter
P. stygicus, P. lecontei and P. foveocollis at George Lake are similar as they were very active at
low temperatures (-2° to 5°C Carter, unpublished) in November. These were in contrast to
Patrobus septentrionis larvae which were certainly subterranean. The condition of cold
hardiness in larvae which overwinter would be advantageous to species living in boreal
environments often with cold autumns preceding very cold winters. For instance, at George
Lake, it allowed larvae of P. stygicus and P. lecontei to disperse from marsh habitats (in which
during the late summer they hatched from eggs) to drier ones more suitable for overwintering.
PHYLOGENY
The following evolutionary interpretation of life history features and other ecological
characters in based on Darlington’s hypothesis of the phylogeny of the Patrobus species except
that Patrobus fossifrons and Patrobus stygicus are regarded as distinct species rather than
subspecies (see Lindroth, 1961). Darlington used morphological and zoogeographical
characters. He wrote that the genus Patrobus probably originated in Asia and that the
ancestors of the present day North American species migrated into the continent through the
north at three times.
He proposed that the stock to arrive first divided into a boreal line
(proto -fossifrons-lecontei) and a temperate line (proto- longicornis) which gave rise to
Patrobus longicornis. The former stock “then divided into a western species (proto -fossifrons,
probably west of the rockies) and an eastern species (proto -lecontei, probably east of the
rockies)”. Later proto -fossifrons entered the range of P. lecontei by migrating eastward.
Eastern and western populations of proto -fossifrons became isolated and differentiated, giving
rise to the ancestors of present day populations of P. fossifrons and P. stygicus. Darlington
included these four species in the subgenus, Neopatrobus.
Morphological changes (for example, in the male genitalia, pronotum, leg size and colour)
which took place during the evolution of the Neopatrobus group proceeded from a primitive
extreme in P. longicornis to the derived extreme in P. stygicus. The series of characters can be
described as a morphocline (Maslin, 1952).
The ecological trends of the ancestors of this first Patrobus stock that reached North
America may be as follows. Patrobus longicornis frequents open habitats, mainly in temperate
region(s) but appears not to be directly dependent on water (Lindroth 1961). This species
appears to be the most primitive one of Neopatrobus (Darlington, 1938) and so the ancestors of
this line may have frequented such habitats.
The lecontei-fossifrons-stygicus line diverged from temperate habitats into wet boreal ones,
with the proto -fossifrons-stygicus stock becoming the more hygrophilous. The ancestors of P.
stygicus probably dispersed into wetter habitats (marsh) than those frequented by the ancestors
of P. lecontei. Today, at George Lake at least, this is reflected in the segregation of P. stygicus
summer adults from P. lecontei adults during the breeding season.
These habitat associations can be described in terms of an ecocline. This dine proceeds from
the primitive extreme in P. longicornis which does not occur in wet habitats to P. lecontei the
largely transition zone dweller, at the borders of standing waters, to the derived extreme in P.
stygicus which is a transition zone and marsh dweller and is found in standing waters. Thus,
this ecocline extends from the primitive condition of least hygrophily to that of most
hygrophily. It runs in the same direction as the morphoclines identified by Darlington (1938).
The habitat preference ecocline is summarized in a phylogenetic diagram in Figure 3.
Ecology of four Patrobus species
241
Patrobus foveocollis and P. septentrionis are presumed to have migrated to North America
later than the Neopatrobus stock (Darlington, 1938), and P. foveocollis remained in drier
boreal habitats than those entered by the Neopatrobus stock. The Patrobus septentrionis stock
converged ecologically on species of Neopatrobus and invaded wet boreal habitats. Both P.
foveocollis and P. septentrionis retained the primitive condition of overwintering as larvae and
adults.
ACKNOWLEDGEMENTS
I am indebted to G.E. Ball for his supervision and encouragement. Thanks are due to: G.
Pritchard, G.E. Ball and J.R. Spence for criticisms of drafts of the manuscript; H. Goulet for
discussions and invaluable assistance with identifications; Sv.G. Larsson for providing P.
septentrionis larvae; N. Chymko for technical assistance and Mr. E. and Mrs. M. Donald, the
late Mr. G. Donald and Mrs. G. Donald, neighbours of the field station, for help in many ways
and their generous hospitality. Financial support was received through a National Research
Council of Canada grant, A- 1939, held by G.E. Ball, and from the Boreal Institute of the
University of Alberta.
REFERENCES
Ball, G.E. 1966. A revision of the North American species of the subgenus Cryobius
(Pterostichus, Carabidae, Coleoptera). Opuscula Entomologica Supplementum 18,
Lund. Sweden, 166 pages.
Carter, A. 1971. Some aspects of the comparative ecology of populations of the four Patrobus
species (Coleoptera: Carabidae: Patrobini) at George Lake, Alberta. M.Sc. Thesis,
University of Alberta.
Carter, A. 1980. Agonum retractum (Coleoptera: Carabidae) in an aspen woodland in the
Canadian Rockies: population biology and chemical cycling. Pedobiologia 20:101-1 17.
Cole, L.E. 1946. A study of the cryptozoa of an Illinois woodland. Ecological Monographs
16:49-86.
Cole, L.E. 1954. The population consequences of life history phenomena. Quarterly Review of
Biology 29: 103-137.
Darlington, P.J. Jr. 1938. The American Patrobini (Coleoptera, Carabidae). Entomologica
Americana 18:135-183.
Dawson, N. 1965. A comparative study of the ecology of eight species of fenland Carabidae
(Coleoptera). Journal of Animal Ecology 34:299-314.
Elton, C.S. and R.S. Miller. 1954. The ecological survey of animal communities with a
practical system of classifying habitats by structural characters. Journal of Ecology.
42:460-496.
Forsskahl, B. 1972. The invertebrate fauna of the Kilpisjarvi area, Finnish Lapland 9.
Carabidae, with special notes on ecology and breeding biology. Acta Societa pro Fauna
et Flora Fennica 80: 99-119.
Graham, P. 1969. A comparison of sampling methods for adult mosquito populations in central
Alberta, Canada. Quaestiones Entomologicae. 5:217-261.
Greenslade, P.J.M. 1964. Pitfall trapping as a method for studying populations of Carabidae
(Coleoptera). Journal of Animal Ecology 33:301-310.
Greenslade, P.J.M. 1965. On the ecology of some British carabid beetles with special reference
Quaest. Ent. 1981, 17 (3,4)
242
Carter
to life histories. Transactions of the Society for British Entomology 16:149-179.
Kiihnelt, W. 1941. Revision der Laufkafergattungen Patrobus and Diplous. Annalen des
Naturhistorischen Museums in Wien 51:151-192.
La Roi, G.H. 1968. Taiga. In Alberta, a natural history. W.G. Hardy, Editor. M.G. Hurtig,
Edmonton, 343 p.
Larsson, Sv.G. 1939. Entwicklungstypen und Entwicklungszeiten der Danischen Carabiden.
Entomologiske Meddelelser 20: 277-560.
Lindroth, C.H. 1949. Die fennoskandischen Carabidae, Eine Tiergeographische Studie. Ill
Allgemeinen Teil - Goteborg. 91 1 p.
Lindroth, C.H. 1961-1969. The ground beetles (Carabidae excl. Cicindelinae) of Canada and
Alaska. In six parts as supplementa of Opuscula Entomologica Part 1, Supplementum
35, 1-XLVIH; Part II, Supplementum 20, 1961, pp. 1-200; Part III, Supplementum 24,
1963, pp. 201-408; Part IV, Supplementum 29, 1966, pp. 409-648; Part V,
Supplementum 33, 1968, pp. 649-944; Part VI, Supplementum 34, 1969, pp. 945-1192.
Lindroth, C.H. 1963b. The fauna history of Newfoundland, illustrated by carabid beetles.
Opuscula Entomologica Supplementum 23, 112 p.
Maslin, T.P. 1952. Morphological criteria of phyletic relationships Systematic Zoology.
1:49-70.
Mayr, E. 1963. Animal species and evolution. Harvard University Press, Cambridge, Mass.
Moss, E.H. 1953. Marsh and bog vegetation in northwestern Alberta. Canadian Journal of
Botany 31:448-470.
Moss, E.H. 1955. The vegetation of Alberta. Botanical Review 21: 493-567.
Murdoch, W.W. 1967. Life history patterns of some British Carabidae (Coleoptera) and their
ecological significance. Oikos 18:25-32.
Sokal, R.R. and F.J. Rohlf. 1969. Biometry. The principles and practice of statistics in
biological research. W.H. Freeman, San Francisco. 776 p.
Southwood, T.R.E. 1966. Ecological methods. Methuen, London. 391 p.
Thiele, H.U. 1977. Carabid beetles in their environments: a study in habitat selection by
adaptations in physiology and behaviour. Springer Verlag, Berlin 369 p.
SPECIES LARVAE PUPAE
General habitats Overwintering Main period of Active on soil surface Pupation Main sites
stage(s) seasonal activity in Autumn
P. stygicus Transition to deep Larvae, Adults ( 1 ) July- ? yes late May- late June Hummocks, bark in
marsh (2) Sept.- Dec. transition
Ecology of four Patrobus species
243
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Figure 1. George Lake field station showing wet habitats and study areas. The station numbers of study areas
were designated by the grid reference of the S.W. corner of the quadrant(s) in which each occurred. The
habitats in which the study areas occurred and their station numbers were: adjoining the lake, main marsh,
Salix and pools in sedge meadows (30, 31, 45 and 46); forest margin (29S), grassy forest clearings (83 W and
84W); small marsh (22N) and sedge meadows (22, 41N).
Ecology of four Patrobus species
247
Quaest. Ent. 1981, 17 (3,4)
Figure 2. Occurrence of larvae and adults of Patrobus species in wet habitats at George Lake.
248
Carter
DECIDUOUS FORESTS,
NEAR RIVERS
Figure 3. Diagram of the possible phylogeny of the North American Patrobus (Neopatrobus)
species, after Darlington (1938). A summary of the geographical distributions and habitat
preferences of these species is included.
Quaest. Ent. 1981, 17 (3,4)
CLASSIFICATION, PHYLOGENY, AND ZOOGEOGRAPHY OF HYDATJCUS LEACH
(COLEOPTERA: DYTISCIDAE) OF NORTH AMERICA1
RE. Roughley
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Quaestiones Entomologicae
17:249-309 1981
D.H. Pengelly
Department of Environmental Biology
University of Guelph
Guelph, Ontario, Canada
N1G2W1
ABSTRACT
The five Nearctic species of Hydaticus Leach are assigned to the subgenera Guignotites
Brinck and Hydaticus sensu stricto. These are H. (H.) modestus Sharp, 1882 (=H. rugosus
Poppius, 1905 new synonomy; type area- Ust Aldan, U.S.S.R.), H. (H.) piceus Leconte, 1863,
and H. (H.) cinctipennis Aube, 1838 which is recognized as a valid taxon. The Nearctic
species of H. (Guignotites) are H. (G.) rimosus Aube, 1838 and H. (G.) bimarginatus (Say,
1831). Because of the similarity between H. rimosus (type not located ) and H. bimarginatus a
neotype is designated for H. bimarginatus from the LeConte collection. All of these taxa are
keyed, diagnosed and described in terms of morphological features of adults, geographical
distribution and habitat. Hydaticus riehli Wehncke, 1876 ( type area— Cuba) is considered to be
a species of uncertain placement.
A reconstructed phylogeny of genera and subgenera of Hydaticini indicated that this tribe
is comprised of two genera: Hydaticus Leach and Prodaticus Sharp; and that Hydaticus
includes the subgenera: Hydaticus (s. str.), Guignotites Brinck, Hydaticinus Guignot, and
Pleurodytes Regimbart, new status. The reconstructed phylogeny of Hydaticini and the species
group of Hydaticus s. str. which includes Nearctic species is used in conjunction with other
features of these taxa to construct an hypothesis accounting for the differentiation and
distribution of these taxa. The hypothesis is that the ancestral stock of Hydaticini is of
gondwanian origin, and that present-day Africa has been a zoogeographic centre from which
taxon pulses have originated. The Nearctic Hydaticus fauna is comprised of three separate
colonizations of North America: via a North Atlantic land bridge (ancestral stock of H.
piceus-H. cinctipennis) via Central America after a closure of the Panamanian portal
(ancestral stock of H. modestus). Subsequent vicariance of the two ancestral stocks has
produced the present pattern.
‘Part of an M.Sc. thesis submitted to the Faculty of Graduate Studies of the University of
Guelph by the first author
250
Roughley and Pengelly
RESUME
Les cinq especes ^’Hydaticus nearctiques sont assignees aux sous-genres Guignotites Brinck et Hydaticus sensu
stricto. Ce sont: H. (H.) modestus Sharp, 1882 (= H. rugosus Poppius, 1905, synonyme nouveau; region typique- list
Aldan, U.R.S.S.), H. (H.) piceus LeConte, 1863, et H. (H.) cinctipennis Aube, 1838, cette derniere espece etant reconnue
comme valide. Les especes nearctiques d’ H. (Guignotites) sont H. (G.) rimosus Aube, 1838 et H. (G.) bimarginatus (Say,
1831). Etant donne ressemblance entre H. rimosus (type non localise ) et H. bimarginatus, un neotype a ete designe pour
H. bimarginatus dans la collection LeConte. Une cle d' identification de tous les taxons est presentee, ainsi qu’un
diagnostic et une description des caracteres morphologiques des adultes, la repartition geographique et ihabitat de
chaque taxon. La position taxonomique r/’Hydaticus riehli Wehncke, 1876 (region typique: Cuba) est consideree comme
incertaine.
Une analyse phylogenetique des genres et sous-genres d’Hydaticini indique que cette tribu comprend deux genres,
Hydaticus et Prodaticus Sharp, et qu 'Hydaticus inclut les sous-genres Hydaticus (s. str.), Guignotites Brinck,
Hydaticinus Guignot et Pleurodytes Regimbart, statut nouveau. L’arbre phylogenetique des Hydaticini et des especes
^’Hydaticus s. str. nearctiques est utilise en combinaison avec d’autres caracteristiques de ces taxons pour edifier une
hypothese expliquant leur differentiation et leur repartition geographique. L’hypothese est la suivante: le groupe
ancestral d’Hydaticini este d’origine gondwandienne, et I’Afrique actuelle a ete un centre evolutif ou des “vagues”
successives de taxons ont origine. La faune nearctique ^’Hydaticus est composee de trois “ vagues " distinctes de
colonisation de VAmerique du Nord: une qui entra par la connection Nord-Atlantique (ancetres de la lignee H. piceus-H.
cinctipennisj, une qui arriva par I’Amerique centrale apres la fermeture de I’isthme de Panama (ancetre de la lignee H.
bimarginatus-rimosusj, et une qui penetra par la connection beringienne (H. modestus). Ulterieurement, la vicariance des
deux groupes ancestraux produisit le patron de repartition actuel.
TABLE OF CONTENTS
Introduction . 250
Materials and Methods . 251
Natural History . 254
Taxonomic Characters and Terms . 255
Classification . 257
Genus Hydaticus Leach . 257
Key to the species of Hydaticus of North America . 259
Species of Uncertain Placement . 277
Phylogeny . 277
Zoogeography . 285
Acknowledgements . 287
References . 288
Figures . 294
Index . 313
INTRODUCTION
The genus Hydaticus Leach, 1817, as it is now known, occurs on all continents of the world
except Antarctica. Zimmermann( 1920) recorded 104 species in his world catalogue, only four
of which were recorded from North America. Later authors (Leng, 1920; Blackwelder, 1939;
Blackwelder and Blackwelder, 1948) concerned with North American taxa, modified the list
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
251
and increased the total to five taxa. The only comprehensive treatments of adults of the world
fauna of Hydaticus are those of Aube (1838) and Sharp (1882). The last treatment of
Hydaticus in North America, by Crotch (1873), consisted of a literature review and
summation. Regional works treating various taxa of Hydaticus were for the Pacific Northwest
(Hatch, 1953), California (Leech and Chandler, 1956), Baja California (Leech, 1948), Alberta
(Larson, 1975), Utah (Anderson, 1962), North Dakota (Gordon and Post, 1965), Indiana
(Blatchley, 1910), Florida (Leng and Mutchler, 1918; Blatchley, 1919; Young, 1954), Maine
(Malcolm, 1971), Virginia (Matta and Michael, 1977), parts of the West Indies (Chevrolat,
1963; Young, 1953), and Canada (Wickham, 1895; Beaulne, 1917). Original descriptions,
check lists, nomenclatural notes, and natural history notes comprise most of the remaining
literature on North American taxa of Hydaticus.
Although only a few taxa of this genus are involved, there has been some confusion
concerning their proper identities and limits (Wallis, 1939; Leech, 1948; Young, 1954; Larson,
1975). During this work adult specimens of the genus Hydaticus were examined in an attempt
to understand specific limits, variation, and distribution. A phylogeny is reconstructed for the
genera of the tribe Hydaticini, for the subgenera of Hydaticus and for the species groups which
possess Nearctic representatives.
MATERIALS AND METHODS
Specimens of Hydaticus used in this study were borrowed from a number of institutions and
private collections in North America and Europe. In addition, field collecting of specimens was
carried out by the senior author in Ontario in 1975 and 1976; in 1978 specimens were examined
during an extensive trip to various entomological museums in the U.S.A. and Canada. Some
2014 adult specimens of North American Hydaticus were examined. The collections from
which material was borrowed and curators of these collections are listed below in association
with abbreviations used in the text.
BMNH British Museum (Natural History, London, England; M.E. Bacchus;
BMUW University of Washington, Burke Museum, Seattle, Washington 98195; S.A.
Rohwer;
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii 96818; G.A. Samuelson;
CASC California Academy of Sciences, San Francisco, California 94118; D.H.
Kavanaugh;
CISC University of California, Berkeley, California 94720; J.A. Chemsak;
CMNH Field Museum of Natural History, Chicago, Illinois 60605; E.M. Smith;
CNIC Canadian National Collection, Agriculture Canada, Ottawa, Ontario, K1A
0C6; A. Smetana;
CUIC Cornell University, Ithaca, New York 14853; L.L. Pechuman;
DBUM University of Montreal, Montreal, Quebec, H3C 3J7; M. Coulloudon;
DEFW University of Minnesota, St. Paul, Minnesota 55101; P.J. Clausen;
ELSC California State College, Long Beach, California 90804; E.M. Fisher;
EMUS Utah State University, Logan, Utah 84322; W.J. Hanson;
FNYC F.N. Young, Indiana University, Bloomington, Indiana 47401; and University
of Michigan, Museum of Zoology, Ann Arbor, Michigan 48109; I.J. Cantrall;
GWWC G.W. Wolfe, University of Tennessee, Knoxville, Tennessee 37916;
ICCM Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15213; G.E.
Wallace;
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INHS Illinois State Natural History Survey, Urbana, Illinois 61801; D.W. Webb,
including W.U. Brigham collection;
ISUI Iowa State University, Ames, Iowa 50010; R.E. Lewis;
JCAC J.C. Aube, 283 des Franciscains, Quebec, Quebec;
JFBC J.F. Brimley (now part of CNIC);
JFMC J.F. Matta, Old Dominion University, Norfolk, Virginia 23508;
KSUC Kansas State University, Manhattan, Kansas 66502; H.D. Blocker;
LACM Los Angeles County Museum of Natural History, Los Angeles, California
90007; C.L. Hogue;
MCZC Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts 02138; J.C. Scott and M. Thayer;
MNHP Museum of Natural History, Prague, Czechoslovakia; J. Jelinek;
MSUC Michigan State University, East Lansing, Michigan 48824; R.L. Fisher;
MZHF Museum of Zoology, University of Helsinki, Helsinki, Finland; H. Silfverburg;
NMDC N.M. Downie, 505 Lingle Terrace, Lafayette, Indiana 47901;
NMSU New Mexico State University, Las Cruces, New Mexico 88001; J.R.
Zimmerman;
OSUC Ohio State University, Columbus, Ohio 43210; C.A. Triplehorn;
OSUO Oregon State University, Corvallis, Oregon 97331; J. Lattin;
PMNH Peabody Museum of Natural History, Yale University, New Haven,
Connecticut 06520; C.L. Remington;
ROMC Royal Ontario Museum, Toronto, Ontario, M5S 2C6; G.B. Wiggins;
SCSC St. Cloud State University, St. Cloud, Minnesota 56301; R.D. Gunderson;
SDSU South Dakota State University, Brookings, South Dakota 57006; E.U.
Balsbaugh;
SEMC Snow Museum, University of Kansas, Lawrence, Kansas 66045; G.W. Byers;
SPMC Provincial Museum of Natural History, Wascana Park, Regina, Saskatchewan,
S4P 3V7; R.R. Hooper;
TAMU Texas A & M University, College Station, Texas 77843; H.R. Burke;
UADE University of Arkansas, Fayetteville, Arkansas 72701 ; E.P. Rouse;
UASM University of Alberta, Strickland Museum, Edmonton, Alberta, T6G 2E3; G.E.
Ball;
UBCZ University of British Columbia, Spencer Museum, Vancouver, British
Columbia, V6T 1 W5; G.G.E. Scudder;
UCDC University of California, Davis, California 95616; R.O. Schuster;
UCSE University of Connecticut, Storrs, Connecticut 06268; R.E. Dubos;
UDCC University of Delaware, Newark, Delaware 19711; P.P. Burbutis;
UGIC University of Guelph, Guelph, Ontario, NIG 2W 1 ; D.H. Pengelly;
UICU University of Illinois, Urbana, Illinois 61801; J. Sternberg;
UMDE University of Maine, Orono, Maine 04473; E.A. Osgood, Jr.;
UMRM University of Missouri, c/o S.O. Swadener, INHS;
UCRC University of California, Riverside, California 92502; S I. Frommer;
USNM United States National Museum, Washington, D.C. 20560; P.J. Spangler;
UVCC University of Vermont, Burlington, Vermont 05401 ; R.T. Bell;
UWOC University of Western Ontario, London, Ontario, N6A 3K7; W.W. Judd;
WEHC W.E. Hilsenhoff, University of Wisconsin, Madison, Wisconsin 53706.
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
253
Standardized techniques were used in the dissection of the copulatory apparatus of adult
males. The technique used for preparation of the aedoeagus was that of Smetana (1971, pp.
10-11).
Illustrations of male copulatory apparatus were drawn to scale using a Bausch and Lomb
microprojector. Figures of the dorsal view of the left half of the body, and hind leg were drawn
using a Leitz Wetzlar binocular, stereoscopic microscope equipped with a 20 x 20 square,
ocular reticule. All illustrations are oriented with the anterior end of the specimen toward the
top of the figure. Scanning electron micrographs of various body parts were taken with an
“ETEC autoscan” after plating the structures with gold.
Measurements of body proportions were taken using a stereoscopic microscope fitted with
an ocular reticule. Measurements were taken only when both end points of the structure were
contained within the grid and were in sharp focus. Only mature specimens, in which elytra were
joined completely along the suture and in which the head was drawn tightly into the pronotum,
were used for measurements. The largest and smallest specimens of each species were chosen
for the range of measurements. Measurements were made in terms of whole or parts of squares
of the reticule and converted to millimeters. In addition, length of the median lobe was taken
for adult males of each species.
Samples from various localities were selected for measurement on the basis of available
material. In general, 10 males and 10 females were measured for each locality; however, for
two species this was not possible, and most of the appropriate specimens were measured. Two
measurements were made on each specimen of the sample; viz. total length of body (TL) and
greatest width of body (GW).
A detailed description of the genus lists characteristics common to all species considered in
this study. The following information is provided for each species treated: citation of the
original description and references to other published taxonomic treatments, discussion of type
material, summation of diagnostic characteristics, short descriptions of adults that accentuate
the distinction among various species, notes on variation, notes about natural history based on
either published records or field observations, distribution, chorological relationships,
phylogenetic relationships, and material examined.
All locality records and the general Nearctic distribution given for each species were based
on specimens examined. Some literature records could be based on misidentifications but they
have been recorded for those areas outside the geographic limits of this study. In the locality
lists of specimens examined, the following information is provided: Country, State or Province,
County or Regional Municipality or District, locality, date, collector, collector’s remarks,
acronym of the collection from which the specimens were received, and the number of
specimens received from that collection. Data were recorded as they appeared on the label
except that the first two digits of the year were deleted and the months were converted to
Roman numberals (i.e., 21 May, 1967 became 21.V.67). In a series of specimens from a single
locality, the label with the most complete data for that locality was the one recorded. Complete
locality records for such common species as H. modestus, H. piceus , and H. bimarginatus are
omitted. Copies of these records are stored at the Department of Environmental Biology,
University of Guelph and at the Strickland Museum, University of Alberta. Persons desiring
copies of these lists should contact the authors or these Departments.
The greatest problem facing any taxonomist is determining the best manner in which to
recognize a biological species from series of museum specimens. Various techniques are used
for inferring species status by means of clues supplied by the museum specimens. Our
Quaest. Ent., 1981, 17 (3,4)
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procedure involved grouping specimens by similarity of the male aedoeagus, especially the
median lobe. The underlying assumption is that dissimilar median lobes of various phena,
especially if correlated with consistent differences in other characters, represent reproductive
isolating mechanisms necessary for fulfilment of the biological species definition (Mayr, 1963,
pp. 91, 663) (for a more detailed discussion, especially of exceptions, see Erwin, 1970; Noonan,
1973; Whitehead, 1972). The next step involved associating females with males of that same
taxon.
Each species of Hydaticus of North America is sympatric with at least one other species in
some part of its range. This area of sympatry was considered a test of species status versus
subspecific status (Mayr, 1969; Ross, 1974). If no intermediate specimens were observed,
species status was considered to be confirmed. The areas of sympatry are indicated under the
treatment of each taxon in the section on chorological relationships.
In practice, identification of North American specimens of Hydaticus is not difficult once
the specific limits of variation are understood. Each taxon has distinctive features which should
allow ready assignment of specimens.
The subgenera of Hydaticus proposed by Guignot (1950) are used herein because they are
monophyletic and represent natural evolutionary units. There is the probability of increased
information from such a taxonomic division, and subgeneric characters are used within the key
to species. The characters separating two subgenera were recently clarified, expanded, and
strengthened by Franciscolo (1968).
NATURAL HISTORY
Very little is known about natural history of North American species of Hydaticus , but
European authors have contributed more to the knowledge of natural history of this genus.
Galewski (1971), in his work on Dytiscidae of Poland, recorded the preference of Hydaticus for
ditches, pools, and ponds with abundant vegetation, a deep layer of plant debris or detritus and
low acid content. Adults frequent temporary pools in early spring to take advantage of available
food, expecially the larvae of snow-melt mosquito populations, but they spend the summer in
more permanent waters. The breeding season of the European species is from late spring to
early summer, mainly in May and June. This breeding season often coincides with development
of preferred oviposition sites, which are aquatic, vascular plants, belonging to genera such as
Alisma, Iris, and Typha.
Adult females of Hydaticus possess genital valves which are long, narrow, and knife-like,
but without serrations (Boving 1912; Galewski (1971)). This piercing type of ovipositor is well
adapted for endophytic, egg deposition (Balduf, 1935).
Larvae of Hydaticus are good swimmers, when compared to most other larvae of
Dytiscidae, because of the dense fringe of swimming hairs on all legs and on the last two
abdominal segments. The three larval stages are passed within the littoral zone of the breeding
areas, although some individuals venture into open water. The large tracheae of larvae are filled
with air; consequently larvae float to the surface when not swimming. Vegetation is used as
resting and feeding sites, but the larvae are awkward when crawling. The prey of larvae are
small naiads of Odonata and Ephemeroptera, and larvae of Trichoptera and Diptera (Galewski,
1971). Pupation occurs on land near the larval habitat in the typical dytiscid manner. In
England, Balfour-Browne (1950) recorded emergence of adults from pupae during August and
September. Many newly emerged adults invade temporary pools in late summer and autumn
(Galewski, 1971). Adults overwinter under leaf litter and moss on land, often a considerable
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
255
distance from the nearest body of water (Galewski, 1964, 1971).
There are few known natural enemies of Hydaticus. The gregarine, Bothriopsis histrio was
recorded from individuals by Balduf (1935). Pujatti (1953) recorded the metacercariae of the
trematode genus Lecithodendrium. The chalcidoid Mesotocharis syclospila Forst.
(Hymenoptera: Eulophidae) parasitized eggs of Hydaticus within the stems of Alisma when
that part of the plant was above water (Balduf, 1935).
TAXONOMIC CHARACTERS AND TERMS
Sexual dimorphism. - Adults of Hydaticus possess five main characters which are sexually
dimorphic, other than those of the male and female genitalia. 1) In the North American taxa
studied, female specimens are, on average, shorter and narrower and have a general outline
which is more symmetrically oval than that of males. 2) Male specimens are smooth except for
two rows of widely spaced, shallow punctures on each elytron. Most females have sculpturing
on the prothorax and/or elytra. The sculpturing or rugosity varies from shallow, widely spaced,
longitudinal depressions to deep, closely spaced, irregular folds or furrows. Rugosity varies
markedly within and among populations; general trends are noted under each species
description and discussion. 3) Protarsal claws of males are longer and are bent more sharply at
the base than are their female counterparts. Mesotarsal claws are of different lengths and differ
in amount of curvature between male and female specimens. Unlike other genera of Dytiscidae,
claws of males differ only slightly among taxa. 4) The three basal tarsomeres of the forelegs of
males are expanded into circular acetabula (Fig. 2), and posses sucker-like setae which adhere
to the pronotum of the female during copulation. Four large suckers are located on the basal
segment and these are supplemented by 16 to 18 smaller suckers. 5) The three basal
mesotarsomeres of adult males are expanded into elongate, oval acetabula. Mesotarsal suckers
are of about equal size inter se. Eight suckers are on the anterior side of the tarsal midline and
seven on the posterior side (Fig. 3). The number of suckers on each acetabulum is variable and
is difficult to ascertain on many specimens as the suckers are easily broken. Students of
Dytiscidae (Wickham, 1893; Chatanay, 1910; Balfour-Browne, 1940, 1950; Larson and
Pritchard, 1974) have studied adaptations within the family. Their works indicated that the
form of the front tarsi of adult males and the arrangement and number of sucker-like setae
were diagnostic for higher taxa. Within the North American males of Hydaticus this character
was uniform yet varied. The same basic arrangement is found throughout; however,
arrangement and number of modified setae varied between legs of an individual and within and
among species. These were without diagnostic value in the taxa studied; however, trends in the
number of suckers and their size and pattern are used as a phylogenetic character.
Head. - In Nearctic Hydaticus , colour pattern and the presence or absence of large spots or
maculae on the frons is an important diagnostic character. The frons varied in colour from a
black band against a yellow ground colour to reddish-brown throughout, with infuscated areas.
Pronotum. - Colour of pronotum varied from yellow to black. In most species there is a
basal, black band of variable width. This band is useful in recognition of some specimens of H.
rimosus because it extends to the anterior margin in the form of a wide band. H. piceus and H.
cinctipennis specimens have a more or less unicolourous pronotum except for discal
infuscations. Curvature of the posterior margin of the pronotum is useful for separation of
subgenera. In adults of subgenus Guignotites, the posterior margin is sinuate. The portion along
the midline is extended posteriorly, and the portions lateral to the midline are shortened, but
the postero-lateral corners are recurved into an acute angle (Figs. 37, 43, 48, 49). In adults of
Quaest. Ent., 1981, 17 (3,4)
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Roughley and Pengelly
the subgenus Hydaticus, the portions lateral to the midline are not as conspicuously shortened
in length and the postero-lateral corners of the pronotum are truncate (Figs. 7, 8, 9, 22, 30).
Metatarsal claws. - Shape of the claws of the hind legs is an aid in distinguishing between
some males of the two species of the subgenus Guignotites. In H. bimarginatus , the anterior
(shorter, inferior) claw is deflected downward at the apex (Fig. 44), whereas in H. rimosus it is
not (Fig. 55). This character was considered unreliable because of the possible effect of
abrasion on shape of the apex of the claws. Some males of H. bimarginatus lacked the apical
deflections. The relative length and shape of the metatarsal claws are given in the description of
each taxon.
Metatibial chaetotaxy. - Balfour-Browne (1950) emphasized the use of arrangement,
number, and size of spines on the legs for separating the higher taxa of Dytiscidae. Guignot
(1950) and Franciscolo (1968) used chaetotaxy of metatibiae for separating the subgenera of
Hydaticus. In North American specimens, the row of spines on the posterior (upper, nearest
the venter) side of the metatibia is either in a straight line and parallel to the outer margin of
the tibia (subgenus Hydaticus , Figs. 10, 23, 31); or the row is curved inward basally, and thus
does not form a parallel line with the outer margin (subgenus Guignotites, Figs. 38, 50). Within
the subgenus Guignotites, the number of large spines on the anterior (lower) side of the
metatibiae is important in separating the two species. These spines were broken off in some
individuals and if so, the large pits where the spines originated were counted. The most reliable
way to determine number of spines per tibia is to average the number of spines on both
metatibiae of the same specimen. The number of spines varies between specimens and between
metatibiae of the same specimen, but the specific limits do not overlap and they are in accord
with other taxonomic characters. In H. bimarginatus there are a number of smaller spines
basally which were not counted. Under each species in which the lower, metatibial spines were
used for diagnosis, a table is provided of the average number of spines per metatibia from the
specimens used for the descriptive measurements.
Metasternum. - The metasternum of dytiscid adults is displaced by the anterior extension of
the hind coxae, resulting in the formation of a relatively narrow, transverse sclerite which is
indented on each side of the median portion. The area lateral to the median indentation is
termed the “metasternal wing” (Balfour-Browne, 1950). The straight anterior margin of the
metasternal wing is a diagnostic characteristic of members of the genus Hydaticus.
Dorsal colour pattern. - Franciscolo (1968) suggested that colour patterns in the subgenus
Guignotites were useful in the recognition of some of the species only. The dorsal colour
pattern, however, is useful in the identification of North American material due, in part, to the
small number of taxa concerned. Specimens of H. piceus and H. cinctipennis are yellowish to
reddish-brown, whereas those of the other taxa are black. Adults of all taxa have the general
colour disrupted by various amounts of yellow which varies from an indistinct lateral border in
H. piceus, to lateral and/or sub-lateral stripes among other taxa, to a highly developed pattern
in some females of H. modestus (Fig. 9).
Male genitalic appendages. - The terms used for the genitalic sclerites were those of
Edwards (1950), except for the specialized structures of Hydaticus, as discussed by Guignot
(1933) whose terms were retained.
The male genital capsule, in situ, is rotated 90° to the left( i.e., left paramere is next to
sternum 7). This sinistral twist of the genitalic armature is common to all Adephaga, except
Gyrinidae (Edwards, 1950; Balfour-Browne, 1950). Components of the genitalic capsule are
illustrated in Figs. 4 and 5, and consist of: 1) sternum 8 which is joined to a semi-circular
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
257
anterior arch; 2) the basal portion of sternum 9, pleuron 9, and tergum 9 which are fused into a
dorso-ventral circular ring surrounding the more posterior structures in repose; 3) an elongate
rod or spicule lying on the membrane between the parameres which is possibly a posterior
section of sternum 9 separated from an anterior portion; 4) the preputial covering which is
strengthened but not sclerotized at the apical portion of a membrane which joins the parameres
ventrally; 5) the epipenite, on the dorsal side of the preputial covering, next to the median lobe;
6) symmetrical, hairless parameres which are parallel and of equal length; 7) the median lobe
which extends slightly beyond the parameres and articulates with them basally. All of these
structures are joined by an ensheathing membrane, except for the dorsal side of the parameres
and the median lobe. The aedoeagus is composed of median lobe, parameres, and epipenite.
The position and form of epipenite are important for subgeneric diagnoses. Males of the
subgenus Hydaticus have the epipenite on the preputial covering (i.e., on the opposite side of
the preputial covering to the median lobe, Figs. 4 and 5). In males of the subgenus Guignotites
the epipenite is internal, on the preputial covering, next to the median lobe. The transition from
external to internal position of the epipenite on the preputial covering reverses the orientation of
the median and lateral arms of the epipenite (Figs. 4 and 41). The epipenite in males of the
subgenus Guignotites is generally not as heavily sclerotized as that of males of subgenus
Hydaticus. In the subgenus Hydaticinus, represented only by the South American H. rectus ,
the epipenite is still less sclerotized and is inserted into the apex of the preputial covering.
The shape of the epipenite and of the apex of the median lobe are important in species
diagnosis. The epipenite is a trilobed structure consisting of a median arm and a pair of lateral
arms. Length of the lateral arms, when compared to length of the median arm, and shape of the
three arms are important in specific assignment. In side view, the apex of the median lobe
varies from truncate to acutely angled. The median lobe possesses lateral flanges (Figs. 4 and
5) of a tough membranous substance. The degree of posterior extension of these flanges in
relation to the apex of the median lobe is a useful character.
CLASSIFICATION
Genus Hydaticus Leach
Hydaticus Leach, 1817: 69, 72 [nec Hydaticus Schoenherr, 1825 (Coleoptera: Curculionidae)]. Nearctic references only -
Aube, 1838: 155 (ex parte). - Crotch, 1873: 403. - Sharp, 1882: 907, 908. - LeConte and Horn, 1883: 67. - Horn, 1883:
380. - Wickham, 1895: 149. - Blatchley, 1910; 232. - Beauline, 1917: 124. - Blatchley, 1919: 314. - Leech, 1948: 419. -
Young, 1953: 6. - Hatch, 1953: 235. - Young, 1954: 1 13. - Leech and Chandler, 1956: 332. - Anderson, 1963; 56. - Gordon
and Post, 1965: 23. - Malcolm, 1972: 30. - Larson, 1975: 405. - Matta and Michael, 1977: 48.
Type-specimens. - Dytiscus transversalis Pontoppidian, 1763; designated by Curtis, 1825:
95 (attributing the species to Fabricius), and by Crotch, 1873: 403. Hope, 1939: 131 cited
Dytiscus hybneri Fabricius, 1787( = Dytiscus seminiger Degeer, 1774) as generitype
Duponchel, 1845 (6): 728 designated Dytiscus fasciatus Fabricius, 1775 [=Sandracottus
fasciatus (Fabricius, 1775)] as generitype but this species was not listed by Leach, 1817 (from
Leech, 1948: 419).
Diagnostic combination. - Dytiscinae with the following combination of characters: outer
margin of metasternal wings straight; outer (shorter) spurs at apex of metatibiae acute; apex of
prosternal process rounded; and posterior margins of four basal tarsomeres of hind legs with
Quaest. Ent., 1981, 17 (3,4)
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Roughley and Pengelly
dense fringe of flat, golden hairs.
Description. — Adults. Size medium (North American specimens, TL from 10.9 to 15.4 mm, GW from 5.9 to 8.0),
body form ovate (ratio GW:TL 0.47 to 0.58), outline continuous, widest just behind middle. Colour various, from
yellowish-brown to black; surface shining or dulled by rugosity.
Microsculpture singular, isodiametric, very fine (difficult to see except at high magnification), small pores scattered
throughout. Sculpture of elytron consisting of two linear rows of shallow punctures, not apparent on some females because
of deep, irregular wrinkles on elytron and/or pronotum.
Head. - Form as in subfamily, except inner anterior margin of eyes slightly emarginate, but not as in Colymbetinae
(Balfour-Browne, 1950). Labrum excised at middle, yellowish to reddish-brown, with brush of fine, short hairs. Clypeus
indistinct, fused with frons, yellowish to reddish-brown. Frons of most specimens darkened basally, reddish-brown to black,
many specimens with yellowish maculae (Figs. 6,47). Occiput short, reddish-brown to black. Palps and antennae yellowish
to reddish-brown, segments often infuscated apically. Structure of mouthparts not examined for specific or subgeneric
differences.
Thorax. - Pronotum with lateral margins acute but not margined; wider at base than at head, with lateral outline
rounded; antero-lateral margins produced anteriorly as acute projections; base continuous with bases of elytra, posterior
margin more or less sinuate, projected posteriorly opposite scutellum, curved forward lateral to scutellum. Scutellum
visible, about 1.5 times as long as wide. Prosternal process with ventral surface convex basally to flat apically, apex broadly
rounded, marginal bead complete except apically. Metasternum with short, deep notch for reception of prosternal process;
metasternal wings with anterior margin straight, not extended to epipleura, posterior margin sinuate. Metacoxal plates as
wide or wider than long (ratio from 1.0 to 0.77); metacoxal processes margined, rounded, separated by posterior incision;
metacoxal lines convergent on metacoxal processes and divergent on metacoxal plates, effaced anteriorly.
Legs. - Foretarsi of males with tarsomeres 1, 2, and 3 expanded as a broad circular acetabulum (Fig. 1). Tarsomere 5
longest, others subequal in length. Four rows of sucker-like, circular, modified setae or cupules on acetabulum; ventral
surface of tarsomere 1 with two rows of four and five suckers each, tarsomere 2 with one row of seven suckers, tarsomere 3
with one row of six suckers. Tarsomere 4 with brush of short, golden setae along antero-ventral margin, tarsomere 5 with
two long parallel rows of similar setae ventrally. Anterior claws slightly longer and less robust than posterior claws. Male
protibiae and profemora hollowed out ventrally for reception of acetabula of protarsi and protibiae respectively (Fig. 1).
Antennal comb and marginal setae of protibiae numerous and longer than those of mesotibiae. Protibiae with pegs along
posterior margin near apex, opposed to pits along dorsal surface of tarsomeres 1 and 2 (Fig. 1) (Larson and Pritchard,
1974). Mesotarsi of males with tarsomeres 1, 2, and 3 moderately expanded as an elongate oval acetabulum (Figs. 2 and
3); tarsomere 5 longest, and others subequal, tarsomere 4 of some specimens longer than 2 or 3. Ventral surface of
tarsomere 1 with seven suckers, 2 and 3 with four suckers. Acetabulum with long lateral setae (Fig. 2). Mesotibiae and
mesofemora not modified for reception of mesotarsi or mesotibiae, respectively. Tarsomere 4 with brush of short golden
setae along antero-ventral margin, tarsomere 5 with two long parallel rows ventrally. Claws of middle legs longer than
those of forelegs, more evenly arcuate, subequal in length, and of similar shape.
Protarsi of females not expanded; tarsomere 5 longest, others subequal. Protibiae and profemora not hollowed out
ventrally. Claws subequal in length and similar in shape. Mesotarsi of female not expanded; tarsomere 5 longer than 1, 1
longer than remaining ones which are subequal. Mesotarsal claws longer than, or subequal to, protarsal claws. Otherwise,
legs of females similar to those of males.
Elytra. - Continuous with outer margin of pronotum at base, widest just behind middle, apices rounded or sinuate
(Fig. 18). Shining or dulled by rugosity baso-laterally (Fig. 17). Epipleuron extended to about middle of last visible (6th)
abdominal segment. Lateral margins with irregular row of long, fine setae extending from basal third almost to apex. Rows
of translucent, rectangular spots beginning marginally in basal third, curving inward posteriorly and in most specimens
confluent with division between lateral and sub-lateral stripes if the latter are present. Colour yellowish-brown to black,
many specimens with yellow marginal stripes, and/or sub-lateral stripes, latter recurved suturally at base in some taxa, but
in adults of most taxa curved away from margin posteriorly. Some specimens with transverse, yellow fasciae sub-basally,
with longitudinal vittae from fasciae ending sub-apically (Fig. 9).
Taxonomic placement. - Most authors placed Hydaticus in the tribe Hydaticini of the
subfamily Dytiscinae, except Balfour-Browne (1950), who included it in the subtribe
Hydaticina of the tribe Dytiscini. The only other genus in the tribe Hydaticini is the monobasic
Prodaticus with P. pictus Sharp, 1882, known from Persia, Baluchistan, India and Arabia
(Zimmermann, 1920). The dibasic Pleurodytes Regimbart with P. dineutoides (Sharp, 1882)
known from Java and Borneo (Zimmermann, 1920; Vazirani, 1969) and P. epipleuricus
(Regimbart, 1891) known from Tennasserim, Malewan (Vazirani, 1969) is also a member of
Hydaticini. However, it does not deserve generic status and Pleurodytes is treated here as a
subgenus (NEW STATUS) of Hydaticus s. lat. (see PHYLOGENY). The monobasic
Notaticus with N. fasciatus as type species was described as belonging to the tribe Hydaticini.
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
259
Spangler (1973) placed N. fasciatus as a junior synonym of Aubehydrus speciosissimus
Guignot 1942 and retained it within the subfamily Aubehydrinae. A key to the world genera of
Hydaticini was presented by Zimmermann, 1919: 225, and by Zimmermann and
Gschwendtner, 1937: 4.
KEY TO THE SPECIES OF HYDATICUS OF NORTH AMERICA
1 (0) Pronotum with postero-lateral corners sharply and distinctly acute (best seen in
side view), posterior pronotal margin markedly sinuate and recurved laterally
(Figs. 37, 43, 48, and 49); metatibia with row of spines on disc of posterior
(upper) surface curving inward basally, not parallel to outer tibial margins
(Figs. 38 and 50); and epipenite at apex of preputial covering. (Subgenus
Guignotites Brinck) 4
1" Pronotum with postero-lateral corners truncate, posterior pronotal margin
moderately sinuate but not revurved laterally (Figs. 7, 8, 9, 22, and 30);
metatibia with row of spines on disc of posterior surface straight, parallel to
outer tibial margin (Figs. 10, 23, and 31). Epipenite anterior to apex of
preputial covering (Figs. 4 and 5). (Subgenus Hydaticus ) 2
2 (L) Elytra with broad, lateral, yellow stripes, without sub-lateral stripes but some
females vittate (Figs. 7, 8 and 9); colour black; frons black with two, narrow,
transverse, yellow maculae (Fig. 6). Epipenite of aedoeagus W-shaped, lateral
arms as long as or longer than median arm (Fig. 13); median lobe with lateral
flanges extending almost to apex (Fig. 12)1 H. modestus Sharp
7! Elytra with broad, lateral, yellow stripes absent, narrow, indistinct, marginal
area present or absent; colour reddish-brown; frons not black, without narrow,
transverse, yellow maculae. Epipenite W-shaped, but length of lateral arms 0.66
or less than that of median arm 3
3 (20 Elytra with sub-lateral, longitudinal, yellow stripes, with or without basal
fasciae (Fig. 22); form elongate oval, more parallel sided. Epipenite of
aedoeagus with lateral arms less than 0.50 length of median arm (Fig. 26); apex
of median lobe acute in side view (Fig. 24) H. cinctipennis Aube
3' Elytron without sub-lateral, longitudinal stripes, yellow marginal area present
or not, basal fasciae absent (Fig. 30); form oval, sides more arcuate. Epipenite
of aedoeagus with lateral arms about 0.66 length of median arm (Fig. 34); apex
of median lobe with projection in side view (Fig. 32) H. piceus LeConte
4 (1) Median lobe of aedoeagus angulate at apex in side view (Fig. 39). Anterior
(lower) disc of metatibiae with an average2 of more than 10 large spines (x =
1 1.8; max. = 14; min. = 9) (Fig. 45); basal black band of pronotum restricted
in most specimens to basal 0.33, extended in few specimens to anterior margin
as broad band; elytron with sub-lateral stripes without inward extensions of
yellow, transverse fasciae absent (Fig. 37) H. bimarginatus Say
4' Median lobe of aedoeagus truncate at apex in side view (Fig. 51). Anterior disc
of metatibiae with average of less than 10 large spines (x = 6.2; max. — 10;
1 While this manuscript was in press, a paper by Nilsson (1981) appeared which suggests that
the valid name of this taxon is H. aruspex Clark, 1864.
2 one-half the total number of spines on both metatibiae.
Quaest. Ent., 1981, 17 (3,4)
260
Roughley and Pengelly
min. = 2) (Fig. 56); basal black band of pronotum often extended to or almost
to anterior margin; elytra with sub-lateral stripes, sometimes with inward
extensions of yellow, transverse fasciae present in some specimens (Figs. 48, 49)
H. rimosus Aube
Subgenus Hydaticus
Guignot, 1950: 104. - Franciscolo, 1968: 48.
Diagnostic combination. - Frons black or infuscated basally, reddish-brown. Frons, when
reddish-brown, lacking distinct, contrasting maculae; maculae represented by variable
infuscations (Figs. 21 and 29). Frons, when black, with distinct, yellow maculae (Fig. 6).
Pronotum with posterolateral corners truncate, pronotal margin moderately sinuate
posteriorally but not recurved laterally (Figs. 7, 8, 9, 22 and 30). Row of spines on disc of
posterior surface of metatibiae straight (Figs. 10 and 23) or only slightly curved inward basally
(Fig. 31). Elytra black to yellowish-brown; stripes absent, or only marginal, or only sub-lateral
present; transverse basal fascia present or absent. Elytra, when black, with yellow colouration
in form of transverse basal fasciae, and longitudinal vittae in some female specimens (Fig. 9).
Epipenite of male genitalic capsule on opposite side of preputial covering to the median lobe
(Figs. 4 and 5); heavily sclerotized.
Hydaticus ( Hydaticus ) modestus Sharp3
Figs. 4-19. Distribution map, Fig. 20
Hydaticus modestus Sharp, 1882: 650. — Wallis, 1939: 126, 127. -- Hatch, 1953: 235. — Leech and Chandler, 1956: 332.
— Anderson, 1962: 73. — Gordon and Post, 1965: — Larson, 1975: 405.
Hydaticus americanus Sharp, 1882: 651. — Zimmermann, 1919: 225. - Wallis, 1939: 126, 127.
Hydaticus stagnalis , Crotch, 1873: 404. — Horn, 1883: 280. — Wickham 1895: 150. — Blatchley 1910: 232, 233. —
Beaulne, 1917: 124. — Zimmermann, 1919: 225, 226 {ex parte). — Zimmermann and Gschwendtner, 1937: 10 {ex parte).
— F. Balfour-Browne, 1950: 300 {ex parte). — Zaitzev, 1953: 307 {ex parte). — Watts, 1970: 727. — nec Fabricius, 1787:
191.
Hydaticus laevipennis , Sharp, 1882: 651. - Blatchley, 1910: 233. - Beaulne, 1917: 124. - nec Thomson, 1867: 88.
Hydaticus rugosus Poppius, 1905: 23, 24. NEW SYNONOMY. Zaitzev, 1910: 44. — Zimmermann and Gschwendtner,
1937: 17.- Zaitzev, 1953:330.
Hydaticus bimarginatus , Wickham, 1895: 150. — nec Say, 1931: 5.
Notes on synonomy and type material. - Holotypes of H. modestus and H. americanus
were examined by Larson (1975) and were not re-examined in this study. The holotype (9) of
H. rugosus was borrowed from the Zoological Museum, University of Helsinki, Helsinki,
Finland. It is labelled as follows: Ust Aldan; FI. Lena; B. Poppius: 877 (pink label); Mus. Zool.
H:fors, Spec, type No. 1953, Hydaticus rugosus Popp.
Nomenclature of Nearctic H. modestus and Palaearctic H. stagnalis (Fabricius, 1787) has
long been confused. J. Balfour-Browne (1944, p. 355) proposed H. continentalis as a new name
for H. stagnalis which was described originally as a member of Dytiscus and thus the latter
name is itself a junior homonym of Dysticus stagnalis Fourcr., 1785, which is itself a junior
synonym of Dytiscus semisulcatus O. Muller, 1776. Most subsequent European workers — e.g.,
F. Balfour-Browne (1950) and Guignot (1947), did not accept the name H. continentalis ,
3While this manuscript was in press, a paper by Nilsson (1981) appeared which suggests that
the valid name of this taxon is H. aruspex Clark, 1864.
Table 1. Variation in the length and greatest width of body of selected samples of North American taxa of Hydaticus (Coleoptera: Dytiscidae).
Measurements in mm.
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
261
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(continued on next page)
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Hydaticus Leach (Coleoptera: Dytiscidae) of North America
263
Quaest. Ent., 1981, 17 (3,4)
Table 2. The numbers of Hydaticus (H.) modestus Sharp with the development of yellow on the elytra, characterized as vittate (Vj.fasciate (F), and
non-fasciate ( N ); and of elytral sculpture, characterized as rugose (R), and smooth (S).
264
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Hydaticus Leach (Coleoptera: Dytiscidae) of North America
265
presumably because the North American H. modestus was, in their opinions, incompletely
studied (see F. Balfour-Browne, 1950, pp. 299-300; and Guignot, 1947, p. 228, footnote #2)
even though Wallis (1939) had presented evidence that the two were separate. The
distinguishing features used by Wallis were that H. modestus is narrower and more
parallel-sided than H. stagnalis and males of H. modestus are not vittate as are males of H.
stagnalis. The anterior metatarsal claw of H. modestus is proportionately shorter and weaker
than the posterior claw when compared to that of H. stagnalis. The elytral apices are sinuate in
females of H. modestus (Figs. 9 and 18). All of these differences were confirmed in this study
and, in addition, the aedoeagus of males differed consistently. The epipenite of H. continentalis
(Fig. 15) has the middle arm much more expanded apically, and the lateral arms are shorter in
relation to the median arm, when compared to H. modestus (Fig. 13). The median lobe of male
H. continentalis (Fig. 16) is more rounded basally and is much longer (x = 2.9 mm, N = 5)
than that of H. modestus males (x = 2.7 mm, N = 12). Thus, the correct name for the
Palaearctic form is H. continentalis.
Specimens of H. modestus run consistently to H. rugosus Poppius 1905 and not to H.
continentalis (= H. stagnalis) in Zaitzev’s key (1953). The type of H. rugosus was examined
and found to be within the range of variation of similar rugose specimens of H. modestus from
Alaska, Alberta, and the North West Territories. Thus, H. rugosus is considered to be
conspecific with H. modestus. This synonomy supports the distinctness of H. modestus from H.
continentalis, because H. rugosus was described by Poppius who had an adequate knowledge of
H. continentalis. The type of H. rugosus is from List Aldan which is in Yakutskaya region of
the Central Siberian Uplands of the U.S.S.R. This specimen is illustrated in Figure 9.
Wallis (1939) doubted the validity of H. cinctipennis Aube 1838 and suggested that it was
possibly conspecific with H. modestus. However, he believed that there was insufficient
evidence for placing the names in synonomy. It would appear that Wallis had not seen Aube’s
rather detailed original description of H. cinctipennis as he cited only Sharp’s (1882)
subsequent description and the taxonomic notes of Zimmermann (1919). In the present study,
H. cinctipennis is considered as a valid and distinct taxon, separate from H. modestus.
Blatchley (1910), as noted by Larson (1975), seemed to have assigned vittate females to H.
stagnalis and the males and non-vittate females to H. laevipennis. Larson (1975), who
examined the types of H. modestus and H. americanus , stated that the latter is a lightly
sculptured female of H. modestus which is the valid name because of page priority.
Diagnostic combination. - Adults are recognized by the narrow, transverse, yellow maculae
on the frons (Fig. 4). They are the only North American members of the subgenus Hydaticus
with a black ground colour, lateral stripes, and vittate females (Fig. 9). Median lobe and
epipenite of aedoeagus are distinctive.
Description. — Length from 12.8 to 15.4 mm, width from 6.8 to 8.0 mm, other measurements in Table 1. Form
ovate, moderately convex. General colour black. Head black, except for clypeus and two narrow, yellowish to
reddish-brown, transverse maculae on frons (Fig. 6). Pronotum yellowish to reddish-brown except for narrow, transverse,
black band at base, band not extending to lateral margins and restricted to basal third (Fig. 7). Elytra piceous to black;
lateral yellow stripes distinct, slightly recurved suturally at base, marginal, ending before elytral apex except in very few
specimens, appearing divided medially because of rows of translucent rectangles (Fig. 7). Elytra, except for lateral stripes,
piceous to black (Fig. 7), or with transverse, basal fasciae (Fig. 8), or some adult females with fasciae and two to 10
longitudinal vittae (Fig. 9), fasciae and vittae yellow to reddish-yellow. Ventrally, prosternum yellow to reddish-yellow;
remainder of thorax black; abdominal sterna black, margins of many specimens piceous and with yellow to piceous
maculations laterally. Profemora and mesofemora yellowish-red with infuscations, tibiae more infuscated, tarsi darkest.
Hind legs piceous to black. Rugosity absent or present, in the form of a few, shallow depressions baso-laterally on elytra or
deep, irregular wrinkles on pronotum and most of elytra except area along suture (Figs. 9, 17). Anterior metatarsal claw,
bent downward at tip, less than 0.50 length of posterior claw. Median lobe of aedoeagus moderately long (x = 2.7 mm, N
Quaest. Ent., 1981, 17(3,4)
266
Roughley and Pengelly
= 12), truncate basally, apex notched in side view (Fig. 11). In ventral view, lateral flanges extending almost to tip,
apex of median lobe not modified (Fig. 12). Epipenite with lateral arms equal or subequal in length to median arm,
expanded medially, thickened and narrowed basally (Fig. 13). Parameres narrow, acute at apex, with narrow,
translucent flange on apical third (Fig. 14). j
Variation. - Adults of H. modestus vary in the amount of rugosity and yellow coloration on
the elytra. Both of these characteristics show patterns of geographical variation. Data for
selected samples of H. modestus are presented in Table 2 and shown schematically in Figure j
19.
The pronotum and elytra of males are smooth. In females, however, rugosity varies from nil
to almost all of pronotum and elytra being covered by deep, irregular wrinkles (Figs. 9, 17).
Elytra of females from British Columbia, Washington, Oregon, and California are
predominantly smooth, whereas those from Alaska and east of the Rocky Mountains to
southeastern Manitoba are consistently rugose in sculpture. Specimens from scattered localities
in northern Ontario and northern Quebec could indicate an eastward extension of this form. A
few rugose females were seen in population samples from Wisconsin, Minnesota, and
Michigan. Females from eastern Manitoba, eastern and mid-western U.S.A., southern Quebec,
and southern Ontario are commonly smooth, whereas specimens from the southeastern part of
the range are exclusively smooth (Table 2, Fig. 19).
Elytral colouration is more complex than is rugosity: females show three states - vittate,
fasciate, and non-fasciate; males show two states — fasciate and non-fasciate. The vittate
condition is characterized by marginal stripes, transverse basal fasciae and from two, but
commonly eight to ten, narrow longitudinal vittae (Fig. 9). The fasciate condition is
characterized by stripes and basal fasciae only (Fig. 8). The non-fasciate condition is
characterized by marginal stripes only. The degree of concentration of the three female
conditions in each geographic area ranges from: vittate — 59% in New England, New York, and
Pennsylvania to 80% in Manitoba and Saskatchewan; fasciate — 5% in northwestern U.S.A.
and British Columbia to 36% in midwestern U.S.A. ; non-fasciate — 3% in midwestern U.S.A.
to 21% in northwestern U.S.A. and British Columbia. The fasciate condition for males ranged
from 91% in northwestern U.S.A. and British Columbia to 64% in midwestern U.S.A. (Table 2,
Fig. 19). There is some correlation between the abundance of fasciate males and vittate females
within areas, throughout the range. In specimens from northwestern U.S.A. and British
Columbia 91% of males were fasciate and 74% of females were vittate. About half of males
from Alberta were fasciate and about half of females were vittate. In specimens from
Michigan, two-thirds of males were fasciate and two-thirds of females were vittate.
Natural history notes. - Galewski (1964, 1971) presented evidence that adults of European
Hydaticus overwintered out of water as they were found in forest debris or litter. Preliminary
evidence for H. modestus indicates a similar pattern; for instance, adults from Framingham,
Massachusetts were labelled “by sifting”, and “under stone, high dry hill” and were collected
in January, March and April. Specimens from Arlington, Massachusetts were labelled “moss
roots” and were taken in March and December. Fletcher and Gibson (1908) recorded
specimens from moss but did not give time of year. A specimen was collected 13.ix.79 in leaf
litter at George Lake, Alberta approximately 3 m from water line.
Flight records are 13.iv.24 at Framingham, Massachusetts; 21.vi.08 at Algonquin, Illinois;
30.viii-3.ix.69 at Chaffey’s Locks, Leeds Co., and 14.ix.73 at Harrow, Ontario; 3.vi.75 at St.
Cloud and l.ix. 70 at Itasca State Park, Minnesota, and 29.ix.56 at Creston, British Columbia.
Although these records are few, the general indication is of two major periods of flight, spring
and fall, perhaps indicating pre- and post-overwintering movements. However, it is also possible
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
267
that these beetles were intercepted during flight to temporary ponds or between ponds. A
number of specimens were recorded by Hatch (1924) as occurring in beach drift in Charlevoix
Co., Michigan.
Larson (1975) noted that H. modestus was found most commonly in the forested regions of
Alberta. Adults were taken in dense detritus, or emergent vegetation along margins of ponds.
The senior author has taken them in similar situations in Ontario. In Mer Bleu marsh, near
Ottawa, specimens were collected among emergent Carex, in the drainage ditch of a small
beaver pond. Near Moffat, Halton R.M., Ontario, specimens were collected among dead
Typha leaves of a permanent woodland pool on 24.iii.76. Andrews (1923) collected adults by
dredging aquatic plants at Whitefish Point, Chippewa Co., Michigan. James (1970) included
H. modestus in his key to the aquatic beetles of vernal woodland pools in Hastings Co.,
Ontario. Mature larvae of H. modestus were described by Watts (1970) (as H. stagnalis ) but
the adequacy of this description was criticized by Galewski (1975). Records for teneral adults
are: 24.vi.33 at Beach, Illinois; lO.viii at L’Anse, Michigan; 14.vii.27 at Long Beach, Long
Island, New York; 12.vii.56 at Chatterton, 2.vii.l9 at Port Stanley and 4.vii.57 at Spanish, all
in Ontario.
Larson and Pritchard (1974) suggested that, for the males, there was a stridulatory function
for the dorsal surface of the expanded protarsi and the accompanying pegs on the protibiae.
Distribution. - Map, Fig. 20. In North America, this species is transcontinental and ranges
as far south as California, Missouri, and the New England states. Zaitzev (1953) recorded H.
rugosus from Ust’-Aldan, Yakutsk, Bulun, and Irkutsk, in Siberia, U.S.S.R. As well as the
type of H. rugosus , the following specimens from the U.S.S.R. were examined: Sibir oc.,
Barnaul am Ob, v.20, leg. Babiy, (CUIC), 2; Barnaul am Ob, 25.iv.18, Babiy, (CUIC), 1.
Chorological relationships. - H. modestus is sympatric with H. piceus in northeastern
U.S.A. and from Alberta to Quebec in Canada. Possibly, H. modestus is sympatric with H.
continentalis in Siberia, U.S.S.R. (Zaitzev, 1953).
Phylogenetic relationships. - H. modestus is more closely related to a complex of
Palaearctic Hydaticus , centering around H. continentalis, than it is to other Nearctic
Hydaticus.
Material examined. - The number of specimens examined was 378 66, and 500 99.
Hydaticus ( Hydaticus ) piceus LeConte
Figs. 1, 2, 21-27. Distribution map, Fig. 28.
Hydaticus piceus LeConte, 1863: 23. - Crotch, 1873: 404. - Sharp, 1882: 653. -- Wickham, 1895: 150. - Beaulne, 1917:
125.
Hydaticus piceous Gordon and Post, 1965: 23, misspelling.
Notes on synonomy and type material. - LeConte, in the original description, gave a general
description of the distribution as “Middle States and Canada”, but did not give a specific
locality. The holotype is a male in the Museum of Comparative Zoology, Harvard University,
Cambridge, Massachusetts. It is labelled as follows: 111.; Type 6086 (red label).
Diagnostic combination. - The predominantly reddish-brown colour, absence of elytral
stripes, shape of the median lobe and epipenite of the aedoeagus of males distinguish members
of this species. Adults of H. piceus have been confused, in collections, with those of H.
cinctipennis. The former are distinguished by smaller size, more oval shape, and lack of
sub-lateral stripes and basal fasciae.
Quaest. Ent., 1981, 17 (3,4)
268
Roughley and Pengelly
Description. — Length 11.6 to 14.0 mm, width 6.6 to 7.9 mm, other measurements in Table 1. Form oval,
moderately convex. General colour reddish-brown. Head yellowish to piceous posteriorly; infuscation of frons generally in
form of inverted “Y” (Fig. 21), but in some specimens joined by indistinct, fuscous areas extended inward from eyes in the
form of indistinct maculae. Pronotum yellowish to dark brown, lighter laterally, often with baso-central infuscation or
blotches (Fig. 22). Elytra yellowish to dark reddish-brown, with small, isolated, obsolete irrorations; irrorations denser
basally and laterally; lateral, yellow stripes absent, but lateral, yellowish margin present or absent, when present
indistinctly delimited (Fig. 22). Ventrally, prosternum lightest in colour, yellow to yellowish-red; remainder of thorax
piceous to black; abdominal sterna intermediate in colour, some specimens with yellow maculations laterally. Profemora
and mesofemora of many specimens yellowish with infuscations, tibiae and tarsi usually darker. Hind legs corresponding to
abdominal colour. Rugosity, baso-laterally on elytra and laterally on pronotal disc of females, shallow, and in some
specimens reduced to isolated, elongated depressions. Anterior metatarsal claw abruptly bent before tip, posterior claw
with tip deflexed. Median lobe of aedoeagus moderately long (x = 2.8 mm, N = 10), rounded basally; apex of median
lobe, in side view, with projection (Fig. 24). In ventral view, lateral flanges narrow and extending almost to tip, apex of
median lobe not modified (Fig. 25). Epipenite with lateral arms about two-thirds of the length of median arm (Fig. 26).
Parameres narrow, acute at apex, with narrow, translucent flange on apical half (Fig. 27).
Variation. - Colour of adults varies markedly. Many samples from single localities exhibit
extremes of colour, although specimens from Ontario and Quebec are consistently darker.
Presence or absence of marginal, yellowish area of the elytra is not related to geographic
localities. This yellow area contrasts markedly in many adults with colour of the suture but
colours evenly intergrade between the suture and the lateral margin.
Natural history notes. - Needham and Williamson (1907) recorded adults of H. piceus
from a permanent spring-fed pond at Lake Forest, Illinois where they were taken among Typha
stalks in water as deep as 1 m. Sherman (1913) supported this record in his discussion of
Dytiscidae of meadow ponds. Adults were recorded at large, coloured search-lights at Niagara
Falls, Ontario by Stirret (1936). Only two other records of flight are available: St. Paul,
25.vi.21 and Albert Lea, 10.vii.23; both localities are in Minnesota.
During the fall of 1975 and spring of 1976 adults were observed near Moffat, Halton R.M.,
Ontario. The habitat was a permanent woodland pool with deep, organic detritus and diverse
flora of aquatic vascular plants. In September and October, 1975, adults of H. piceus, assumed
to be newly emerged, were collected among emergent Carex. In April and May of 1976,
specimens were collected consistently among Typha stalks. In the laboratory, females
oviposited into the leaf petioles of Alisma plant ago-aquatica L. (Alismaceae). Eggs hatched
after about two weeks and the larvae were fed on mayfly naiads and mosquito larvae collected
from the pond. Larvae of H. piceus are agile swimmers and were observed to frequent open
water as well as dense vegetation in the laboratory and in pond situations.
Distribution. - Map, Fig. 28. The general range of this species extends from central Alberta
and southern Manitoba southward to Missouri, eastward to the New England states and
northward into Quebec. The Alberta locality (Sturgeon River at bridge 2mi. N.W. of Calahoo,
l.vi.77, K.A. Shaw, in Myriophyllum sp. at river edge, [(UASM), 3<3<3] was unexpected since
the most westerly previous record had been Winnipeg, Manitoba. This species was not found
during the extensive survey by Larson (1975). Subsequently, specimens from Buchanan,
Hudson Bay and Prieceville, Saskatchewan (SPMC) were examined, indicating a sporadic,
localized distribution of this species along the southern edge of the boreal forest of western
Canada.
Chorological relationships. - H. piceus is sympatric with H. cinctipennis in the more
southerly part of its range. It is also sympatric with H. modestus in northeastern United States,
and in Alberta, Manitoba, Ontario, and Quebec in Canada.
Phylogenetic relationships. - This species could be closely related to H. cinctipennis’,
however, the shared derived characters uniting these two species are relatively weak. As well,
the relationship of these two species to other Hydaticus is obscure.
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
269
Material examined. - The number of specimens examined was 169 92 and 206 66.
Hydaticus ( Hydaticus ) cinctipennis Aube
Hydaticus cinctipennis Aube, 1838: 191, 192. --Sharp, 1882: 651 . — Zimmermann, 1919: 225. -Wallis, 1939: 127.
Notes on synonomy and type material. - The type-area specified by Aube was the United
States and the Antilles.
Mr. J.T. Huber kindly searched for the type of H. cinctipennis in various European
museums, including the Aube and Dejean collections of the Paris Museum, the collections of
the National Museum of Belgium, Brussels, and those of the British Museum of Natural
History, London, but without success.
H. cinctipennis has been a source of taxonomic confusion, possibly because of its rarity in
collections. Wallis (1939) hesitated to synonomize this name with that of H. modestus because
he had not seen specimens which agreed exactly with Sharp’s (1882) description and because
Zimmermann (1919) considered H. cinctipennis to be valid. Aube’s original description does
seem to apply very well to the specimens studied although the type series was not located.
Diagnostic combination. - Large size, predominantly reddish-brown colour, short but
distinct, yellow, sub-lateral stripes on the elytra and form of male aedoeagus distinguishes
adults of this species from others of the genus in North America.
Description. - Length 12.8 to 14.7 mm, width 6.9 to 7.6 mm, other measurements in Table 1. Form ovate, moderately
convex. General colour reddish-brown. Frons yellowish-brown to dark-brown posteriorly, infuscation irregular but in form
of indistinct maculae posteriorly in some specimens (Fig. 29). Pronotum of most specimens lighter laterally, some with
baso-central infuscation (Fig. 30). Head and pronotum of some specimens unicolourous, yellowish-orange. Elytra
reddish-brown to dark brown; sub-lateral, yellow stripes distinct, not recurved suturally at base, marginal basally but
curved away from margin posteriorly, ending from two-thirds to three-quarters of elytral length; area between stripes and
outer margins darker; transverse fasciae present (Fig. 30) or absent at base. Ventrally, prosternum lightest in colour,
yellowish to yellowish-brown; remainder of thorax dark reddish-brown to black; abdominal sterna of many specimens
darkest, some with yellow maculations laterally. Profemora and mesofemora of many specimens yellowish with
infuscations, tibiae and tarsi of same colour or darker apically. Hind legs corresponding to abdominal colour. Rugosity
absent, or on females: baso-laterally on elytra and laterally on disc of pronotum, shallow and, on some specimens, reduced
to isolated, elongate depressions. Anterior metatarsal claw about 0.40 length of posterior claw, both slightly bent at tip,
anterior claw more acutely bent in females than in males. Metatibiae ventrally with row of spines only slightly curved
inward at base (Fig. 31), but not as noticeably as in adults of subgenus Guignotites. Median lobe of aedoeagus long x =
3.3 mm, N = 5), broadly rounded basally, apex acutely angled in side view (Fig. 32). In ventral view, lateral flanges
ending well before tip, median lobe broadened apically (Fig. 33). Epipenite with lateral arms about same length as median
arm but extending apically less than half the length of median arm (Fig. 34). Parameres broad, with wide, translucent,
membranous flange on posterior half (Fig. 35).
Variation. - Specimens from southern United States (Florida, Georgia, Mississippi) are
smaller than more northern specimens (Table 1). Also, the head and pronotum of southern
specimens are more uniformly orangish in marked contrast with darker, elytral colour. Elytra
of most of the northern specimens are more uniformly coloured with respect to pronotal colour.
Transverse fasciae at the base of the elytra were evident in 7 of 45 specimens examined.
Fasciae are isolated, pale spots near the base, or full, distinct transverse bands as in Figure 30.
Presence or absence of fasciae was not related to the sex of the specimen.
Natural history notes. - Immature stages of H. cinctipennis are unknown. Habitat of adults
is also unknown.
Distribution. - Map, Fig. 28. The general distribution of H. cinctipennis is inferred from
scattered localities as being the Atlantic coastal plain from New York south to Florida and west
along the Gulf coastal plain to Mississippi, and north in the Mississippi River valley to
Tennessee.
Quaest. Ent., 1981, 17 (3,4)
270
Roughley and Pengelly
Aube (1838) wrote that he had seen a specimen from the Antilles. This is an interesting
record since no other North American species of Hydaticus ( s . str.) shows a tendency toward
toleration of brackish-water situations which would be useful in colonizing and exploiting
aquatic habitats within the Antilles. The only species of Hydaticus known with certainty from
the Antilles are in the subgenus Guignotites, and these occur commonly in brackish water.
Fleutiaux and Salle (1890, p. 352) provided a possible explanation for this anomaly. Felix
Lherminier collected extensively in Guadeloupe and in South Carolina and, upon his death, his
collections were sent to Chevrolat and Dupont. The collections, however, were mixed and
certain species from the U.S.A. were labelled as if they had come from Guadeloupe. Aube
(1838, p. viii-ix) acknowledged use of Chevrolat and Dupont collections. Therefore, it is
possible that the specimen(s) Aube recorded from the Antilles actually came from South
Carolina, U.S.A. which is well within the known range of H. cinctipennis.
Chorological relationships. - This species is sympatric with H. piceus in the northerly part
of its range and with H. bimarginatus in the southern and southeastern United States.
Material examined. - The number of specimens examined was 23 66 and 22 $2 from the
following localities:
UNITED STATES OF AMERICA
CONNECTICUT. Middlesex Co.: Cockaponsett St. Forest, 24.V.49, G.E. Pickford, fire pond, s. section, (PMNH), 1.
DELAWARE. Sussex Co.: Glasgow, 29.vi.53, 30.viii.50, H.E. Milliron, electric light, (UDCC), 2; Rehoboth Beach,
-.viii.34, (ICCM), 1.
FLORIDA. Baker Co.: near Manning, xii. 30.47, F.N. Young, (FNYC), 1. Walton Co.: near Bruce on Fla. Hwy. 20,
x.17.41, F.N. Young, hog wallow, (FNYC), 1.
GEORGIA. Clinch Co.: 2.6 mil. W. Clinch Co.-Ware Co. line on Hwy #177, 8.vi.75, G.W. Wolfe, (GWWC), 1. Lowndes
Co.: (county record only), v. 13.63, E. Hazard, (OSUC), 1. Pierce Co.: Blackshear, viii. 17.50, P.J. Spangler, (USNM),
1.
MASSACHUSETTS, (state record only), (ICCM), 1. Bristol Co.: Dartmouth, 20.x. 06, (MCZC), 1; Fall River, iv. 19.18,
N.S. Easton, (CNIC), 1; Freetown, viii. 16. 12, N.S. Easton, (CASC), 1; No. Attleboro, 6.ix.20, C.A. Frost, (MCZC),
1; Westport, iv. 1 8.03, (MCZC), 1. Middlesex Co.: Lexington, 20.iv.30, Darlington, (MCZC), 1. Norfolk Co.:
Brookline, (MCZC), 1; Stoughton, -.vii.20, Blake, (USNM), 1.
MISSISSIPPI. Jackson Co.: Ocean Springs, vi. 14.31, H. Dietrich, (CUIC), 1.
NEW JERSEY. Atlantic Co.: Atlantic City, 7.4.01, (USNM), 1. Burlington Co.: Atsion, vi. 11.45, J.W. Green, (CASC),
1 . Morris Co.: Great Swamp at end of White Bridge Rd., 2.ix.75, G.W. Wolfe, (GWWC), 1 .
NEW YORK. Richmond Co.: Staten Island, (USNM), 2, (MCZC), 1, -.vii.91, iv.16.05, -.viii.25, (USNM), 3, iv.16-05,
E. Shoemaker, (USNM), 2; Bull’s Head, 1.4.21 (UASM), 1. Suffolk Co.: Long Island, M.L. Linell, (USNM), 1;
Forrest Park, v.21.04, (USNM), 1; Orient, ix.3.49, R. Latham, (CUIC), 2; Riverhead, vi.5.36, vi.9.36, R. Latham,
(CUIC), 2.
PENNSYLVANIA. Somerset Co.: Windber, 8.xi.24, (ICCM), 1.
RHODE ISLAND. Newport Co.: Tiverton, v.8.09, (MCZC), 1 .
SOUTH CAROLINA. Charleston Co.: McClennanville, 2-3.vi.73, R. Turnbow, lite, (UMRM), 1.
TENNESSEE. White Co.: Swamp along Rt. 42, 7 mi. N. Sparta, 19.vi.76, G.W. Wolfe, (GWWC), 2.
VIRGINIA. Nansemond Co.: Dismal Swamp, iv.16-17.65, P.J. Spangler, (USNM), 2; Holland, vi.20.55, (JFMC), 1.
Sub genus Guignotites
Subgenus Guignotites Brinck, 1943: 141, new name for subgenus Isonotus Houlbert, 1934: 124, preoccupied by Lepeletier
and Serville, 1828 (Coleoptera: Scarabaeidae). - Guignot, 1950: 104. - Franciscolo, 1968: 48.
Diagnostic combination. - Frons piceous to black basally; yellowish anteriorly, with or
without distinct maculae (Figs. 47, 36). Pronotum with postero-lateral corners sharply and
distinctly acute, posterior pronotal margin strongly sinuate and recurved laterally (Figs. 37, 43,
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
271
48 and 49). Row of spines on disc of posterior surface of metatibiae curved inward basally, not
parallel to outer tibial margins (Figs. 38, 50). Elytra black; lateral and sub-lateral stripes
present in most specimens, fused in some specimens to form a single, wide lateral band. Elytra
with increased yellow ornamentation in some specimens (Fig. 48). Epipenite of male genital
capsule on same side of preputial cover as median lobe; lightly sclerotized.
Hydaticus ( Guignotites ) bimarginatus (Say)
Figs. 36-45. Distribution map. Fig. 46.
Dytiscus bimarginatus Say, 1831: 5. — Say, 1834: 442. Hydaticus bimarginatus, LeConte, 1869: 556. — Crotch, 1873:
404. — Sharp, 1882: 654. — Blatchley, 1910: 233. — Leng and Mutchler, 1918:89. — Blatchley, 1919: 314. - Young, 1954:
113. — Matta and Michael, 1977: 48. Hydaticus fulvicollis Aube, 1838: 184. Hydaticus rimosusl Young, 1954: 113; nec
Aube, 1838: 182.
Notes on synonomy and type material. - Say’s type area is “Louisiana” but the types were
lost (Le Conte, 1869a, p. VI). However, specimens in the LeConte collection, MCZC, are
considered characteristic of Say’s species (Lindroth and Freitag, 1969). A male from the
LeConte collection is hereby designated as NEOTYPE (MCZC type number 32443). It bears
an orange, circular label (for southern States), and another label “ bimarginatus 4”. It is well
within the range of variation described by Say. It lacks the left mesotibia and mesotarsus, and
claws of the left metatarsus. A neotype is needed to clarify the concept of this taxon because of
the similarity to H. rimosus and because the type of H. rimosus was not found. Aube (1838)
recorded the type area of H. rimosus as the United States, but type material was not located.
The most commonly cited original description of H. bimarginatus is that of Say (1834).
This is possibly because the 1834 description is the one quoted by LeConte (1869b). However,
Scudder (1899) discussed a paper written by Say in 1831 which contained the description of
Dytiscus bimarginatus (see also Bequaert (1951) and Leech (1970, p. 241, footnote 1).
In the original description, Aube suggested that H. fulvicollis could be identical to H.
bimarginatus. Subsequent authors have considered the names to be synonymous. Because Aube
gave a general description of the distribution as the United States and because H. bimarginatus
is the only similar species in the type area, the two are considered synonymous herein, although
the type of H. fulvicollis was not located.
Specimens from Broward Co., Florida, which Young (1954) identified as MH. rimosus ? ”
because of their colour pattern, are similar to a few other specimens from Texas, Louisiana,
Maryland, and Virgina. However, shape of apex of median lobe of the aedoeagus and number
of large spines on the metatibiae places them within the limits of H. bimarginatus as
interpreted here.
Young (1954) suggested that H. rimosus and H. bimarginatus could be subspecies. This
could be so, for there is extensive overlap in colour pattern and a close similarity in other
morphological characters. However, H. rimosus and H. bimarginatus are considered
specifically distinct because of subtle, yet consistent, morphological differences. In addition,
there was a small area of sympatry within the West Indies.
Sharp (1882) noted differences in body shape of H. bimarginatus and H. rimosus. The
pronotum of H bimarginatus is more abruptly curved outward at the anterior projections and
the elytra are less distinctly widened post-medially. The pronotum of H. rimosus is more linear
from the anterior projections to the base and the elytra are distinctly widened post-medially. In
dorsal profile the sides of H. bimarginatus are more parallel than those of H. rimosus. These
differences are noticeable in the sympatric specimens from Cuba; however, these differences
Quaest. Ent., 1981,17(3,4)
American taxa of Hydaticus ( Guignotites ) (Coleoptera: Dytiscidae )
272
Roughley and Pengelly
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Hydaticus Leach (Coleoptera: Dytiscidae) of North America
273
are too subtle to be used as diagnostic characters.
Diagnostic combination. - Adults of this species are recognized by combination of the
following characters: small size, undeveloped sub-lateral, elytral stripes, average of more than
ten large spines on lower disc of metatibiae, and angulate tip of median lobe of aedoeagus.
Description. — Length 1 0.9 to 13.1 mm, width 5.8 to 7.2 mm, other measurements in Table 1. Form ovate,
moderately convex. General colour black. Clypeus and frons yellowish to yellowish-red, except posterior, transverse black
band and antero-lateral projections (Fig. 36), some specimens with maculae on frons enclosed by black areas. Pronotum
yellowish to yellowish-red, except for transverse, black band at base; band not extended to lateral margins, band as wide as
distance between inner limits of basal, recurved portions of elytral stripes and restricted to basal third in most specimens
(Fig. 37), but in some specimens extended as a narrow band to anterior border. Elytra piceous to black; lateral and
sub-lateral, yellow stripes distinct; lateral stripes marginal, separated from sub-laterals or joined to them in many
specimens, in most specimens originating in basal third and ending sub-apically with yellow lobes (Fig. 37) and/or spots;
sub-laterals recurved suturally at base, sub-marginal at base, but curved inward from margin posteriorly, ending from
two-thirds the length of elytra to near apex of elytra, medial and post-medial, sutural extensions of yellow absent; yellow
transverse, basal fasciae absent. Ventrally, prosternum lightest in colour, yellow to yellowish-red; remainder of thorax
black; abdominal sterna piceous to black. Profemora and mesofemora yellow with varied amounts of infuscation, tibiae
more infuscated, tarsi darkest. Hind legs piceous to black, corresponding to adominal colour. Rugosity of pronotum of
females absent or, in most specimens, in form of isolated, circular areas not in contact with anterior or lateral borders of
pronotum (Fig. 43); rugosity of elytra absent from most specimens but in some in form of elongate, shallow depressions
baso-laterally. Anterior metatarsal claw about half the length of posterior claw, both claws bent downward at tip (Fig. 44),
anterior more so in females than in males. Anterior disc of metatibiae with large spines (Fig. 45), (x = 12.2, max. = 16,
min. = 8, N 1 150). Median lobe of aedoeagus short (x = 2.0 mm, N = 10), unevenly rounded basally, apex, in side view,
angulate (Fig. 39). In ventral view, lateral flanges ending sub-apically, apex of median lobe not modified (Fig. 40).
Epipenite with lateral arms extending about two-thirds the length of median arm, narrowed apically; median arm wide,
rounded at tip, thickened medially (Fig. 41). Parameres narrow, acute at apex, with narrow, translucent flange on apical
third (Fig. 42).
Variation. - Adults are quite variable with respect to form of lateral, and sub-lateral elytral
stripes and amount of darkening of pronotum. Most specimens have yellow marginal stripes
beginning at about one-third of elytral length and extending to apex or ending sub-apically.
Lateral stripes on some specimens have two to four apical extensions of yellow, although these
are reduced to isolated spots in a few specimens. Sub-lateral stripes of most specimens have
inner margins curvilinear and in some specimens these stripes are abruptly narrowed, usually at
about two-thirds of elytral length. Inner edges of sub-lateral stripes are uneven in some
specimens but distinct median and post-median sutural extensions of yellow are absent. In some
specimens, lateral and sub-lateral stripes are united as a single, broad, lateral band (Fig. 37).
Black colour at base of pronotum is relatively uniform in width but varies markedly in
length. In most specimens, it is restricted to the basal third (Fig. 37), but in some it extends as
much as three-quarters of pronotal length as a broad, semi-lunar area, or as a narrow, black
band from wide, basal area to anterior margin. Black area does not extend the entire length of
the pronotum as a wide, black band as in some specimens of H. rimosus (Fig. 49). Commonly,
the frons is immaculate (Fig. 36), but inward, lateral and central, anterior infuscations enclose
indistinctly defied maculae. These infuscations vary in darkness and in very few specimens
approach the wide, black area seen in H. rimosus (Fig. 47).
Number of large spines on disc of lower surface of metatibiae (Fig. 45) varies from nine to
18, with the average consistently greater than 10 in more than 150 specimens examined. In
addition, males with an average of 10 to 12 spines were found to have the tip of median lobe of
aedoeagus angulate in side view. Numbers of large spines on left and right metatibiae of
specimens used for descriptive measurements are presented in Table 3.
Natural history notes. - Young (1953, 1954) noted that most specimens were found in clear
temporary fresh-water ponds, but also recorded occurrence of a few specimens in
brackish-water habitats. Young (1954) regarded H. bimarginatus in Florida as an erratic
Quaest. Ent., 1981, 17 (3,4)
274
Roughley and Pengelly
occupant of the lowland and upland regions, but noted that it was more abundant in flatwoods
situations.
Matta (1973) found adults of H. bimarginatus in woodland pools and in thick vegetation of
non-acidic, fresh-water ditches, also beetles were collected in a deep, permanent, sand-bottom
pool with no vegetation and in a small bog characterized by large clumps of Typha latifolia L.
and Juncus effusus L. in the Dismal Swamp of Virginia.
Label data indicate a wide variety of aquatic habitats, from saline (brackish pools, brackish
water, salt marsh), to permanent (woods pond, tupelo swamp, sink hole pond), to temporary or
disturbed (canal, pool in canal, puddle in forest stream, temporary pond). Judging from the
frequency of specimens labelled “electric light”, “black light”, and “black light trap”, ranging
from February to November, adults of this species apparently come to light more readily than
those of any other North American species of Hydaticus. Eggs, larvae, and pupae are
undescribed.
Geographical distribution. - Map, Fig. 46. The general range of this species includes the
Atlantic and Gulf coastal plains of the U.S.A. (New York south to Florida and west to Texas)
and parts of the West Indies.
Chorological relationships. - H. bimarginatus is sympatric with H. cinctipennis over the
whole range of the latter; and with H. piceus in the northern part of the Atlantic coastal plain.
It is sympatric with H. rimosus in Cuba and the Bahama Islands, however, no zone of sympatry
with H. rimosus was found on the continent. More intensive collecting along either side of the
Mexico-U.S.A. border should provide valuable information.
Phylogenetic relationships. - H. bimarginatus and H. rimosus are sister species that
represent an invasion from the Neotropical realm.
Material examined. - The number of specimens examined was 331 66, and 313 29.
Hydaticus ( Guignotites ) rimosus Aube
Figs. 3, 47-56. Distribution map, Fig. 46.
Hydaticus rimosus Aube, 1838: 182, 183. — Chevrolat, 1863: 202, 203. — Sharp, 1882: 654.
Notes on synonomy and type material. - Aube’s type of H. rimosus was not located. In his
original description he stated that H. rimosus was found in Mexico and the Antilles. He
included a possible record for Paraguay.
As interpreted here, H. rimosus does not occur in continental U.S.A. H. stagnalis of Horn
(1894) and H. bimarginatus of Horn (1896) from San Jose del Cabo, Territory of Baja
California, Mexico are most likely variants of H. rimosus with less developed markings of the
sub-lateal stripes (Leech, 1948).
Diagnostic combination. - Adults are recognized by combination of: small size, often highly
developed sub-lateral elytral stripes, average of less than ten large spines on lower disc of
metatibiae, and truncate tip of median lobe of aedoeagus.
Description. — Length 11.1 to 1 3.7 mm, width 6.2 to 7.0 mm, other measurements in Table 1. Form ovate,
moderately convex. General colour black. Head black except clypeus and two, large, oval maculae on frons, yellowish to
yellowish-red (Fig. 47). Pronotum yellowish to yellowish-red, except transverse, black band at base; band not extended to
lateral margins, in most specimens as wide as distance between inner limits of basal, recurved portions of elytral stripes; in
many specimens extending anteriorly as a broad, black band (fig. 49), and in some specimens enclosing a broad area of
anterior margin. Elytra piceous to black; lateral and sub-lateral yellow stripes distinct, lateral stripes marginal, separated
from sub-laterals in some specimens or both stripes joined, originating in basal third in most specimens and ending
sub-apically with yellow lobes (Fig. 48) and/or spots; sub-lateral stripes sub-marginal at base, recurved suturally, and
curving away from margin posteriorly (Fig. 49), ending from two-thirds the length of elytra to near apex, inner edges of
sub-lateral stripes undeveloped (Fig. 49), or with sub-basal, inward extensions connected with basal, transverse fasciae,
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
275
and in a few specimens with medial and post-medial lobes of yellow present as only isolated spots, lobes as wide as
stripe in most specimens (Fig. 48); yellow, transverse, basal fasciae present (Fig. 48) or absent (Fig. 49). Ventrally,
prosternum lightest in colour, yellowish to yellowish-red; remainder of thorax black; abdominal sterna piceous to black.
Profemora and mesofemora yellowish-red with variable amounts of infuscation, tibiae more infuscated, tarsi darkest.
Hind legs piceous to black, corresponding to abdominal colour. Rugosity, in most specimens, restricted to pronotum of
females; varying from small isolated areas laterally to broad bands from anterior to posterior margins, in most
specimens not involving lateral margins. Anterior, metatarsal claw about half the length of posterior claw, bent
downward at tip in females but not in males (Fig. 55). Anterior disc of metatibiae with average of less than ten large
spines (Fig. 56), x = 6.7, max. = 10, min. = 2. N = 42). Median lobe of aedoeagus short x = 2.2 mm, N = 5),
unevenly rounded basally, apex truncate in side view (Fig. 51). In ventral view, lateral flanges ending sub-apically, apex
of median lobe not modified Fig. 52). Epipenite with lateral arms sub-equal to or slightly longer than median arm,
rounded apically; median arm wide, rounded at apex, thickened medially (Fig. 53). Parameres narrow, acute at apex,
with narrow, translucent flange on apical half (Fig. 54.)
Variation. - No specimens of H. rimosus included in this study were without distinct
maculae on the frons. Most had two anterolateral projections of black from the lower band of
maculae (Fig. 47). Size of the black area at base of pronotum was usually greater in H.
rimosus than in H. bimarginatus. In most specimens of H. rimosus , the length of the black
area is greater than one-half of pronotal length, ending sub-apically in most specimens (Fig.
48), and in some specimens involving width of pronotum posterior to head. Lateral and
sub-lateral yellow elytral stripes are extremely varied in size and extent (compare Figs. 48, 49).
Specimens with the least developed stripes resemble those of H. bimarginatus. Basal recurved
portions of sub-lateral stripes, in a few specimens, have short, posterior prolongations. The area
between basal, recurved portions and sub-basal, inward extensions, in some specimens, is also
yellow. Sub-basal inward extensions of yellow and basal transverse fasciae vary from complete
and distinct, to isolated spots, to obscure yellowish-brown areas. Lateral and sub-lateral stripes,
in a few specimens, are joined as wide, lateral stripes, much as in some specimens of H.
bimarginatus , however, in H. rimosus the wide stripes are usually accompanied by yellow
ornamentation. Apical portions of lateral stripes are dilated in most specimens (Fig. 48).
As in H. bimarginatus , the number of large spines on disc of the lower surface of metatibiae
(Fig. 56) varies appreciably. However, there are limits between the two species. Although some
tibiae have only two, others have 10 spines, the average number of spines per tibia of the
specimens observed was consistently less than 10 (Table 3).
Natural history notes. - The immature stages of H. rimosus are undescribed and very little
is known about the habitat of the adults. Specimens from Nayarit, Mexico were labelled “pool
in drying stream bed” and “pool in stream”. Adults were collected at light in late July and
early August in Sinaloa, Mexico.
Geographical distribution. - Map, Fig. 46. This species is found in the Antilles, Mexico,
and southward to at least Honduras. Aube’s record of H. rimosus from Paraguay is based on a
single female specimen with a smooth pronotum. This could represent a distinct taxon as Aube
(1838: 184) suggested because in this study no specimens of H. rimosus were received from
south of Honduras, but of those received from other areas, the pronotum of at least a few
females was smooth.
Chorological relationships. - H. rimosus could be a southern sub-species of H.
bimarginatus as suggested by Young (1954) because differences between adults of the two
species are subtle. Intermediate specimens have not been discovered; however, too few
specimens were seen from northern Mexico to indicate zones of contact. The two species appear
to be sympatric in Cuba and the Bahama Islands.
Phylogenetic relationships. - H. rimosus and H. bimarginatus are closely related species
that, in aggregate, have their closest relatives within the Neotropical Region.
Quaest. Ent., 1981, 17 (3,4)
276
Roughley and Pengelly
Material examined. - The number of specimens examined was 37 68, and 35 $2 from the
following localities:
BAHAMAS, BRITISH WEST INDIES
NEW PROVIDENCE: 4 mi. SW Nassau, iv.8.53, E.B. Hayden, (FNYC), 1.
BRITISH HONDURAS
Punta Gorda, vii.34, (CASC), 1.
CUBA
CAMAGUEY: Camaguey, xii.20.23, J. Acuna, (UGIC), 2. LAS VILLAS: Buenos Aires,
Trinidad Mts., v.8-14,36, Darlington, 2500-3500’, (MCZC), 4. ORIENTE: Cuabitas, Stgo.
de Cuba, v.51, P. Alayo, (UGIC), 1; Cauto El Cristo, (Cauto R.), viii.12.36, (UGIC), 1.
Rangel Mts., P. de Rio, viii.24.36, Darlington, about 1500’, (MCZC), 1. Upper Ovando R.,
vii. 17-20.36, Darlington, 1000-2000’, (MCZC), 1.
GUATEMALA
El Salto, Esquintla, .34, F.X. Williams, (FNYC), 1. Naranjo, El Peten, iii.20.22, H.F. Loomis,
(FNYC), 1. Peten Tikal, iv.9.56, T.H. Hubbell & I.J. Cantrall, at light at camp, (FNYC),
1.
HONDURAS
DEPT. MORAZAN: Zamorano, Esc. Agr. Pan., vii. 6. 48, T.H. Hubbell, 2600’, (carbonal),
(FNYC), 1.
MEXICO
(Country record only), (MUSC), 1. BAJA CALIFORNIA: betw. San Jose del Cabo and
Triunfo, (CNIC), 1; San Jose del Cabo, Fuchs, (CASC), 1. CAMPECHE: Hopelchen, 18
mi. E., xi.28.63, K.L. McWilliams, (NMSU), 1. CHIAPAS: Chuatemoc, viii.28.63, K.L.
McWilliams, (NMSU), 1. COAHUILA: Matamoros, .v., (CASC), 1. COLIMA: 30 mi.
NE Colima, xii.4.48, H.B. Leech, (CASC), 2, (FNYC), 1; Colima, 20 mi. W., vi.6.63, K.L.
McWilliams, (NMSU), 2. JALISCO: La Huerta, 6 mi. N., x.25.66, A.H. Smith & J.R.
Zimmerman, roadside puddle, (NMSU), 2; La Huerta, 6 mi. N. & 2 mi. E., iii.22.71, J.R.
Zimmerman, (NMSU), 1. NAYARIT: 24 mi. N. Acaponeta, vii. 63, F.D. Parker & L.A.
Stange, (UCDC), 1; 20.3 mi. W. Compostela, vi. 19.67, A.R. Hardy, (UCRC), 1; Sierra de
Zapotan, xi.42, E. Paredes, pool in stream, (CASC), 3; 20 mi. SE Tepic, 23.ix.48, pool in
drying steam bed, (CASC), 3. OAXACA: Oaxaca, 20.vii.37, Embury, 3000’, (CASC), 1.
SAN LUIS POTOSI: N. Morelos, iii.21.59, (NMSU), 3; Paletla, xii.19.40, F.N. Young,
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
277
(FNYC), 2; Tamazunchale, Quinta Chilla Cts., 19.vi.71, A. Newton, Trop. sub-evergreen
for., ca. 600’, blacklight trap, (INHS), 2. SINALOA: Coyatitan, 3 mi. E. on road to San
Ignacio, iv.9.75, J.R. Zimmerman, (NMSU), 1; 5 mi. N. Mazatlan, vii.22.72, J. & M.A.
Chemsak, A. & M.M. Michelbacher, (CISC), 1, vii.25.73, J. Chemsak, E.G. Linsley &
A.E. Michelbacher, at lite, (CISC), 1, vii.26.73, J. Chemsak, at lite, (CISC), 1, vii.28.73, J.
Chemsak, at lite, (CISC), 1, vii.29.66, J. Chemsak & J. Doyen, white lites, (CISC), 1,
30.vii.64, W.C. McGuffin, (CNIC), 2, viii.5.64, J.A. Chemsak & J. Powell, black and white
lights, (CISC), 1, viii.5-7.64, H.F. Howden, (CNIC), 1. TABASCO: Villahermosa, 5 mi. S.,
viii.26.63, K.L. McWilliams, (NMSU), 3. TAMAULIPAS: ditch N. of Mante, vi.12.60,
F.N. Young, (FNYC), 3. VERACRUZ: J.D. Covarrubia, 1 mi. N., viii. 26.62, J.R.
Zimmerman, (NMSU), 1; Lake Catemaco, D.C. Robinson, (TAMU), 4; 13 km. WNW
Potrero, vii. 16.48, H.B. Leech, (CASC), 1. YUCATAN: Chichen Itza, Xtolok Cenote,
vi.28.32, E.P. Creaser, (MCZC), 1; Progreso, 3 mi. S., xi.24.63, K.L. McWilliams,
(NMSU), 3.
SPECIES OF UNCERTAIN PLACEMENT
A female specimen of Hydaticus (G.) grammicus Germar, 1830, is labelled: FLA:
Highlands Co.; Archbold Biol. Sta.; 13-X-1964; P.H. Arnaud, Jr., (CASC), Zaitzev (1953)
reported this species from southern U.S.S.R., central and southern Europe, Iran, and Japan.
The Florida specimen appears to be identical to specimens received from European collections.
Most likely it is mislabelled but it could be an isolated occurrence of the species in North
America. Leech (1970) discussed some intriguing additions to the California water beetle
fauna which could have been introduced by means of aquaria supplies.
The name Hydaticus riehli Wehncke, 1876, was not associated with any specimens studied.
The type locality is Cuba but the collection of the Academia de Ciencias, Havana, Cuba
contained no specimens assigned to this name. Sharp (1882, p. 782) was of the opinion that the
species belonged to Thermonectus Dejean.
PHYLOGENY
Introduction
Methods, principles, usefulness, and importance of cladistic analysis for extant faunas are
presented by Hennig (1966), Brundin (1966), Ball and Erwin (1969), Erwin (1970),
Whitehead (1972), Noonan (1973), and Kavanaugh (1972), 1978). Darlington (1970) and
Ashlock (1980) discuss the limitations of cladistic techniques.
Cladistic techniques were used to discover relationships but the formal classification is not
cladistic. Characters and character states used in phylogenetic analysis are presented in Table 4
and cladograms are presented as Figs. 57 and 58. Each character is numbered and the derived
or apotypic state is represented by a filled circle on the cladogram. The ancestral or plesiotypic
state is represented by an open circle. Characters which have three states are considered to have
both a derived and a highly derived state; the last is designated by a prime mark(')-
An attempt to reconstruct the evolutionary history of a group, by cladistic techniques,
employs analysis of transformation series (morphoclines) of two or more character states.
Quaest. Ent., 1981, 17 (3,4)
Hydaticus, and species of Hydaticus (s. str.) (Coleoptera: Dytiscidae
278
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Hydaticus Leach (Coleoptera: Dytiscidae) of North America
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Hydaticus Leach (Coleoptera: Dytiscidae) of North America
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Polarity or direction of each morphocline is critical to an accurate reconstruction because taxa
are grouped on the basis of apotypic rather than plesiotypic character states. Polarity of
characters is often determined by frequency of occurrence. A character state distributed among
all or most members of a monophyletic group is considered to have been present in the common
ancestor of that group. Therefore, ex-group comparisons reveal the plesiotypic character state.
Another technique, called in-group comparison, is used to establish modification and
remodification of characters or polarity.
In determining the relationships of members of Hydaticus (s. str.), distribution of character
states within the subgenus must be known (in-group). Similarly, relationships within Hydaticus
(s. lat .) require knowledge of character states within Hydaticini (ex-group). This should be
supplemented by knowledge of character states among an immediate, higher taxon (ex-group,
viz. Dytiscinae) such that an evaluation could be made of the number of times a character has
been modified - i.e., group trends (Ross, 1974, p. 158). The manner in which character polarity
was determined is given in Table 4.
All characters are not of equal value in reconstructing a phylogeny. For instance, gain of a
character should be considered of more value than loss of a character, and gain of a complex,
intricate character of more value than gain of a simple character. Ranking characters in terms
of value for determining relationships is termed weighting. Character weighting allows
separation of taxa which are similar because of parallel or convergent evolution. Hecht and
Edwards (1976) proposed five weighting categories based on increasing information; a higher
category indicating more information. Ball (1978) used a similar system for Trichopselaphus
Chaudoir (Coleoptera: Carabidae).
The following list of types and characteristics of the weighting system is that of Ball (1978)
modified slightly to apply to the character states present in Hydaticini.
Loss of a structure.
Simplification or reduction of a complex character.
Simple change, involving sclerotization, colour or position.
a. Two states.
b. Three states.
Parts of a functional complex.
The weight of each character used in phylogenetic analysis of members of Hydaticini is given in
Table 4. Specimens of taxa listed below were used for analysis, in addition to the North
American taxa.
Hydaticus ( Guignotites ) dorsiger Aube
I.
II.
III.
IV.
Prodaticus pictus Sharp
Hydaticus ( Hydaticus ) bowringi Clark
H. continentalis J. Balfour-Browne
H. (//.) histrio Clark
H. (H.) seminiger (Degeer)
H. (H.) transversalis (Pontoppidian)
H. (//.) vittatus- group
Hydaticus ( Hydaticinus ) rectus Sharp
Hydaticus ( Pleurodytes ) dineutoides Sharp
H. ( G .) exclamationis Aube
H. ( G .) flavolineatus Boheman
H. ( G .) fractivittis Guignot
H. ( G .) grammicus Germar
H. ( G .) leander Rossi
H. (G.) matruelis Clark
H. ( G .) palliatus Aube
H. ( G .) subfasciatus LaPorte
Analysis of Characters
Some of the characters used in the phylogenetic analysis require explanation because of
possible alternate explanations of character transformation. The analysis would be greatly
Quaest. Ent., 1981, 17 (3,4)
282
Roughley and Pengelly
different if these character states were interpreted differently and therefore the reasons for one
interpretation should be provided.
Mesotarsal acetabulum of male ( characters 1, 2, 3 and 4, Figs. 2 and 3). - These characters
could represent a functional unit, however, they do not show a direct, dependent relationship -
e.g., a large mesotarsal brush of the apotypic “group” state does not always occur in
conjunction with larger mesotarsal suckers. Therefore, the characters are believed to have I
evolved independently and are weighted separately for phylogenetic analysis.
The presence of the brush on mesotarsomere 1 of male adults remains to be tested as a
synapomorphy for all members of Hydaticus ( s . lat.j. Franciscolo (1968) did not illustrate the
mesotarsal brush of any species he studied, except H. (G.) pullatus Guignot (Fig. 14).
However, the brush in the plesiotypic “linear” state is very difficult to see unless one is looking
for it. For instance, an obscure, linear brush is present on the male mesotarsal palettes of H.
(G.) dorsiger, H. (G.) flavolineatus, H. (G.) grammicus and H. (G.) leander even though it is
not shown in Franciscolo’s Figs. 20, 15, 17 and 10 respectively.
The size of suckers on the male mesotarsal acetabulum has been divided subjectively into
three classes based on comparison of mesotarsal suckers to the smaller suckers of the protarsal
palette. If the mesotarsal suckers were much smaller than the protarsal suckers they were
classed as “small”; if they were slightly smaller than or equal to the protarsal suckers they were
classed as “large”; and, if slightly larger than the protarsal they were classed as “very large”.
Perhaps these are not natural groups, but they do appear to be correlated with other characters
in defining lineages.
The number of rows of suckers of the male mesotarsomeres 2 and 3 is difficult to analyze.
Reduction of number of rows is not correlated to increasing size of individual suckers. In fact,
the opposite trend appears — decreasing number of rows and decreasing sucker size — implying
that the male mesotarsus has a different function or a difference in degree of function among
some lineages of Hydaticus (s. lat.).
Row of spines on venter of metatibia. - (character 5, Figs. 10, 23, 31, 38 and 50). The
principal differences used by Guignot (1950) to distinguish Guignotites from other subgenera
of Hydaticus (s. lat.), was a “curved” (Figs. 38 and 50) rather than a “straight” (Figs. 10 and
23) row of spines ventrally on the metatibia. Members of Pleurodytes also possess the apotypic
“curved” state of this character. However, possession of either state of this character is not
definitive subgenerically because H. cinctipennis (Fig. 31), which belongs to Hydaticus (s. str.),
on the basis of a suite of other characters such as position, shape and degree of sclerotization of
the epipenite, also possess a curved row of metatibial spines.
Vittae and apical sinuation of elytra (characters 9, 10 and 13, Figs. 7, 8, 9 and 18. - H.
(//.) continentalis and H. ( H .) modestus have forms in which the elytra are marked with
longitudinal, yellow vittae. In H. ( H .) continentalis most specimens of both sexes are vittate
whereas in H. (H.) modestus only the female specimens are vittate although the percentage of
population samples vary widely (Fig. 19). This sexual distinction is an example of divergent
evolution which could be expected among predaceous water beetles which respond primarily to
visual stimulae. At some time in the past, a complete distinction between the sexes with respect
to vittae would be hypothesized - i.e., females exclusively vittate. Females of some populations
of H. modestus, as mentioned above, have now passed this intermediate state and become
increasingly male-like in colour pattern - i.e., non-vittate.
A similar divergence in sexual characteristics has occurred in the apical sinuation of elytra
of females of H. modestus. Males and females of H. continentalis have rounded elytral apices
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
283
similar to those of male specimens of H. modestus (Figs. 7 and 8). Females of H. modestus,
however, possess an apical sinuation of the elytra (Figs. 9 and 18), possibly allowing distinction
between the sexes by the adult beetles.
Position and sclerotization of epipenite (characters 15 and 15, Figs. 4 and 5). - All males of
Dytiscinae possess a dorsal sclerite on the male aedoeagus which probably is homologous to the
epipenite of male hydaticines. A dorsal positioning of the epipenite therefore would be
plesiotypic. A switch to a ventral epipenite among members of certain subgenera of Hydaticus
could represent a difference in the copulatory function of this sclerite. Correlated to change in
position is a trend toward less heavy sclerotization of the epipenite which culminates in its
absence from males of Hydaticinus.
Width of ventral flange of paramere (character 20, Figs. 14, 27, 35, 42 and 54). - Most
males of Dytiscinae do not possess a ventral flange of unsclerotized chitin on the paramere such
as is present among hydaticines. This flange, in the apotypic state, is as wide as or wider than
the adjacent sclerotized portion of the paramere (Figs. 14, 27 and 35). This state is
characteristic of Hydaticus (s. str.j.
End-point and terminal width of lateral flanges of median lobe (characters 22 and 23, Figs.
12, 25 and 33). - Most males of Hydaticini possess lateral flanges on the median lobe which
taper to a fine point and terminate sub-apically. This represents the plesiotypic state of both
characters. Within the species group of Hydaticus (s. str.), which contains Nearctic taxa, these
two characters vary in a mosaic pattern (Fig. 58). Both characters have become apotypic
independently, however, the combination of apotypic states of both characters (Fig. 12) serves
to demonstrate the monophyly of one complex of species.
Hydaticini as a monophyletic unit
All members of Hydaticini examined during this study possess two unique character states
which could be interpreted as evidence of a monophyletic assemblage. 1) The suture between
the metepisternum and the metasternal wing is straight (Balfour-Browne, 1950, p. 254). 2) A
character state which is of great value because of its complexity is the presence, on all males
examined, of a possible stridulatory apparatus (Larson and Pritchard, 1974, Figs. 38-42). This
apparatus consists of a row of pegs on the tibia (plectrum) opposed to pits on protarsomere 2
(file) (Fig. 1).
Relationships of Higher Taxa of Hydaticini
Proposed relationships of the higher taxa are represented in Fig. 57. Hydaticini are
represented by two genera, one of which includes four subgenera.
Adults of Prodaticus pictus are markedly plesiotypic. The only proposed synapotypy is that
the median lobe is modified from its ancestral form (character 24). This apotypy is
unconvincing because deviations from the plesiotypic, parallel-sided shape of the median lobe
have taken place many times within Hydaticini and Dytiscinae (group trend). Thus, P. pictus
appears to represent a taxon little differentiated from the original body plan of Hydaticini, at
least, in adult form.
Pleurodytes is a highly autapotypic sister group of Guignotites in that they possess a
ventrally positioned epipenite and a curved, ventral row of metatibial spines. The latter
character is not confined to these subgenera since males of H. (H.) cinctipennis have a slightly
curved row of metatibial spines. This curvature is judged to be acquired secondarily in H.
cinctipennis because in all other characters it is a member of Hydaticus (s. str.).
Quaest. Ent., 1981, 17 (3,4)
284
Roughley and Pengelly
Franciscolo (1968) inferred that a species group of Guignotites centered around H. leander
Rossi was the most plesiotypic of the entire genus. However, based on the phylogenetic analysis
presented, Guignotites and Pleurodytes should be considered relatively advanced groups of
Hydaticus ( s . lat.). The leander species group is monophyletic, as shown by Franciscolo,
because of the lack of a posterior row of bristles on the male protarsal acetabulum. However,
this character state must be considered apotypic because the “bristled” state occurs in
Prodaticus and Hydaticus (s. str.). This interpretation is parsimonious in explaining the
complexity of the apex of the median lobe from the simpler forms found in members of
Hydaticus s. str., through “bristled” Guignotites to the more complex forms among
“bristleless” Guignotites (compare Figs. 12, 25, 33, 40 and 52 herein to Figs. 96-107 and
119-132 of Franciscolo, 1968).
Relationships of Hydaticus s. str.
The reconstructed phylogeny of some of the members of the subgenus Hydaticus is shown in
Fig. 58. Only a few taxa were available for study, however, at least one monophyletic species
group could be defined. Also, a structure is provided on which other taxa can be placed as they
become available.
Two major complexes are suggested by the phylogeny: 1) the cinctipennis-piceus complex
and 2) the transversalis-semininger-continentalis-modestus complex. The cinctipennis-piceus
complex is, perhaps, not monophyletic. In Fig. 58 this complex is isolated from the remainder of
the species group on the basis of a colour character and a transformation series. This grouping
is maintained in apparent conflict with good structural characters (relative width and length of
lateral flanges of the median lobe of the male - characters 22 and 23). The conflict is
supported by correlation with a high degree of autapotypy of H. cinctipennis and distribution
patterns. These correlations are suggestive of an early isolation of the common stem of H.
cinctipennis and H. piceus.
Male members of the European H. transversalis also exhibit a high degree of autapotypy in
regard to characters of the aedoeagus. In general facies and elytral colour pattern, this taxon is
closely related to the seminiger-continentalis-modestus part of the species group. It is possible
that H. transversalis represents a more recent lineage than the
seminiger-continentalis-modestus group except that an Upper Miocene fossil has been assigned
to this taxon (Galewski and G/azek, 1978).
The seminiger-continentalis-modestus group is isolated on the basis of apotypic states of
characters 22 and 23. Under “Analysis of Characters” these two characters were presented as
varying in a mosaic pattern except in this group where both characters are found in apotypic
forms. If the pattern is truly mosaic and is shared only coincidentally by these three taxa, then
the group could be paraphyletic.
H. continentalis and H. modestus are almost certainly sister species because they are the
only taxa belonging to Hydaticus (s. str.), observed to possess vittate elytra. If, however, the
ancestral stock of any part of the transversalis-seminiger-continentalis-modestus complex can
be demonstrated to have possessed vittate elytra then continentalis and modestus would be
grouped on the basis of plesiotypic rather than apotypic character states.
Relationships of Guignotites
Most species of Hydaticus (s. lat.), belong to this subgenus and, as such, the complexity of
their relationships is outside the scope of this study. American species are few in number, with
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
285
eight or nine described taxa, as compared to the fauna of other regions — e.g. at least 60 species
in the Ethiopian region (Franciscolo, 1968 and Zimmermann, 1920). The understanding of the
South American fauna is inadequate and the taxa are so poorly delimited that the formulation
of phylogenetic hypotheses is not possible. Characters of possible phylogenetic importance can
be gleaned from Franciscolo (1968) and the present study. Preliminary analysis of the available
material indicates that the two Nearctic taxa assigned to this subgenus are closely related but a
particular species group containing them cannot be discerned.
ZOOGEOGRAPHY
Introduction
Dytiscidae probably arose before the end of the Jurassic (Crowson, 1975). This conclusion
comes in part from the position of the family in the Adephaga, which must have been one of the
first lineages to have diverged from the remainder of the Coleoptera. As well, many groups of
Dytiscidae show gondwanian distribution patterns although subsequent dispersal and
vicariance by means of continental movement have obscured some patterns, as have extinctions.
A serious problem for zoogeographic interpretation is the lack of phylogenetic analysis of
genera and tribes of Dytiscidae. Preliminary analysis of some tribes and genera of Dytiscidae
show that a good working hypothesis for dytiscid zoogeography would be one similar to that of
Noonan (1979) for anisodactyline carabid beetles.
Some evidence for a gondwanian origin of dytiscids is provided by the observation that most
of the Nearctic fauna is derivable from Palaearctic and Neotropical sources (Wolfe, 1979);
North America has very few endemic tribes and genera. Matthews (1979) discusses the major
routes from Eurasia to North America. One or more of the three North Atlantic routes were
probably used by dytiscids during Cretaceous to Eocene times. Invasions from eastern Eurasia
were possible via the Bering land bridge which is thought to have existed at sporadic intervals
from Cretaceous through Pleistocene times (Matthews, 1979).
Faunal affinities to the Neotropical realm indicate that a number of lineages have entered
North America from the south. These invasions have probably come about after one of the
many closures of the Panamanian portal which range in age from late Pliocene to the present
(Childs and Beebee, 1963).
General Patterns of Distribution
Two of the five higher taxa [ Prodacticus and H. ( Pleurodytes )] considered in this study are
confined to small areas within the Oriental region, and another [H. ( Hydaticinus ) from
Argentina north to Trinidad] to the Neotropical realm. Presence of both relict, plesiotypic, and
apotypic taxa, as well as the greatest diversity of Hydaticini in the Palaeotropics leads to the
hypothesis that Hydaticini have arisen and diversified on the continental land mass of Africa
when that continent was part of Gondwanaland. Africa represents the zoogeographic centre
from which taxon “pulses” (Erwin, 1979) have originated. These “pulses” represent forms
capable of rapid colonization of new areas presumably because of entering a new adaptive zone
which has allowed them to out-compete the established inhabitants if any were present. The
first “pulse” of Hydaticini could have given rise to Prodaticus which could have dispersed from
Africa into the Oriental region where it is found today. The next “pulse” is represented by
Hydaticus ( s . str.), which could have replaced Prodaticus in Africa first and secondly in the
Quaest. Ent., 1981, 17 (3,4)
286
Roughley and Pengelly
Palearctic region as the former taxon began to colonize that area. Subsequently, Hydaticus ( s .
str.). was replaced in the tropics by the more advanced Guignotites as the latter began to
diversify.
Distribution of Hydaticus (s. str.) in North America
Hydaticus ( s . str.), is primarily Holarctic in distribution. The three species of Hydaticus (s.
str.) in the Nearctic Region are of quite different ages. H. modestus is a recent arrival whereas
H. piceus and H. cinctipennis represent much older invasions.
The common ancestor of H. piceus and H. cinctipennis probably entered North America
from western Eurasia. The most plausible route would seem to be one of the three North
Atlantic routes discussed by Matthews (1979) ranging in age from Cretaceous to Eocene (Figs.
58 and 60). The H. cinctipennis-piceus ancestor having entered North America diverged into
the two extant taxa. A North Atlantic route for the common ancestor of these two species is
suggested by their present distributions. The two species are confined to the eastern U.S.A.,
and the Atlantic and Gulf Coastal distribution of H. cinctipennis represents a relict
distributional pattern. The vicariant event which most likely came to separate H. cinctipennis
and H. piceus was the fluctuation of epicontinental seas in the southeastern United States in
the Pliocene and Pleistocene (Howden, 1963). Ancestral H. cinctipennis was isolated in central
Florida by marine transgressions whereas the ancestral H. piceus was diverging on the
remainder of the continent to the north of the seaway. This reconstruction presupposes that H.
piceus and H. cinctipennis represent a monophyletic grouping. Further information on the
phylogeny of Hydaticus ( s . str.), will be of great value in testing the zoogeographic hypothesis.
The second invasion of North America by Hydaticus (s. str.), is represented by H.
modestus, a relatively recent species (Figs. 58 and 60). One clue which suggests that H.
modestus is recent is its widespread distribution which is characteristic of more recent taxa.
This species, derived from a common ancestor of it and H. continent alis, entered North
America by means of the Bering land bridge possibly as late as the Pleistocene. The vicariant
event which separated the two taxa was probably been eustatic, sea-level fluctuation which led
to sporadic closings of the Bering land bridge.
Once H. modestus entered North America something can be deduced of its recent history
from distribution of its two female elytral morphs (Fig. 19). Analysis of distribution of
“rugose” and “smooth” elytral sculpture implies two different full-glacial Wisconsin refugia.
These are the Beringian refugium for the rugose form and the eastern-deciduous forest
refugium for the smooth form (Fig. 59 A). Post-glacial warming and retreat of ice northward
would have allowed the smooth form to colonize North America in a westward direction
whereas the Beringian population remained isolated in northwestern North America (Fig.
59B). This Beringian population presumably became adapted to a shorter growing season and
developed rugose elytra of females. In post-glacial times this Beringian morph has colonized
those areas of North America with continental type climates whereas the smooth morph has
predominated in more temperate areas.
Distribution of Guignotites in North America
As mentioned above, under the Relationships of Guignotites, phylogeny of this subgenus is
quite involved and complicated. The fauna within the Americas is only a very small part of the
total fauna, members of which occur world-wide. A preliminary analysis shows the American
fauna to represent a plesiotypic level within Guignotites which can be interpreted as indicating
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
287
that Guignotites was present in South America before that continent separated from Africa.
Members of Guignotites occurring in North America would then represent a northward
extension of the Neotropical fauna after one of the many closures of the Panamanian portal.
This zoogeographic hypothesis will be tested as further knowledge is gained about phylogeny of
the subgenus. For instance, the hypothesis would be disproved if the North American
Guignotites are not closely related to the South American Guignotites or if the South American
Guignotites are not representative of a gondwanian colonization of South America.
If the common ancestor of H. bimarginatus and H. rimosus did enter North America after
the closure of the Panamanian portal then they probably represent a vicariant species pair.
Both species have an affinity for forest-pond situations and therefore a vicariant event such as
development of a grassland barrier between the southeastern United States ( H . bimarginatus)
and Mexico ( H . rimosus ) in the Miocene (Martin and Harrell, 1957; Rosen, 1978; and Allen
and Ball, 1980), could have brought about the speciation. This hypothesis would require that
the Caribbean islands were colonized secondarily after the speciation event. Another hypothesis
would be that speciation took place in Central America by means of one of the events described
by Rosen (1978) and that H. bimarginatus has colonized southeastern United States by means
of island-hopping across the Caribbean. This hypothesis seems less likely because of the
absence of H. bimarginatus from critical staging points such as Jamaica. The island-hopping is
invoked, however, to explain the presence of H. rimosus in Cuba although it is also absent from
Jamaica but it is present in other critical staging areas such as the Yucatan Peninsula. A
summary of the zoogeographic and phylogenetic relationships of North American Guignotites
is shown in Fig. 60.
ACKNOWLEDGEMENTS
We would like to express our appreciation to the members of RER’s examining committee:
Dr. M.K. Sears, Department of Environmental Biology, University of Guelph, and Dr. A.
Smetana, Biosystematics Research Institute, Agriculture Canada, Ottawa. The kindness of Dr.
P. Alayo, Academia de Ciencias, Havana, Cuba during RER’s visit there in January, 1976 was
most welcome. Translations of German by Dr. Smetana, French by Mr. J.T. Huber, and Polish
by Dr. H.J. Liu, presently at Alberta Environment, Vegreville, Alberta was very helpful.
Technical assistance with scanning electron microscope work was provided by P. Pearson, and
G. Subden, University of Guelph and G. Braybrook, University of Alberta. Mr. R. Hurst did a
professional job of the drawings. Mr. J. Scott, University of Alberta, has done his usual
admirable job in the production of plates. We would like to extend our sincere thanks to our
colleague, J.T. Huber, for searching for type specimens in various European museums.
Colleagues at both the Universities of Guelph and Alberta have given assistance in many
ways, notably their enthusiasm. Among them are: J.S. Ashe, Dr. G.E. Ball, A. Borkent, J.M.
Cumming, J.T. Huber, W.D. Husby, J.D. Lafontaine, S.A. Marshall, E. Maw, K. Shaw, and
N.D.G. White. The senior author is extremely grateful to his wife, Marsha, who provided
invaluable assistance throughout this study. We appreciated the time and effort of the people
who arranged the loans of specimens.
Financial support was provided in the form of Agriculture Canada grant #4033, and NRC
grant #A9552 held by D.H. Pengelly; additionally the senior author has been supported at the
University of Alberta by NSERC grant #A-1399 held by Dr. G.E. Ball.
Quaest. Ent., 1981, 17 (3,4)
288
Roughley and Pengelly
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Figs. 4 and 5. Genital appendages of male of Hydaticus ( Hydaticus ) modestus Sharp. Fig. 4. Ventral view. Fig.
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297
Figs. 10 to 14. Parts of the body of Hydaticus ( Hydaticus ) modestus Sharp. Fig. 10. Left metafemur and metatibia,
posterior surfaces. Fig. 1 1. Median lobe of aedoeagus, side view. Fig. 12. Median lobe of aedoeagus, ventral view of apex.
Fig. 13. Epipenite of aedoeagus. Fig. 14. Left paramere of aedoeagus. Figs. 15 and 16. Parts of the aedoeagus of Hydaticus
{Hydaticus) continentalis J. Balfour-Browne. Fig. 15. Epipenite. Fig. 16. Median lobe, side view.
Quaest. Ent., 1981, 17 (3,4)
298
Roughley and Pengelly
Figs. 17 and 18. Scanning electron micrographs of elytron of a female of Hydaticus ( Hydaticus ) modestus
Sharp. Fig. 17. Rugose elytron. Fig. 18. Sinuate apex of elytron.
Hunpxt F.nt 1981 17 H.4^
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
299
Fig. 19. Geographic distribution of certain elytral characteristics of selected samples of specimens of Hydaticus
(. Hydaticus ) modestus Sharp.
300
Roughley and Pengelly
Fig. 20. Distribution of Hydaticus ( Hydaticus ) modestus Sharp, within North America.
CiuaP.Rt.Fnt. 1981 17 ( 3 4^
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
301
Figs. 21 to 27. Parts of the body of Hydaticus ( Hydaticus ) piceus LeConte. Fig. 21. Right half of head. Fig.
22. Left half of body. Fig. 23. Left metafemur, and metatibia, posterior surfaces. Fig. 24. Median lobe of
aedoeagus, side view. Fig. 25. Median lobe of aedoeagus, ventral view of apex. Fig. 26. Epipenite of aedoeagus.
Fig. 27. Left paramere of aedoeagus.
302
Roughley and Pengelly
Fig. 28. Distribution of Hydaticus ( Hydaticus ) cinctipennis Aube, and Hydaticus ( Hydaticus ) piceus LeConte.
Hunoct Fnt 1QR1 17
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
303
Figs. 29 to 35. Parts of the body of Hydaticus ( Hydaticus ) cinctipennis Aube. Fig. 29. Right half of head. Fig.
30. Left half of body. Fig. 31. Left femur and metatibia, posterior surfaces. Fig. 32. Median lobe of aedoeagus,
side view. Fig. 33. Median lobe of aedoeagus, ventral view of apex. Fig. 34. Epipenite of aedoeagus. Fig. 35.
Left paramere of aedoeagus.
304
Roughley and Pengelly
Figs. 36 to 42. Parts of the body of Hydaticus ( Guignotites ) bimarginatus (Say). Fig. 36. Right half of head.
Fig. 37. Left half of body. Fig. 38. Left metafemur, and left metatibia, posterior surfaces. Fig. 39. Median lobe
of aedoeagus, side view. Fig. 40. Median lobe of aedoeagus, ventral view of apex. Fig. 41. Epipenite of
aedoeagus. Fig. 42. Left paramere of aedoeagus.
Quaest. Ent., 1981, 17 (3,4)
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
305
Figs. 43 to 45. Scanning electron micrographs of Hydaticus ( Guignotiies ) bimarginatus (Say). Fig. 43. Head
and pronotum of female. Fig. 44. Metatarsal claws of male. Fig. 45. Disc of metatibia, anterior surface, of
male, showing 13 large spines.
306
Roughley and Pengelly
!
Fig. 46. Distribution of Hydaticus ( Guignotites ) bimarginatus (Say), and Hydaticus ( Guignotites ) rimosus
Aube.
Ouaest. Ent.. 1981, 17 (3.4)
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
307
Figs. 47 to 54. Parts of the body of Hydaticus (Guignotites) rimosus Aube. Fig. 47. Right half of head. Fig. 48.
Left half of body. Fig. 49. Left half of body. Fig. 50. Left metafemur, and left metatibia, posterior surfaces.
Fig. 51. Median lobe of aedoeagus, side view. Fig. 52. Median lobe of aedoeagus, ventral view of apex. Fig. 53.
Epipenite of median lobe. Fig. 54. Left paramere of aedoeagus.
308
Roughley and Pengelly
Figs. 55 and 56. Scanning electron micrographs of Hydaticus ( Guignotites ) rimosus Aube. Fig. 55. Metatarsal
claws. Fig. 56. Disc of metatibia, anterior surface, showing 5 large spines.
Quaest. Ent., 1981, 17 (3,4)
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
309
Fig. 57. Reconstructed phylogeny of the higher taxa of Hydaticini. Open circles represent the plesiomorphic,
filled circles the apomorphic states presented in Table 4.
310
Roughley and Pengelly
/
Fig. 58. Reconstructed phylogeny of the species-group of Hydaticus ( Hydaticus ) which contains North
American taxa. Open circles represent the plesiomorphic, filled circles the apomorphic states presented in Table
4.
Quaest. Ent., 1981, 17 (3,4)
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
311
to
CK
lO
Fig. 59. Zoogeographic derivation of pattern of distribution of smooth and rugose forms of elytral sculpture of females of Hydaticus ( Hydaticus ) modestus Sharp. Fig. 59A. Maximum
distribution of glacial ice in North America at the peak of Wisconsin glaciation (after Nimmo, 1971, fig. 664). At this time the rugose form was within the Beringian refugium and the
smooth form in the eastern-deciduous forest refugium. Fig. 59B. Approximate ice positions at about 11,000 years before present (after Prest, 1969). During retreat of ice sheets the
smooth form dispersed westward whereas the Beringian population remained isolated. Later the Beringian, rugose form colonized central North America to achieve the present
distribution shown in Fig. 19.
312
Roughley and Pengelly
Fig. 60. Summary of the phylogenetic and zoogeographic relationships of North American Hydaticus. The
epipenite of the aedoeagus of the male is used as a pictorial representation of species. The subgenus Hydaticus
has probably had two separate invasions of North America and the subgenus Guignotites has possibly only
invaded once.
Quaest. Ent., 1981, 17 (3,4)
Hydaticus Leach (Coleoptera: Dytiscidae) of North America
313
Index to Names of Taxa
FAMILY GROUP TAXA
Aubehydrinae, 259
Carabidae, 281
Coleoptera, 257,281
Curculionidae, 257
Diptera, 254
Dytiscidae, 254, 255, 256
Dytiscinae, 281
Ephemeroptera, 254
Eulophidae, 255
Hydaticini, 258,259,281,285
Hymenoptera, 255
Odonata, 254
Trichoptera, 254
GENERA AND SUBGENERA
Dytiscus Linnaeus, 260
Guignotites Brinck, 255, 256, 257, 259,
269, 270, 282, 283, 284, 286, 287
Hydaticinus Guignot, 257, 283, 285
Hydaticus Leach, 250, 251, 254, 255, 256,
257, 258, 259, 265, 266, 267, 268, 270,
274, 283, 284
Hydaticus s. lat., 258,281,282,284
Hydaticus s. str., 270, 281, 282, 283, 284,
285,286
Hydaticus Schoenherr, 257
Isonotus Houlbert, 270
Pleurodytes Regimbart, 258, 282, 283,
284, 285
Prodaticus Sharp, 258, 284, 285
Thermonectus Dejean, 277
Trichopselaphus Chaudoir, 281
SPECIES AND SUBSPECIES
americanus Sharp,
Hydaticus, 260, 265
aruspex Clark, Hydaticus, 259, 260
bimarginatus LeConte,
Hydaticus, 287
bimarginatus Say, Dytiscus, 271
bimarginatus Say,
Hydaticus, 253,256,259,270,271,
273, 274, 275
bimarginatus Wickham,
Hydaticus, 260
cinctipennis Aube,
Hydaticus, 255, 256, 259, 265, 267,
268, 269, 270, 274, 282, 283, 284, 286
continentalis
Balfour-Browne,
Hydaticus, 260,265,267,281,282,
284
dineutoides Sharp,
Hydaticus, 281
dineutoides Sharp,
Pleurodytes, 258
dorsiger Aube, Hydaticus, 282
epipleuricus Regimbart,
Pleurodytes, 258
exclamationis Aube,
Hydaticus, 269,281,286
fasciatus Fabricius,
Dytiscus, 257
fasciatus Fabricius,
Sandracottus, 257
fasciatus Zimmermann,
Notaticus, 258, 259
flavolineatus Boheman,
Hydaticus, 281
fractivittis Guignot,
Hydaticus, 281,282
fulvicollis Aube, Hydaticus, 271
grammicus Germar,
Hydaticus, 277,281,282
histrio Clark, Hydaticus, 281
hybneri Fabricius, Dytiscus, 257
laevipennis Sharp,
Hydaticus, 260, 265
leander Rossi, Hydaticus, 281, 282, 284
matruelis Clark, Hydaticus, 281
modestus Sharp, Hydaticus, 253, 256,
259, 260, 265, 266, 267, 268, 269, 282,
283, 284, 286
palliatus Aube, Hydaticus, 281
piceous Gordon and Post,
Hydaticus, 267
piceus LeConte, Hydaticus, 253, 255,
314
Roughley and Pengelly
256, 259, 267, 268, 274, 284, 286
pictus Sharp, Hydaticus, 281
pictus Sharp, Prodaticus, 258, 281, 283
pullatus Guignot,
Hydaticus, 282
rectus Sharp, Hydaticus, 281
riehli Wehncke, Hydaticus, 277
rimosus Aube, Hydaticus, 255, 256, 260,
271,273,274, 275,287
rimosus Young, Hydaticus, 271
rugosus Poppius,
Hydaticus , 260, 265, 267
seminger (Degeer),
Hydaticus, 281,284
seminiger Degeer, Dytiscus , 257
semisulcatus O. Muller,
Dytiscus, 260
stagnalis Crotch,
Hydaticus , 260
stagnalis Fabricius,
Hydaticus, 260, 265, 267, 274
stagnalis Fourcrier,
Dytiscus , 260
subfasciatus LaPorte,
Hydaticus, 281
transversalis
(Ponloppidian),
Hydaticus, 281,284
transversalis Pontoppidian,
Dytiscus , 257
vittatus-group, Hydaticus, 281
Quaest. Ent., 1981, 17(3,4)
315
BOOK REVIEW
TSCHIRNHAUS, M. von 1981. Die Halm- und Minierfliegen im Grenzbereich Land-Meer
der Nordsee. Eine okologische Studie mit Beschreibung von zwei neuen Arten und neuen Fang-
und Konservierungsmethoden (Diptera: Chloropidae et Agromyzidae). Spixiana, Supplement
6. 405 pages containing 89 tables and 25 diagrams; 1 1 plates of photographs.
This work presents part of the results of a recently completed doctoral project commenced in
1965. The length of time required to complete the project is explained by the fact that over half
a million flies had to be identified. Tschirnhaus originally undertook a quantitative ecological
study. What he has produced is much more. For, unlike most quantitative ecologists, he has not
been content with unreliable species determinations or aggregation of his data according to
genera or broader taxonomic units. Identification problems encountered during this project
have prompted him to undertake the taxonomic research necessary to solve them. As a result he
has emerged not only as an outstanding ecologist but also as one of the leading taxonomic
specialists on the Agromyzidae and Chloropidae.
Tschirnhaus’ study sites include tidal salt marshes, dykes and brackish water reservoirs
along the west coast of Schleswig-Holstein and Denmark. Comparisons are made between
grazed and ungrazed land, as well as between different plant communities. The sampling
methods include the rearing of adults from collections of larvae and from vegetation samples
isolated in the laboratory, and field trapping with sweep nets, coloured trays, emergence traps
(“photoeclectors”) and wind funnels.
Since coloured tray traps have not been used before in quantitative studies of small Diptera,
this technique and the interpretation of its results are discussed at length. A new technique for
producing microslides for preserving the genitalia of taxonomic specimens is also described.
This involves use of a viscous embedding medium (“Zeiss W 15”) which is enclosed in a rigidly
hardening medium called “Eukitt”. This technique should be tried by workers on very small
insects whose genitalia are too easily lost when stored in microvials.
The sampling data has been analysed from the following synecological aspects: flight season,
dominance dynamics, comparative abundance in coastal and inland regions, distribution
according to habitat type, dispersal intensity, sex ratio, activity level and population size. The
comparison with inland regions discloses some results of the author’s studies in Norway and the
Alps, as well as at inland localities in Schleswig-Holstein. The analysis of dispersal also uses
data from three lightships moored in the North and Baltic Seas.
Surprising differences between species in all the above parameters are revealed by
Tschirnhaus’ analysis, which thus demonstrates the inadequacy of the usual ecologists’
approach of sorting samples only to broad taxonomic units. For instance, the colour tray data
show that uniform colour preferences cannot be assumed for closely related species or even for
the sexes of the same species. Likewise there is no uniformity in activity response to weather
changes, a few species showing unexpected behaviour such as increasing their flight activity
under steady rain. Of particular interest to biogeographers is the demonstration that dispersal
of these small flies in “aeroplankton” has nothing to do with the effects of storms, but is a
voluntary phenomenon occurring during fine summer weather among species whose population
densities have become high. Species which did not achieve high population densities were rarely
taken far from their breeding habitat, no matter how wind exposed.
Following his synecological discussions Tschirnhaus presents taxonomic and autecological
treatments of ten species of agromyzids whose larvae feed on halophytes (namely Aster
tripolium, Artemisia maritima, Atriplex spp., Triglochin spp. and Juncus gerardii). These
316
treatments are extremely detailed and include a complete critical review of previous literature. ,
Two of the fly species ( Napomyza maritima, a stem-borer in the Artemisia , and Metopomyza
junci , a leaf-miner on the Juncus ) are new to science. The description of the new Metopomyza j
is preceded by a discussion of other species of this little studied genus, in the course of which an
expanded description is given of the Canadian M. griffithsi Sehgal (Edmonton, known only
from the holotype).
Evidently this publication still presents only a small part of the new taxonomic information
accumulated during Tschirnhaus’ studies, since we are promised further publications describing
eight more new agromyzids and four new chloropids listed as nomina nuda. It is to be hoped
that these further publications will be available as soon as possible, since it is not satisfactory to
have published information associated with nomina nuda.
Tschirnhaus’ work needs to be brought to the attention of Canadian entomologists because
of its possible relevance to the survey of prairie insects which the Biolgical Survey of Canada is
planning. His sampling methods have been developed for exclusively herbaceous vegetation
(some tall, some very short due to heavy grazing), physically comparable with that of our
praireis. So his conclusions on the usefulness and comparability of various methods are relevant
to the design of any quantitative sampling for the prairie insect survey. I recommend those
involved in organizing the survey to read Tschirnhaus’ work.
I am confident that this publication will become recognized as a classical work of its kind,
one which will set the standard for future ecological studies on phytophagous Diptera. Well
done!
Graham C.D. Griffiths
Department of Entomology
The University of Alberta
Edmonton, Alberta T6G 2E3
317
EDITOR’S ACKNOWLEDGEMENTS
Just about one year ago, I expressed hope that we would be able to complete Volume 17 of
j Quaest. Ent. in 1981. In spite of sustained efforts to do so, we missed our target by six months,
j Nontheless, our contribution for 1981 is now complete and the first issue of Volume 18 is well
underway. Perhaps we will manage to complete this one in the appropriate year.
Until we began to use computers in production of the journal, issues were always just about
on time. I suppose that appearance of these complex machines coincided with an increase in
complexity of our world, so that without computers, we would not be able to cope, at all. Put in
another way, I doubt that use of computers is the causative factor of delay in publication of
Quaest. Ent., or of the many other scientific journals that seem to be having problems similar to
ours. In due time, perhaps computers will enable us to deal effectively with the complexities of
existence with which we are now besieged.
Be that as it may, I am indebted to the following for their prompt assistance in reviewing the
manuscripts that appeared in volume 17:
D. A. Craig, Department of Entomology, University of Alberta;
R. H. Gooding, Department of Entomology, University of Alberta;
C. W. Hogue, Los Angeles County Museum, Los Angeles, California;
R. J. Mackay, Department of Zoology, University of Toronto, Toronto, Ontario;
D. Rosenberg, Freshwater Institute, Environment Canada, University of Manitoba,
Winnipeg, Manitoba;
J. A. Shemanchuk, Research Station, Agriculture Canada, Lethbridge, Alberta; and
J. R. Spence, Department of Entomology, University of Alberta.
I also record my appreciation to members of my Department who assisted with the
publishing process. Jean-Francois Landry provided French translation of several abstracts.
John S. Scott helped with illustrative matters, and he, Danny Shpeley, Robert S. Anderson, and
Andrew P. Nimmo read proof, as required.
Ms. Suseela Subbarao, now in her second year as Publications Manager, performed
admirably the many tasks that this position entails. Her patience, knowledge, and skill are
much appreciated.
G. E. Ball
Ecology of four Patrobus species
319
INDEX
Aechlimann, J.P., 170
Aedes, 149, 152, 155, 156, 157, 158, 159,
160, 161, 165, 166
Aedes (Aedes) cinereus
Meigen, 160, 166
Aedes ( Aedimorphus )
vexans (Meigen), 158,159,165,166
Aedes (Ochlerotatus)
campestris Dyar and
Knab, 157, 158, 159, 165
Aedes ( Ochlerotatus )
canadensis (Theobald), 155, 156, 162
Aedes ( Ochlerotatus )
cataphylla Dyar, 156, 157, 158, 159,
165
Aedes ( Ochlerotatus )
communis (de Geer), 155, 156, 162,
165
Aedes ( Ochlerotatus )
diantaeus Howard,
Dyar and Knab, 160
Aedes ( Ochlerotatus )
dorsalis (Meigen), 158,159,166
Aedes ( Ochlerotatus )
euedes Howard, Dyar
and Knab, 148, 149, 152, 157, 158,
162, 165
Aedes ( Ochlerotatus )
excrucians (Walker), 148,158
Aedes ( Ochlerotatus ) fitchii
(Felt and Young), 157,158,165
Aedes (Ochlerotatus)
flavescens (Muller), 159,165
Aedes (Ochlerotatus)
hexodontus Dyar, 156,162
Aedes (Ochlerotatus)
impiger (Walker), 160
Aedes (Ochlerotatus)
implicatus Vockeroth, 157, 160, 162
Aedes ( Ochlerotatus )
increpitus Dyar, 157,162
Aedes (Ochlerotatus)
intrudens Dyar, 160
Aedes (Ochlerotatus)
melanimon Dyar, 159
Aedes (Ochlerotatus)
mercuratorDyav, 148,149,152,157,
158, 160, 162, 165
Aedes (Ochlerotatus)
nigromaculis (Ludlow), 160
Aedes (Ochlerotatus)
pionips Dyar, 162
Aedes (Ochlerotatus)
pullatus Coquillett, 156,162,165,
166
Aedes (Ochlerotatus)
punctorYArby, 156,162
Aedes (Ochlerotatus)
schizopinax Dyar, 156,162
Aedes (Ochlerotatus)
spencerii (Theobald), 159, 165
Aedes (Ochlerotatus)
sticticus (Meigen), 160
Aedes (Ochlerotatus)
trichurus (Dyar), 160,162
Aedes barri Rueger, 148, 162
Aedes beklemishevi
Denisova, 148
Aedes cataphylla Dyar, 1 8 1
Aedes fitchii (Felt and
Young), 181, 182
Aedes Meigen, 181
Aedes riparius Dyar and
Knab, 148, 162
Aedes stimulans (Walker), 149, 158, 162
Aedes vexans (Meigen), 181, 182
Agonum Bon., 215,216
Agonum cupreum Dej., 215
Agonum placidum Say, 215
Agrotis ipsilon (Hiifngel), 216
alaskaensis (Ludlow),
Culiseta (Culiseta), 161
albertae Kevan and
Cutten-Ali-Khan,
Oreadomyia , 121, 125, 126, 127
Allan, J.D., 143
Amara apricaria Payk., 214
Amara Bon., 214,216
Quaest. Ent. 1981, 17 (3,4)
320
Carter
INDEX
Amara carinata Lgc., 213,214,215
Amarafracta Lee., 213,214
Amara lacustris Lee., 213,214
Amara latoir Kby., 214
Amara littoralis Mnh, 214
Amara obesa Say, 213,214
Amara quenseli Schnh., 213,214
Amara torrida Panz., 214
amplus (Thaver), Baetis , 130
amputatus Say, Harpalus, 214
Anaphes pratensis Foerster, 169, 170
Anopheles , 161
Anopheles ( Anopheles )
ear lei Vargas, 161
antarctica (Jacobs),
Belgica , 125
apricaria Payk., Amara, 214
aquatilis Wahlenb., Carex , 236
arcticum Malloch,
Simulium , 189, 191, 192, 198, 199,
204, 206, 207, 208, 209
assimilis Chaudoir,
Patrobus, 239
atherodes Spreng., Carex , 236
atrorufus Stroem, Patrobus , 239
Aubin, A.,
see Maire, A., 165
Baetidae, 130
flaef/s, 130,138,142,143
Baetis alpinus, 138
Baetis amplus (Thaver), 130
Baetis bicaudatus Dodds, 130, 133, 136,
138, 142, 143
Baetis foemina
McDunnough, 130
Baetis hudsonicus Ide, 130
Baetis insignificans
McDunnough, 130
Baetis lapponicus
(Bengtsson), 130, 138
Baetis lapponicus group, 1 30
Baetis lutheri, 138
Baetis macani bundyae , 138
Baetis rhodani group, 130, 138
Baetis tricaudatus Dodds, 131, 133, 136,
138, 142, 143
Baetodes Needham and
Murphy, 130
Ball, G.E., 235,241
barri Rueger, Aedes, 148, 162
Beegle, C.C.,
see Best, L.R., 216
beklemishevi Denisova,
Aedes , 148
Belgica , 125
Belgica antarctica (Jacobs), 125
Bembidion bimaculatum
Kby., 213
Bembidion Latr., 213, 214, 216
Bembidion nitidum Kby., 213, 216
Bembidion obscurellum
Mtsch., 213,214,216
Bembidion oppositum Say, 213, 214, 216
Bembidion rupicola Kby., 213
Bembidion timidum Lee., 213, 216
Berner, L.,
see Edmunds, G.E., Jr., 130, 138, 143
see Ward, J.V., 143
Best, L.R., 216
bicaudatus Dodds, Baetis , 130, 133, 136,
138, 142, 143
bimaculatum Kby.,
Bembidion , 213 i
bivittatum Malloch,
Simulium , 191, 192, 198, 199, 202,
206
Borkent, A., 127
Bowman, M.C.,
see Widhaas, D.E., 183
brassicae (Bouche),
Hylemya, 216
Brown, A. W. A., 182
Brundin, L., 126
Brust, R.A.,
see Ellis, R. A., 155
Buchonomyiinae, 126
Ecology of four Patrobus species
321
Bunn, R.W.,
see Hamlin, J.C., 169,170
Burgess, L., 159
Cameron, R.S.,
see Johnson, N.E., 216
Campbell, C.J.,
see Finlayson, D.G., 212
campestris Dyar and Knab,
Aedes (Ochlerotatus), 157, 158, 159,
165
canadensis (Theobald),
Aedes (Ochlerotatus), 155, 156, 162
Carex, 236
Carex aquatilis Wahlenb., 236
Carex atherodes Spreng., 236
Carex rostrata Stokes, 236
carinata Lee., Amara , 213, 214, 215
Carpenter, S.J., 148, 152, 156, 157, 159,
160, 181
Carter, A., 237,240
cataphylla Dyar, Aedes, 1 8 1
cataphylla Dyar, Aedes
( Ochlerotatus ), 156,157,158,159,
165
Chamberlain, T.R., 169,170
Chironomidae, 121, 126, 127
Chironominae, 121
Christie, G. I., 169
cinereus Meigen, Aedes
(Aedes), 160, 166
Clausen, C.P., 169,170
Clifford, H.F., ,131
see Corkum, L.D., 130
Cobb, D.G., 138
cognatus Gyll.,
Trichocellus, 216
Cole, L.E., 236, 237
communis (de Geer), Aedes
(Ochlerotatus), 155, 156, 162, 165
Cook, E.F.,
see Hansen, D.C., 126
Corkum, L.D., 130
INDEX
Coulson, J.R.,
see Stone, A., 191
Ctenicera destructor
(Brown), 216
Culex, 161
Culex (Culex) tarsalis
Coquillett, 161
Culex (Neoculex) territans
Walker, 161
Culiseta, 161, 165
Culiseta (Culicella)
moristans (Theobald), 162
Culiseta (Culiseta)
alaskaensis (Ludlow), 161
Culiseta (Culiseta) incidens
(Thomson), 161
Culiseta (Culiseta) inornata
(Williston), 161
cupreumDey, Agonum, 215
Cutten-Ali-Khan, F.E.A.,
see Kevan, D.K.McE., 121, 125, 126
Cutten, E.A., 126
Dang, P.T.,
see Wood, D.M., 152
Danilov, V.N., 149
Darlington, P.J. Jr., 235, 236, 239, 240,
241
Davies, D.M., ,191,205
see also Wood, D.M., 191
Davies, L., 191
Dawson, N., 238
Day, W.H.,
see Dysart, R.J., 169
decorum Walker,
Simulium, 191, 193, 198, 199, 204,
208
destructor (Brown),
Ctenicera , 216
Diamesinae, 121, 126, 127
diantaeus Howard, Dyar
and Knab, Aedes
(Ochlerotatus), 160
Dicercomyzon, 130
Diplocladius, 126
Quaest. Ent. 1981, 17 (3,4)
'322
Carter
Diptera, 126, 182
Doane, J.F., ,212,216
see also Wishart, G., 216
Dodds, G.S., 130,143
Dondale, C.D.,
see Doane, J.F., 212
dorsalis (Meigen), Aedes
( Ochlerotatus ), 158, 159, 166
Downes, J. A., 126
Drenkelfort, H., 131,143
Dritschilo, W., 212
duplex Shewed and
Fredeen, Simulium, 191,192,193,
199, 202, 205
Dyar, H.G., 148,162
Dysart, R.J., 169
earlei Vargas, Anopheles
( Anopheles ), 161
Ectemnia taeniatifrons
(Enderlein), 191, 193, 198, 202, 204
Eddy, G.H.,
see Gjullin, C.M., 152
Edmunds, G.E., Jr., 130,138,143
Ellis, R. A., , 155
see Wood, D.M., 152
elongatus (Hagen),
Isogenoides, 131, 136
Elton, C.S., 236
236
Enfield, M.A., 148, 149, 157, 158, 160,
162, 165, 166
Epeorus Eaton, 130
Esau, K.L., 212
Essig, E.O., 169
euedes Howard, Dyar and
Knab, Aedes
( Ochlerotatus j, 148,149,152,157,
158, 162, 165
euryadminiculum Davies,
Simulium , 191,192,193,199,202,
204, 205
Euxoa detersa group, 1-113
Index of names, 115-120
Euxoa ochrogaster Guenee, 2 1 6
INDEX
excrucians (Walker), Aedes
( Ochlerotatus J, 148, 158
Felicitomyia, 125
Finlayson, D.G., 212
fitchii (Felt and Young),
Aedes , 181, 182
fitchii (Felt and Young),
Aedes ( Ochlerotatus ), 157, 158, 165
Flannagan, J.F.,
see Cobb, D.G., 138
flavescens (Muller), Aedes
( Ochlerotatus ), 159, 165
foemina McDunnough,
Baetis, 130
Foote, R.H.,
see Stone, A., 191
Ford, H.R.,
see Weidhaas, D.E., 183
Forsskahl, B., 239
fossifrons Eschscholtz,
Patrobus , 236, 240
foveocollis Eschscholtz,
Patrobus , 236, 237, 238, 239, 240,
241
Fowler, H.W., Jr.,
see Horsfall, W.R., 166
Fox, C.J.S., 212,216
fracta Lee., Amara , 213, 214
Frank, J.H., 212,213,214,215,216
Fredeen, F.J.H., , 206, 207, 208, 209
see also Shewed, G.E., 204, 205
Gillett, J.D., 166
Gjullin, C.M., 152
Gooding, R.H.,
seeTawfik, M.S., 148,162
Graham, P., 162,236
Greenslade, P.J.M., 237, 239
Gressitt, J.L.,
see Wirth, W.W., 125
griseum Coquillett,
Simulium , 191, 192, 198, 199, 202,
206
Ecology of four Patrobus species
323
INDEX
Gyorkos, Helen,
see Wood, D.M., 191
Hackman, W., 126
hageni Eaton, Rhithrogena, 131, 136,
138, 142
Hagley, E.A.C.,
see Holliday, N.J., 212
Hamlin, J.C., 169,170
Hansen, D.C., 126
Happold, D.C.D., 148,160
Harpalus amputatus Say, 214
Harpalus herbivagus Say, 213, 214
Harpalus Latr., 214, 216
Harpalus pleuriticus Kby., 214
Hashimoto, E., 126
Hawkes, H.A., 143
Heptageniidae, 130
herbivagus Say, Harpalus , 213, 214
Hesperoperla pacifica
(Banks), 133, 138, 142
Heterocloeon
McDunnough, 130, 138
hexodontus Dyar, Aedes
(Ochlerotatus), 156, 162
Hisaw, F.L., , 143
see Dodds, G.S., 130, 143
Hobbs, G. A., 169
Hocking, B., , 180, 181, 182, 183
see also Klassen, W., 180, 182
Holliday, N.J., 212
Hora, S.L., 130, 143
Horsfall, W.R., 166
Hudson, J.E.,
see Hocking, B., 180, 182, 183
h udsonicus Ide, Baetis, 130
Hughes, D. A., 130
Hydaticus Leach, 251-317
Index of names, 319-320
Hylemya bras sic ae
(Bouche), 216
Hymenoptera, 182
Hynes, H.B.N., 130,142
Hyper a, 170
Hypera nigrirostris (¥ .), 169
Hypera postica (Gyll.), 169,170
Hypera punctata ( Fabr.), 170
Ide, F.P., 143
Idema, R., 127
impiger (Walker), Aedes
( Ochlerotatus ), 160
implicatus Vockeroth,
Aedes ( Ochlerotatus ), 157, 160, 162
incidens (Thomson),
Culiseta (Culiseta), 161
increpitus Dyar, Aedes
( Ochlerotatus j, 157, 162
inornata (Williston),
Culiseta (Culiseta), 161
insignificans McDunnough,
Baetis, 130
intrudens Dyar, Aedes
( Ochlerotatus ), 160
ipsilon (Hiifngel), Agrotis, 216
Ironodes T raver, 1 30
Isogenoides elongatus
(Hagen), 131, 136
Jensen, S.L.,
see Edmunds, G.E., Jr., 130, 138, 143
Johnson, N.E., 216
Jones, L.J.,
see Hamlin, J.C., 169,170
Kevan, D.K.McE., , 121, 125, 126
see also Cutten, E.A., 1 26
Kirk, V.M., 212,214,215
Klassen, W., 180,182
Kuhnelt, W., 239
La Roi, G.H., 236
LaCasse, W.J.,
see Carpenter, S.J., 148,152,156,
157, 159, 160, 181
Lachlania Hagen, 130
lacustris Lee., Amara, 213, 214
Landau, R., 208
lapponicus (Bengtsson),
Baetis, 130, 138
lapponicus group, Baetis, 1 30
Quaest. Ent. 1981, 17 (3,4)
324
Carter
Larsen, J.R.,
see Horsfall, W.R., 166
Larsson, Sv.G., 239
latifolia L., Typha, 236
latoirKby., Amara, 214
lecontei Chaudoir,
Patrobus, 236, 237, 238, 239, 240
Lehmkuhl, D.M, 138
Lepidoptera, 182, 1-120
Leucocytozoon, 205, 206
Lewis, T., 215
Lieberman, F.V.,
see Hamlin, J.C., 169, 170
Lindroth, C.H., 235, 236, 237, 239, 240
littoralis Mnh, Amara , 214
Lofgren, C.S.,
see Weidhaas, D.E., 183
longicornis Say, Patrobus , 239, 240
lucublandus Say,
Pterostichus, 213,214,215,216
luggeri Nicholson and
Mickel, Simulium, 189,191,193,
198, 199, 202, 206, 207, 208, 209
luna (Girault), Patasson , 169, 170
macani bundyae, Baetis, 138
MacArthur, R.H., 215
MacLellan, C.R.,
see Fox, C.J.S., 212,216
Magnarelli, L.A., 166
Mailhot, Y., 165
Maire, A., , 165
see also Mailhot, Y. , 165
Maslin, T.P., 240
Maybee, G.E.,
see Wishart, G., 216
Mayr, E., 239
McAlpine, J.F., 127
McAlpine, J.F., 191
McCafferty, W.P.,
see Morihara, D.K., 130, 138
McDuffe, W.c.,
see Hamlin, J.C., 169,170
McDunnough, J., 138
Mclver, S., 148
INDEX
Mclver, S.B., 143
melanimon Dyar, Aedes
(Ochlerotatus), 159
mercurator Dyar, Aedes
( Ochlerotatus ), 148, 149, 152, 157,
158, 160, 162, 165
meridionale Riley,
Simulium , 191, 192, 193, 199, 202,
205
Metacnephia
saskatchewana Shewell
and Fredeen, 191, 193, 199, 202, 204
Michelbacher, A.E.,
see Essig, E.O., 169
Miller, R.S.,
Moretti, L.J.,
see Horsfall, W.R., 166
Morihara, D.K., 130,138
moristans (Theobald),
Culiseta (Culicella), 162
Moss, E.H., 236
Mount, G.A.,
see Weidhaas, D.E., 183
Miiller-Liebenau, I., 138
Murdoch, W.W., 238, 239
Naturalists, 177
Neopatrobus, 240, 24 1
Newton, R.C.,
see Hamlin, J.C., 169, 170
Nielsen, L.T., 156
nigrirostris (F.), Hypera, 169
nigromaculis (Ludlow),
Aedes (Ochlerotatus), 160
Nimmi, W.O.,
see Hobbs, G. A., 169
nitidum Kby., Bembidion, 213, 216
Noctuidae, 1-120
Nymphomyiidae, 121, 125
obesa Say, Amara, 213,214
obscurellum Mtsch.,
Bembidion, 213,214,216
ochrogaster Guenee,
Euxoa, 216
Odontomesa, 127
Ecology of four Patrobus species
325
INDEX
Oligoneuriidae, 130
oppositum Say, Bembidion, 213, 214, 216
Oreadomyia albertae Kevan
and Cutten-Ali-Khan, 121,125,126,
127
Orthocladiinae, 121, 126, 127
pacifica (Banks),
Hesperoperla, 133, 138, 142
Palaeodipteron, 125
Pat as son, 170
Patasson luna (Girault), 169, 170
Patrobus , 236, 237, 238, 239, 240
Patrobus assimilis
Chaudoir, 239
Patrobus atrorufus Stroem, 239
Patrobus fossifrons
Eschscholtz, 236, 240
Patrobus foveocollis
Eschscholtz, 236, 237, 238, 239, 240,
241
Patrobus lecontei Chaudoir, 236, 237,
238, 239, 240
Patrobus longicornis Say, 239, 240
Patrobus septentrionis
Dejean, 236, 237, 238, 239, 240, 241
Patrobus stygicus
Chaudoir, 236, 237, 238, 239, 240
Percival, E., 130
Permanent International
File of Naturalists, 177
Peters, D.C.,
see Esau, K.L., 212
Peterson, B.V.,
see Davies, D.M., 191,205
see McAlpine, J.F., 191
see Wood, D.M., 191
Philipson, G.N., 143
Phillips, V.,
see Gillett, J.D., 166
Phosterodores, 207
Picard, L.,
see Maire, A., 165
Pielou, E.C., 212,213
PIFON, ,
see Permanent
International File of
Naturalists,, 177
pionips Dyar, Aedes
(Ochlerotatus), 162
placidum Say, Agonum, 215
pleuriticus Kby., Harpalus, 214
Popovici-Baznosanu, A., 143
postica (Gyll.), Hypera , 169, 170
pratensis Foerster,
Anaphes, 169, 170
Pritchard, G., 149, 158, 160, 165, 166
Prodiamesinae, 121, 126, 127
Pseudocloeon Klapalek, 130
Pterostichus Bon., 216
Pterostichus lucublandus
Say, 213,214,215,216
Pucat, A., 148,160,162
pullatus Coquillett, Aedes
(Ochlerotatus), 156, 162, 165, 166
punctata (Fabr.), Hypera , 170
punctor Kirby, Aedes
( Ochlerotatus ), 156, 162
quenseli Schnh., Amara, 213, 214
Rempel, J.G., 148,158,160,162
Rheobaetis , 138
Rhithrogena hageni Eaton, 131, 136, 138,
142
rhodani group, Baetis, 130, 138
riparius Dyar and Knab,
Aedes, 148, 162
Rivard, I., 212,215
Rohlf, F.J.,
see Sokal, R.R., 238
Roman, E.A.,
see Gillet, J.D., 166
rostrata Stokes, Carex , 236
Rothfels, K.H., 208
rugglesi Nicholson and
Mickel, Simulium, 191,192,193,
199, 202, 205,206
rupicola Kby Bembidion, 213
Quaest. Ent. 1981, 17 (3,4)
326
Carter
INDEX
Sabrosky, C.W.,
see Stone, A., 191
Saether, O.A., , 121, 125, 126, 127
see also Brudin, L., 126
see also Brundin, L., 126
Salix, 236, 237, 238
saskatchewana Shewed and
Fredeen , Metacnephia, 191, 193, 199,
202, 204
schizopinax Dyar, Aedes
(Ochlerotatus), 156, 162
Schmidt, H., 131,143
Scholefield, P.J., 149,162,165
sensu Ambiihl, 143
septentrionis Dejean,
Patrobus , 236, 237, 238, 239, 240,
241
Shemanchuk, J.A., 161,162
Shewed, G.E., , 204, 205, 206
see also McAlpine, J.F., 191
Simulium arcticum
Malloch, 189, 191, 192, 198, 199, 204,
206, 207, 208, 209
Simulium bivittatum
Malloch, 191, 192, 198, 199, 202, 206
Simulium decorum Walker, 191, 193,
198, 199, 204, 208
Simulium duplex Shewed
and Fredeen, 191, 192, 193, 199, 202,
205
Simulium
euryadminiculum
Davies, 191, 192, 193, 199, 202, 204,
205
Simulium griseum
Coquillett, 191, 192, 198, 199, 202,
206
Simulium luggeri
Nicholson and Mickel, 189, 191, 193,
198, 199, 202, 206, 207, 208, 209
Simulium meridionale
Riley, 191, 192, 193, 199,202,205
Simulium rugglesi
Nicholson and Mickel, 191, 192, 193,
199, 202, 205, 206
Simulium tuberosum
(Lundstroem), 191, 192, 198, 199,
202, 208, 209
Simulium venustum Say, 191, 193, 198,
199, 204, 208,209
Simulium verecundum
Stone and Jamnback, 191,193,198,
199, 204, 209
Simulium vittatum
Zetterstedt, 191, 192, 193, 198, 199,
202, 206
Sitona, 170
Sleeper, S.E.,
see Thomas, D.S., 212
Sneath, P.H.A., 133
Snoddy, E.L.,
see Stone, A., 191
Sokal, R.R., , 238
see Sneath, P.H.A., 133
Southwood, T.R.E., 237
spencerii (Theobald), Aedes
( Ochlerotatus ), 159, 165
Spinadis Edmunds and
Jensen, 130
sticticus (Meigen), Aedes
( Ochlerotatus j, 160
stimulans (Walker), Aedes, 149, 158, 162
Stone, A., 191,207
Strickland, E.H., 148, 155, 160, 161, 162
stygicus Chaudoir,
Patrobus, 236, 237, 238, 239, 240
taeniatifrons (Enderlein),
Ectemnia, 191, 193, 198, 202, 204
tarsalis Coquillett, Culex
(Culexj, 161
Tawfik, M.S., 148,162
territans Walker, Culex
( Neoculex ), 161
Teskey, H.J.,
see McAlpine, J.F., 191
Tessier, C.,
see Maire, A. , 165
Thiele, H.U., 239
Ecology of four Patrobus species
327
INDEX
Thomas, D.S., 212 162, 191
timidum Lee., Bembidion, 213, 216 see also Davies, D.M., 191, 205
Titus, E.G., 169 see also McAlpine, J.F., 191
torrida Panz., Amara, 214
tricaudatus Dodds, Baetis , 131, 133, 136,
138, 142, 143
Trichocellus cognatus G yll., 216
trichurus (Dyar), Aedes
(Ochlerotatus), 160, 162
Tricorythidae, 130
tuberosum (Lundstroem),
Simulium, 191, 192, 198, 199, 202,
208, 209
Turner, F.B., 212
Typha latifolia L., 236
van Emden, H.F., 215
Venter, G.E., 130
venustum Say, Simulium , 191, 193, 198,
199, 204, 208,209
verecundum Stone and
Jamnback, Simulium, 191, 193, 198,
199, 204, 209
vexans (Meigen), Aedes , 181, 182
vexans (Meigen), Aedes
(Aedimorphus), 158, 159, 165, 166
Virostek, J.,
see Hobbs, G.A., 169
vittatum Zetterstedt,
Simulium , 191, 192, 193, 198, 199,
202, 206
Vockeroth, J.R.,
see McAlpine, J.F., 191
Wada, Y., 148, 162
Wanner, D.,
see Dritschilo, W., 212
Ward, J.V., 143
Weidhaas, D.E., 183
Whitehead, H.,
see Percival, E., 130
Wirth, W.W., , 125
see Stone, A., 191
Wishart, G., 216
Wood, D.M., 127
Wood, D.M., , 148, 149, 152, 157, 160,
Quaest. Ent. 1981, 17(3,4)
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Quaestiones
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A periodical record of entomological investigations,
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QUAEST10NES ENTOMOLOGICAE ISSN 0033-5037
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6»
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Quaestiones
Entomologicae
MW 1 rij j/.. ■
// ° ‘ ’ t'
/
FEB 2 . 1983
\
V
A periodical record of entomological investigations,
published at the Department of Entomology,
University of Alberta, Edmonton, Canada.
VOLUME 18
NUMBERS 1-4
JANUARY-OCTOBER 1 982
CORRIGENDA-Qwaesfiones Erttomologicae, Volume 18
Hilchie, G. J. (1982, 18: 91-126).- Evolutionary aspects of Geographical Variation in Color and
of Prey in the Beewolf Species Philanthus albopilosus Cresson.
pages
106-109
Tables 6 and 7
The headings are transposed for columns indicating sex of “Specimens collected”. Thus,
the numbers in the column to the left refer to females, and those in the column to the right
refer to males.
THEORIES OF SEXUAL DIFFERENCE: THE SEXUAL SELECTION HYPOTHESIS
AND ITS ANTECEDENTS, 1786 - 1919
R.B. Aiken'
Department of Zoology, Erindale College
University of Toronto, Mississauga, Ontario, L5L 1C6
CANADA Quaestiones Entomologicae
18:1-14 1982
“ There is one great difficulty with a good hypothesis. When it is complete and rounded and the corners smooth
and the content cohesive and coherent, it is likely to become a thing in itself, a work of art. It is then like a finished
sonnet or a painting completed. One hates to disturb it. Even if subsequent information should shoot a hole in it, one
hates to tear it down because it was once beautiful and whole.”
John Steinbeck, 1951 (p. 180)
ABSTRACT
Since Hunter’s recognition of primary and secondary sexual characters, biologists have
offered explanations for the question of why males and females of many taxa exhibit striking
structural differences. Natural theologians of the early nineteenth century considered such
differences to represent the intricacy of some creative power. Darwin’s early writing about
sexual selection emphasized intrasexual selection and later incorporated the key ideas of
imbalanced sex ratio ( especially in polygamous animals) and female choice. Darwin’s chief
opponent, A.R. Wallace, grew away from the sexual selection argument and proposed the
vitalistic notion of greater male vigour and that secondary sexual differences were a result of
females needing protective colouration. Critics also attacked female choice as a selective agent
because it implied some aesthetic sense of females and because it was not seen as providing a
sufficiently consistent selective force to act in any direction. The ideas of three such critics, the
Duke of Argyll, St. George Mivart and J. Stolzmann, are examined.
RESUME
Depuis I’epoque ou Hunter reconnut les caracteres sexuels primaires et secondaires, les biologistes ont tente
d’expliquer de plusieurs f aeons le pourquoi des differences structurales souvent frappantes entre males et femelles. Au
debut du XIX,eme siecle, les theologiens de la nature consideraient que de telles differences refletaient la complexity d’un
certain pouvoir createur. Les premiers ecrits de Darwin sur la selection sexuelle insisterent sur la selection intrasexuelle,
et subsequemment incorporerent les idees capitales de proportions inegales des sexes (particlierement chez les animaux
poly games) et de choix des males par les femelles. Le principal adversaire de Darwin, A.R. Wallace, se detacha du debat
entourant la selection sexuelle; il emit I’hypothese que les differences sexuelles secondaires resultent de la necessity pour
les femelles d’avoir une coloration protectrice, et proposa la notion vitaliste de vigueur plus grande chez les males. Les
critiques attaquerent egalement I’idee que le choix des femelles constitue un agent selecteur parce qu'elle implique une
certaine notion d’esthetique de la part des femelles, et parce que ce choix n’etait pas percu comme pouvant exercer une
force selective suffisamment constante pour agir d’une facon directionnelle. Les idees de trois de ces critiques, le due
d' Argyll, Saint-George Mivart, et J. Stolzmann, sont examinees.
'Present address: Department of Entomology, University of Alberta, Edmonton, Alberta,
Canada T6G 2E3
2
Aiken
THE THEORY OF SEXUAL SELECTION
By 1871, Charles Darwin could have, quite easily, retired gracefully, his scientific
reputation established beyond any doubt. He could have abandoned his work of theorizing
about grand schemes of nature and devoted himself to his more restricted studies of
earthworms, orchids and moving plants. He had successfully weathered the storm (in terms of
scientific acceptance) over his theories of descent with modification, having been the focus of
more criticism and debate in a decade than most other scientists would encounter in several
lifetimes. Yet, in 1871, he published ‘The Descent of Man and Selection in Relation to Sex’,
probably not without some inkling of the debates his work would rekindle.
In the Descent, Darwin tried to tie off one of the more annoying loose ends of the Origin of
Species - a theory about the evolution of secondary sexual characters. In retrospect, the title of
the book has proven to have laid the wrong emphasis. By far the greatest amount of controversy
and scientific discussion has revolved not around the schema for the evolution of man but rather
the process by which he envisioned the races of man to have differentiated - sexual selection.
Sexual selection was Darwin’s attempt to explain one of the thorniest problems of animal
morphology - the varied and often bizzare structures, colours and actions that distinguish the
sexes. The theory of sexual selection as it appeared in 1871 (earlier versions of it will be
discussed below) goes somewhat as follows.
In sexually reproducing species, there are often striking differences in what are termed
secondary sexual characters. These differences can be placed (although not absolutely) in two
categories. The first encompasses all those structures (horns, antlers and the like) with which
males fight among themselves. Darwin hypothesized that the strongest, most vigorous males
would, most often, win such encounters and gain more matings than those who were not
victorious. These victorious males would then produce offspring more disposed to developing
such structures and the strength and vigour necessary to use them successfully in fighting.
Into the second category, fall structures and behaviour patterns used directly in courting
females. Those males with brighter colours, longer plumes or more melodious songs would be
preferred and hence chosen by females for mating. The elaboration of such attributes would
occur over several generations through the selective preference of the female. Darwin stipulated
that the necessary pre-conditions for sexual selection were a greater variability of one sex, the
greater proportion of one sex and competition among the members of the more numerous sex -
in most species, the males — for mating. I shall elaborate further on these points below and let
this statement of the theory stand for the present.
PRECEDENTS AND PREDECESSORS
Darwin was not operating in a vacuum. All theorizing must, to some extent, grow out of
what has gone before. If it does not grow directly from previous theory, then it must make use
of known fact. I will examine those that preceded Darwin to gain some understanding of the
facts and theories extant at Darwin’s time and a feeling for the perspective Darwin may have
had.
The key in examining development of the sexual selection hypothesis is not to discuss it in
terms of Darwin’s final (1871) formulation. Rather, the problem is best studied from the
Theories of sexual difference
3
perspective Darwin himself most probably would have had. The question seems to be not
specifically one of why a male is colourful, possesses horns or antlers or displays more readily
but rather a more fundamental amalgam of these points -- why are the sexes so different in so
many species?
A convenient benchmark for discussion of differences between sexes is the work of John
Hunter (1786). Hunter was the first to distinguish primary (those sexual differences present
at birth) from secondary sexual characters (sexual differences attendant with the onset of
sexual maturity). These secondary sexual characters are the subject of all subsequent debate.
Most authors of the late eighteenth and early nineteenth centuries recognized the more
mundane sexual differences (e.g. one sex larger than the other) as well as the more beautiful
and bizarre. Many examples of differences in secondary sexual characters amount to little more
than descriptions of oddities of animal structure that happened to catch the attention of a
particular author. Kirby and Spence (1828) describe the mating pads on the forelegs of the
male water beetle Acilius sulcatus (L.)
“Particular care seems to have been taken by the Creator that when all the above inhabitants are paired, the male
should be able to fix himself firmly by means of his remarkable anterior tarsi... and these asperities &c, in the upper
surface of his mate as not to be displaced by the fluctuations of that element, the reluctance of the coy female or any
other slighter cause.”
Similarly, in describing sexual difference in antennae of moths, they note that those of males
are often more complex and speculate:
“For what end the Creator has so distinguished them is not quite clear; but most probably this complex structure is
for the purpose of receiving from the atmosphere information of the station of the female.”
Not all discussion of sexual differences consisted of such isolated examples: more general
phenomena were discussed.
One class of structural features that attracted the notice of every writer on the subject was
the horns and antlers that males use in combat with each other, particularly at the time of
mating. For example, Hunter (1786) writes
“ ... as males of almost every class of animals are probably disposed to fight, they are, as I have observed, stronger
than the females. In many, there are parts designed solely for that purpose.”
and Kirby (1835) writes
“... the head ... generally only in the male is ornamented. ..these are used by the males in their mutual combats.”
In all passages quoted so far, each author shows a clear idea of the immediate function of
the structure being discussed. The forelegs of male Acilius are to grasp females, the complex
antennae of male moths are for detecting females and male horns and antlers (which I will
refer to collectively as armaments) are for winning fights. The purpose of winning such fights,
however, was stated by fewer. A clear exposition was given by Kirby (1835):
“ ... at the time of pairing, males contend fiercely and sometimes fatally for the females.”
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But for one author, the statements quoted so far seem to be reflective of the state of the art in
dealing with the ‘why’ of such structures. That author was Erasmus Darwin. The portion of his
Zoonomia (1794) relevant to sexual selection is worth quoting at length.
“As air and water are supplied to animals in sufficient profusion, the three great objects of desire, which have
changed the forms of many animals by their exertions to gratify them, are those of lust, hunger, and security. A great
want of one part of the animal world has consisted in the desire of the exclusive possession of the females; and these
have acquired weapons to combat each other for this purpose, as the very thick, shield-like, horny skin of the shoulder
of the boar is a defence only against animals of his own species, who strike obliquely upward, nor are his tushes for
other purposes, except to defend himself, as he is not naturally a carnivorous animal. So the horns of the stag are
sharp to offend his adversary, but are branched for the purpose of parrying or receiving the thrusts of horns similar to
his own, and have therefore been formed for the purpose of combating other stages for the exclusive possession of the
females; who are observed, like the ladies in the times of chivalry, to attend the car of the victor.
The birds which do not carry food to their young, and do not therefore marry, are armed with spurs for the
purpose of fighting for the exclusive possession of the females, as cocks and quails. It is certain that these weapons
are not provided for their defence against other adversaries, because the females of these species are with this
armour. The final cause of this contest amongst the males seems to be, that the strongest and most active animal
should propagate the species, which should thence become improved.”
Here is a clear attempt to go beyond a discussion of immediate function for a structure and seek
a more fundamental reason for its existence. This very passage led modern analysts of the
relationship of between the work of Erasmus and Charles Darwin to conclude that the theory of
sexual selection was not original (Irvine, 1955; King-Hele, 1963). This is only partially true.
The above passage shows that E. Darwin did elucidate the function of male combat before
Charles Darwin, but the female choice part of the theory was original.
Juxtaposed with the analysis of male armament given by E. Darwin (1794), the efforts of
Kirby and Spence (1828) at theorizing pale. The contrast in what we might consider the
‘scientific’ or ‘modern’ approach of Erasmus Darwin and that of Kirby and Spence stems from
the direction in which each author approached the problem. Kirby (1835) had neither the
inclination nor desire to upset the applecart of natural theology. His Bridgewater Treatise was
designed expressly to show “the power, wisdom and goodness of God” through an examination
of biological phenomena. If the thrust of one’s writing is to show that the animate world is a
divine machination then all other explanation becomes superfluous. Just as today, the
adaptation of an animal to its circumstances is assumed a priori to be illustrative of natural
selection, Kirby assumed such adaptation illustrated the magnificent intricacy of God’s
workings. Paley (1822) must be given the last word for the natural theologians. His concise and
single reference to differences between the sexes is,
“ Nor do the works of the Diety want this clearest species of relation. The sexes are manifestly made for each other.
They are for the grand relation of animated nature; universal, organic mechanical; subsisting, like the clearest
relations of art, in different individuals: unequivocal, inexplicable without design.”
There was one other great facet of animal appearance that was somewhat more problematic
— animal colouration. Since colour could not be seen as having a decisive role in the outcome of
any biological interaction, discussions of its function soon came to musings on aesthetics and
standards of beauty. The raison d’etre for colour came under the same nostrum as male
armament - being illustrative of the wisdom of God. Lesser (1791) combined the theological
and aesthetic approaches to animal colour. According to Lesser, brightly coloured insects are
beautiful to charm the eye and butterflies possessing bright underwings flash them so “their
Theories of sexual difference
5
beauty can be admired”. His final conclusion is that since colourful insects are generally of no
use “we should be grateful to the Creator for giving them to us”. Interestingly, it was this very
question of the function of animal colour and beauty that was to cause Darwin much theoretical
trouble.
The final point is one that is somewhat less concrete than the previous arguments and must,
to some extent, be read into the writings of various authors. It is the nearly universal practice of
anthropomorphizing the behavioural roles of the sexes in mating. Males (human or otherwise)
were viewed as aggressors, vying lustily for the attentions of the coy, demure females. Kirby
and Spence (1828) best show this approach in their description of insect song noting that
“female insects are too intent upon their business, too coy and reserved to tell their love even to
the winds”. Even Erasmus Darwin speaks of males fighting for “exclusive possession of the
females”. Apparently social outlook and norms transcended even the theoretical differences
these men might have wrestled with.
Here, too, was a crucial gap in reasoning about sexual interactions that Darwin’s sexual
selection was to fill. All previous authors failed to clarify the logical sequence between winning
a fight and mating with a given female (or females). The assumption seems to have been that it
was absolutely obligatory that a female mate with the victorious male. This is curious in view of
the reference Kirby and Spence (1828) made to the coyness of Acilius females.
DARWIN’S EARLY WRITING ABOUT SEXUAL SELECTION
Darwin’s writings about sexual selection prior to publication of the Descent of Man (1871)
are in five sources: his notebooks of 1838 (de Beer, 1960); his sketch of 1842; his Essay of 1844;
in a joint paper with Wallace (1858); and in Origin of Species (1859). 1
Darwin’s notebooks of 1838 (de Beer, 1960) show marked differences from previous
authors. Three tendencies appear in these notebooks that set the intellectual stage for
development of sexual selection theory.
First, to use Kuhn’s (1970) terms, Darwin appears not to have tried to fit the facts into the
paradigm of such people as Kirby, Lesser and Spence. One of the first references to sexual
differences to appear in Darwin’s notebooks is,
“Gould seems to think that (the) widow bird replaced Birds of Paradise -- if such fantastic sexual ornaments have so
intimate a relation in two continents as to be called into existence on two continents, our ignorance is profound and
such it appears.”
Clearly, he is not satisfied with the old natural theology or aesthetic arguments.
Secondly, Darwin’s continual search for a utilitarian reason for the existence of a particular
structure is evident. He writes that secondary sexual characters are developed
“ ... only when they first become of use”
Finally, Darwin can be seen as attempting to relate diverse types of secondary sexual characters
to a common cause,
'The works of 1858 (with Wallace), 1844 and 1842 are reprinted in Darwin and Wallace
(1958).
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“Dairies Barrington says cock birds attract females by song, do they by beauty - if not war (sic) not.”
Darwin’s theory first appeared in his Sketch of 1842, with reference to conditions necessary for
sexual selection to occur. There is a surplus of males at breeding time and thus males with the
“fullest vigour” will mate. It was a “struggle of war or charms”. Interestingly, at this point
there was no explicit statement of female choice. Darwin seems to assume, as did many of
predecessors, that females became the obligatory possession of victorious males. There was as
yet no role for choice by females in his writings.
His Essay of 1844 expanded some points mentioned in 1842. He reiterated the “struggle by
war or charms” notion and added two important points. The first was his recognition of what
may be a functional as opposed to a strictly statistical sex ratio when he noted that the struggle
would be severest in polygamous animals. Secondly, he noted that sexual selection was not as
rigourous as natural selection since the penalty for failure was not death but fewer offspring.
In the first edition of Origin of Species (1859), Darwin gives essentially the same treatment
to sexual selection as in Darwin and Wallace (1859 - see Darwin and Wallace (1958)). The
argument is developed as follows. Darwin noted that variations often arise in one sex and are
transmitted to that sex alone. Hence, one sex is modifiable “in its functional relations to the
other sex”. He repeats the contention that sexual selection is less rigourous than natural
selection and that the struggle is severest in polygamous animals.
The most significant new point in the Origin is that Darwin makes his first explicit reference
to female choice as a selective agent:
“ ... if a man can in a short time give elegant carriage and beauty to his bantams according to his standard of beauty,
then 1 can see no good reason to doubt that female birds by selecting during thousands of generations, the most
melodious or beautiful males according to their standard of beauty, might produce a marked effect.”
Two points in this passage, one explicit and one implied, caused Darwin considerable
trouble. He explicitly states that choice by the female could produce evolutionary change. Later
authors did not believe that mere capricious choice could effect evolution in any one direction.
Second, Darwin was seen as implying (by his standard of beauty statement) that females
possessed some highly refined aesthetic sense not unlike our own.
Darwin ends his statement of sexual selection with an affirmation of his own belief in the
extent of the process,
“ Thus it is, as 1 believe, that when the males and females of any animal have the same general habits of life but
differ in structure, colour, or ornament, such differences have been mainly caused by sexual selection.”
OPPONENTS AND CONTROVERSIES
Darwin’s chief opponent on sexual selection was, oddly enough, one of his closest colleagues
— Alfred Russel Wallace. Their controversy both reveals a clash between two close friends and
illustrates the danger of an hypothesis “becoming a thing in itself, a work of art.” (Steinbeck,
1951)
Wallace began with, as Vorzimmer (1970) puts it, “disinclination that led to complete
disavowal” toward Darwin’s hypothesis. In their joint publication, Darwin was concerned with
sexual selection while Wallace concentrated on natural selection. Correspondence between
Theories of sexual difference
7
them reveals Wallace’s growing “disinclination” quite clearly. In a letter to Darwin (29 May
1864 (Marchant, 1916)), Wallace questions some of Darwin’s ideas about sexual selection
stating that it would give “uncertain results” in the relation to man and that he (Wallace) has
found “little polygamy in the lowest tribes”. By 1868, Wallace questions some of Darwin’s
basic precepts. He does not understand how minute variations could be sexually selected.
Wallace writes that sexual selection requires a “series of bold and abrupt variations”. He asks
Darwin “how would an inch in a peacock’s tail or one-quarter inch in a Bird of Paradise be
noticed or preferred”. Darwin replied (29 March 1868 (F. Darwin, 1903)) that females would
be assessing the whole appearance of the male and not just comparing relative plume lengths.
Marchant (1916) claims that at this time (1867-1868), Wallace accepted the influence of
sexual selection much more than he did later. Darwin himself (30 April 1868 (F. Darwin,
1903)) complains that Wallace does not allow for the role of colour independent of protection in
print but does so in his letters.
Wallace’s firsf clear break with Darwin on this question came in his letter of 18 September
1868 (Marchant, 1916). In this letter, he clearly spells out his general and specific arguments
of animal colouration.
General Case
1 . Females are exposed to more danger and often live longer.
2. Females thus need more protection.
3. If males and females were separate species, we would think that natural selection had
operated on each.
4. Variation in one sex can often be transmitted to that sex alone. Thus, natural selection can
operate as if the sexes were separate species.
5. Natural selection can’t improve an animal beyond its needs.
6. In protected species, the sexes are similar.
Special Case
1. In weak flying moths, both sexes are mimics.
2. In strong flying moths, only the female mimicks.
3. Females can acquire bright colours.
4. There is no case of strong flying male alone mimicking.
5. Colour is more frequent in the male.
6. A male can’t get more protection than he needs.
After this exchange of letters, the differences between the two men became sharper until
1877 when Wallace wrote Darwin that he was “opposed to voluntary sexual selection”. Darwin
replied that to think a peacock’s tail and its movement during display were due to vitality and
vigour (see below) was “incredible”.
Wallace’s Darwinism (1901) was his grand summary of his evidence against Darwinian
sexual selection. This is a rather more readable and coherent account of his ideas than the
general and specific cases mentioned above. Wallace submitted that, because of their greater
vigour and vitality, males had a natural tendency to develop bright colours. In many species,
these colours developed in association with underlying organs and areas of intense nervous
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activity (e.g., the head). These colours were then passed on to both sexes. Females, however, did
not acquire such colour because of their need for protection and the development of such was
countered by natural selection. In his Theory of Bird’s Nests, Wallace pointed out that females
of many species that incubated in concealed nests were as colourful as their respective males.
He then extended the argument to other groups, especially the Lepidoptera. He pointed out
that in species which need protection (slow, weak fliers), both sexes mimicked other species. In
species which were stronger, faster fliers, only females were the mimics. Both concepts were
joined in what Wallace called his Theory of Protective Resemblance.
Armed with this notion and supported by Bates’ (1863) mimicry hypothesis, Wallace was
able to further generalize about protective function of colour and ornament in a wide variety of
organisms that ran the gamut from birds to caterpillars. A number of papers (Butler, 1869;
D’Orville, 1869; Alex. Wallace, 1869; Weir, 1869) were available noting the value of various
bumps, spines, bristles and colours in various creatures.
The main problem with all this criticism and counter-theorizing was one of misplaced
attack. Darwin tried to explain only the evolution of colours that were secondary sexual
characters. He at no time meant to give a comprehensive theory of animal colouration. As
George (1964) pointed out, Darwin and Wallace began by emphasizing different parts of the
colour question and continued, each in his own track, for several years.
Careful consideration of both theories shows quite readily their differences in emphasis.
Darwin was theorizing as to why males were brightly coloured. All Wallace (1891) could offer
that pertained directly to this point was the vitalistic argument that male colour was due to
“greater vigour and health and generally higher vitality”. Wallace, in his Theory of Bird’s
Nests, had a perfectly reasonable hypothesis as to why females are dull - not the same question
Darwin was trying to answer.
Wallace cannot take full blame for extending the argument into areas in which it did not
belong. An anonymous reviewer in the Edinburgh Review (1871) used presence of colour in
asexual forms (animals on which sexual selection, by definition, cannot operate) and colour in
inanimate objects as refutation of Darwin. The reviewer writes that identity of colour
throughout the realms of animate and inanimate nature (i.e., red is red whether found in a bird
or a rock)indicates similar cause. Perhaps this reviewer was St. George Mivart for in his
Lessons from Nature (1876), he states:
“It is reasonable to suppose that whatever cause has produced brilliant colour in either fishes or caterpillars may have
produced them in both.”
But let us return to Wallace for a moment. The argument that Darwin and Wallace were
examining different parts of the question of animal colour does not explain why the debate
continued for so long. It does not explain why Wallace (who cannot be said to have suffered
from lack of biological insight) put so much effort into what was apparently a “straw man”
controversy. I think responsibility for any disagreement must rest squarely with Wallace.
Darwin did not try to attack Wallace’s ideas but was quite impressed with them. In a letter to
Wallace (5 May 1867 (F. Darwin, 1903)), he wrote that Wallace’s insight into female colour
was “so much clearer and deeper than my own”. All of Darwin’s arguments were in favour of
sexual selection and not against Wallace’s protective resemblance theory.
Deeper reasons for Wallace’s disagreement lie in the realm of speculation. It is possible that
Wallace, having first been forced to share elucidation of the natural selection theory with
Theories of sexual difference
9
Darwin and then considered as the lesser light in the discovery, was trying to establish a major
theoretical advance that was purely his own. The advance he tried to make was that of a
comprehensive theory of animal colouration, a subject that had received little critical analysis
before 1871.
Darwin (1871) admitted that female choice was the weakest part of the sexual selection
hypothesis. Darwin had said that the females of a species could be the selective agent in the
elaboration of male adornment by choosing males with the longest plumes, brightest colours,
etc. To Darwin’s critics, this hypothesis had two rather unpalatable implications.
The first was possession of some aesthetic sense by any non-human female animal. It must
be pointed out that this notion of an aesthetic sense was not explicitly stated by Darwin. He
wrote that females would choose males on the basis of their adornment but, at first, gave little
concrete in the way of mechanism. Headley (1900), in a review of the whole question, wrote
that Darwin needed to show both that females need adequate perception to choose and that
they used that perception in choosing. This did not involve use of some subtle aesthetic sense
but simply elaboration of some sense of colour and hearing. Such development, according to
Headley, did not constitute an aesthetic sense. Similarly, in defence of Darwin, Romanes
(1896) argued that the taste of the female animal was not the same as that in humans and did
not necessarily imply intelligence.
On the other side of the coin, detractors were adamant — female choice did imply aesthetic
sense. Several (e.g., Edin. Rev. (1871))thought Darwin was unfair in erecting what seemed to
be untestable hypotheses. After all, how could one empirically decide what was going on in an
animal’s mind? There seemed to be one consistent characteristic in most of Darwin’s
detractors: they all had counter-explanations of their own. Three authors illustrate this point —
Stolzmann, the Duke of Argyll and Mivart.
In his Reign of Law (1884), Argyll errs by using the result of a process as an explanation for
its mechanism. He argued that to account for beauty as attractiveness to females was beside the
question, since females never mate with the wrong species anyway. His conclusion was,
“ Mere ornament and variety of form for their own sake is the only principle or rule with reference to which the
creative power seems to have worked.”
Mivart (1876) thought that Darwin conceded the point in question when he stated that
variation in structure could occur spontaneously. To Mivart, it was only a matter of degree as to
whether this variation gives a slightly different colour or a fully plumed Bird of Paradise. All
the evidence, Mivart says, points to,
“The existence of some unknown, innate and internal law which determines at the same time colouration and its
transmission to either or to both sexes.”
Mivart clearly doesn’t believe in female choice but the best he can offer as an alternate
explanation is a restatement of the problem.
Stolzmann (1885) was the most rigourous of the three in criticising Darwin and came up
with one of the more imaginative counter-theories. He argued that if one considered the
question from a larger geographical and phylogentic perspective, the notion of female choice
was thrown into doubt. The fact that European birds were generally less colourful than tropical
birds indicated that they (European birds) had less taste. Similarly, the fact that mammals
Quaest. Ent., 1982, 18 (1-4)
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were less colourful than birds indicated that that they too had less taste. The first fact,
Stolzmann argued, was illogical and the second was inconsistent with phylogenetic standing.
Stolzmann’s theory used many of the same precepts as Darwin’s. He said that males were
numerically superior, put less nourishment into eggs and offspring and were less important than
females in perpetuation of the species. They were, in a sense, parasitic on females after mating
was completed. Males were a drain on resources necessary to females and could even disrupt
incubation. All colour and ornamentation, then, performed three functions: it allowed females
to easily see males and avoid them; it allowed males to be easily seen by birds of prey; and it
reduced the chances of a male’s escaping a predator (females already being slower). Behaviour
associated with such structures served to attract attention of celibate males and keep them from
interfering with incubating females. It is unfortunate that Darwin died three years before the
publication of Stolzmann’s views. His response would have been interesting.
But what of Wallace? He was not as involved in the question of aesthetic taste of females as
he was in the question of animal colouration. Interestingly enough, criticisms from Wallace
focussed one ambiguity in the argument about female aesthetic sense. The process by which
female choice was effected was not made clear. Most discussion revolved around the issue of
whether females were excercising some conscious choice or were being excited by and yielding
to a male. Was it selecting or succumbing? Darwin (1871) originally thought it was selection.
He states:
“ No doubt this implies powers of discrimination and taste on the part of the female ... ”
Wallace (1891, 1901) objected to this notion of conscious choice, returning again and again
to the admonition that female choice could not be shown in nature. Wallace stated that while
female birds may be excited by a display of decorative plumage, there was no reason to suppose
that this conferred a mating advantage.
It is difficult to understand Wallace’s reasoning in the light of his own ideas. He stated that
colour and ornament are concomitant with vigour and general health and that it is the most
healthy, persistent males that will mate. Differences between Darwin and Wallace seem to be a
matter of mechanism rather than than basic principles. Indeed, in view of a paper by Darwin
(1876), it seems that Wallace is beating something of a dead horse. In that paper, there is a
rather abrupt change of attitude by Darwin. He wrote, in discussing sexual selection in
monkeys,
“ ... I presume that no supporter of the principle of sexual selection believes that the females select particular points
of beauty in males; they are merely excited or attracted in a greater degree by one male than by another ... ”
In a preface to a paper by Van Dyck (1882), Darwin again stated that it was more correct to
speak of females being excited by males rather than deliberately selecting.
Yet several authors persisted in making this argument for and against conscious female
choice. Romanes (1896) and Montgomery (1910) still referred to the bases of aesthetic
discrimination by females. Montgomery (1910) even took the argument a step further,
entertaining the question of whether a male was conscious of his own beauty and its effect on
the female. After a review of the subject, T.H. Morgan (1919) concluded that the evidence
supported the notion that the female did succumb rather than select. The controversy did have
one extremely beneficial effect: it stimulated great deal of research and careful observation
Theories of sexual difference
11
(e.g., Huxley, 1914; Montgomery 1910; Peckham and Peckham, 1899) and this research
provided a great fund of factual knowledge for Morgan to draw upon. He concluded that more
adorned males aroused females sooner. An aesthetic sense was not at issue for Morgan. He
drew on Montgomery’s (1910) work which showed by experiment that an aroused female shows
little preference for the male that aroused her. Morgan also concluded that the
“ ... purpose of the display may mean no more than a reaction that leads to a result propitious to the species.”
Morgan’s essay was significant in that it collected the observed facts and drew a conclusion that
took the argument out of the realm of the untestable and provided direction for further
research.
There was another aspect of female choice that Darwin’s critics refused to accept. It was
that choice by females could provide a consistent enough selective force to produce modification
of a structure and behaviour pattern in a given direction. One’s point of view on this question
would seem to depend largely on whether one accepted a more mechanistic view of a female
being excited by and succumbing to a male. If this view was accepted, then to imagine all the
females of a species possessing similar nervous, sensory and hormonal systems being excited by
similar displays and structures is not a large step. Add to this that animals deviating too far
from the norm had a greater chance of being culled out by natural selection. This was
essentially Darwin’s position (1876). He concluded,
“It should however be observed in the first place that although the range of variation of a species may be very large,
it is by no means indefinite. I have elsewhere given good instance of this fact in the pigeon of which there are at least
a hundred varieties differing widely in their colours and at least a score of varieties of the fowl differing in the same
manner; but the range of colour in these two species is extremely distinct . Therefore, the females of natural species
cannot have an unlimited scope for their taste.”
Most of Darwin’s antagonists, however, took his earlier statement of female choice. Mivart
(1876) stated that while he did not dispute that birds showed preferences, he could not accept
that “instability of feminine caprice” could produce secondary sexual characters. Similar
sentiments were echoed by Stolzmann (1885). Wallace (1901) seemed to address this question
only in passing. This may seem strange except for one factor. Wallace, in this own theory of
animal colouration, refused to accept that slight differences in males ability to excite females
could lead to a female preference, seemed to have disposed of the question almost before it was
raised.
I began by stating that Darwin realized his female choice hypothesis would not be accepted
readily. He admitted in a letter to Wallace that it was an “awful stretcher”. When all is said
and done, however, I think most of the objection to the idea (especially in its original
formulation) revolves around its lack of demonstrability. A statement appearing again and
again (in various forms) concerns the absence of proof of female choice in nature. It was only
near the end of Darwin’s life that empirical evidence began to emerge (e.g., Van Dyck (1882)).
It is unfortunate that Darwin did not live to see his ideas vindicated, at least in part, by later
workers (Morgan, 1919).
I conclude with a few remarks about a question that I have avoided thus far. Is sexual
selection a valid theory in its own right or merely a form of natural selection? Is Darwin, as the
reviewer in the Edinburgh Review (1871) complained, being unfair in raising new categories? I
think the answer lies both how widely one chooses to examine the question and what time frame
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is used.
If one examines all animate nature, then sexual selection is a special process that applies
only (by definition) to sexually reproducing animals. Natural selection can apply to all forms,
sexual and asexual. But this observation is trivial since Darwin stipulated that it only applied to
sexually reproducing forms. The question of time frame now emerges. If one takes the more
modern view of such authors as Fisher (1930), Williams (1966, 1975) and Dawkins (1976) that
the object of the evolutionary game is to have one’s genes represented in the next generation,
then negative results of natural selection (death of the individual) and sexual selection (failure
to breed) are one and the same. Consequently, the line of demarcation between the two ideas is
now recognized as being conceptually artificial. Historically, however, this is not fair to either
Darwin or his critics. Although ideas that reproductive products from the sexes were somehow
mixed in producing offspring were present (e.g. pangenes (Darwin, 1868), germ plasm
(Weismann, 1904) and idioplasm (Nageli, 1884)), such were viewed as mechanisms for
ensuring species (as opposed to individual) survival. To them, individual death and failure to
reproduce were distinct and thus sexual selection remained distinct from, although subordinate
to, natural selection. The idea that selection operates at the level of the individual received
much of its impetus from the work of Fisher (1930). The final statement of the problem goes to
E.B. Poulton (1890) who best summarizes the thoughts of those who accepted sexual selection.
“Natural selection is a qualifying examination which must be passed by all candidates for honours: sexual selection is
an honours examination in which many who have passed the previous examination will be rejected.”
ACKNOWLEDGEMENTS
I wish to thank Dr. M.P. Winsor, Institute for the History and Philosophy of Science and
Technology, University of Toronto for many fruitful discussions and much constructive
criticism. G.C. Williams reviewed the paper and provided some new and interesting insight.
This paper was written while I was supported by NSERC (Canada) Operating 4946 to G.K.
Morris (University of Toronto). The final stages of manuscript preparation were generously
funded by the Department of Entomology, University of Alberta.
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Anonymous. 1871. Article VIII. Edinburgh Review 134: 99 - 120.
Argyll, Duke of. 1884. The works of the Duke of Argyll containing the Reign of Law, the Unity
of Nature and Primeval Man. John B. Alder. N.Y.
Bates, H.W. 1863. A naturalist on the river Amazons: A record of adventures, habits of
animals, sketches of Brazilian and Indian life and aspects of nature under the equator
during eleven years of travel. J.M. Dent and Sons. London (reprint of the first edition
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Butler, A.G. 1869. Remarks upon certain caterpillars etc. which are unpalatable to their
enemies. Trans, ent. Soc. Lond. 1869: 27 - 29.
Darwin, C. 1859. On the origin of species by means of natural selection, or the preservation of
favoured races in the struggle for life. John Murray. London, (facsimile of the first
edition, Harvard Univ. Press (1964)).
Darwin, C. 1868. Variation of plants and animals under domestication. John Murray. London.
Theories of sexual difference
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Darwin, C. 1871. The descent of man and selection in relation to sex. John Murray. London
(second edition, 1880, Appleton and Co. N.Y.).
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Darwin, E. 1794. Zoonomia. London.
Darwin, F. 1903. More letters of Charles Darwin. John Murray. London.
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Fisher, R.A. 1930. The genetical theory of natural selection. Clarendon Press. Oxford.
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Headley, F.W. 1900. Problems of evolution. Duckworth and Co. London.
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history of insects. London, (fifth edition)
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LIFE CYCLES OF MAYFLIES (EPHEMEROPTERA), WITH SPECIAL REFERENCE TO
VOLTINISM
HUGH F. CLIFFORD
Department of Zoology, University of Alberta
Edmonton, Alberta, Canada
T6G 2E9 Quaestiones Entomologicae
18: 15-90 1982
ABSTRACT
This report is a survey in breadth of mayfly (Ephemeroptera) life cycles and includes data
about 718 life cycles for 297 species. Most of the published reports are for Northern
Hemisphere mayflies. About 60% of all mayfly life cycles were reported as univoltine, 30%
multivoltine, 4%) semivoltine, and 3%o were judged variable, i.e. for a given population of a
single report there would be different types of voltinism. In addition, for many species,
different studies indicated different life cycle types. The survey indicates extensive flexibility
of life cycles for many but not all mayfly species.
The univoltine winter cycle (single generation overwinters in the nymphal stage ) is the most
common single life cycle type. Baetid species, mainly Baetis, are mainly multivoltine.
Ephemerellids, mainly Ephemerella, and heptageniid species are mainly univoltine, members
of these two families accounting for more than half the univoltine cycles reported.
Ephermerids, entirely Ephemera and Hexagenia, are the major semivoltine group. Cycles
exhibiting a long period of egg dormancy, either in summer or winter, are common in
mayflies; for example about 25% of all life cycles are summer cycles with a long period of
egg dormancy in winter.
Yearly nymphal periodicity (or cohort production interval, CPI), an important factor in
secondary production estimates, was calculated for species of all life cycle types. The average
values for Northern Hemisphere mayflies of each life cycle are: 11.0 months for univoltine
winter cycle (no summer egg-dormancy ); 7.8 months for univoltine winter cycle with summer
egg-dormancy; 4.1 months for univoltine summer cycles; and 23.8 months for the semivoltine
cycles.
Each life cycle type has a characteristic seasonal reproductive period (equating
reproductive period with adult periodicity). About half of all Northern Hemisphere mayflies
reproduce in June and July. Multivoltine mayflies have the most extensive reproductive
period and the semivoltine mayflies the most restrictive. Univoltine summer mayflies
generally reproduce later in the year than univoltine winter mayflies. Possible adaptive
strategies of these average reproductive periods are discussed.
Clear relationships between life cycle types and body size of individual species or
phylogenetic position are not evident, but mayfly life cycle types may change predictably from
the tropics to the arctic. Some features of flexible and rigid life cycles are discussed, as is
importance of distinguishing cohorts from generations. Several areas of future research are
suggested, especially the need for complete voltinism data of tropical mayflies.
Key words:
16
Clifford
Ephemeroptera, mayflies, life cycles, voltinism, strategies, reproduction,
production, literature.
RESUME
Ce rapport presente une reconnaissance complete des cycles biologiques des ephemeres ( Ephemeroptera ); il contient
des donnees sur 718 cycles biologiques appartenant a 218 especes. La plupart de ces donnees proviennent d’esp'eces de
I’hemisphere nord. Ces rapports ont indique qu’environ 60% de ces cycles sont univoltins, 30% multivoltins, 4%
semivoltins, et que 3% sont variables, c’est a dire que pour une certaine population, un rapport a indique la presence de
plusieurs types de voltinisme. De plus, pour un grand nombre d’especes, differentes etudes ont rapporte differents types
de cycle biologique, indiquant qu’il existe une grande souplesse, quant au type de cycle, chez un certain nombre d’especes
d’ephemeres.
Le cycle univoltin hivernal (une generation hiberne a I’etat de nymphe) est le type de cycle le plus courrant. Les
especes de Baetidae (principalement Baetisj sont surtout multivoltines, tandis que les especes d’Ephemerellidae
(principalement Ephemerellaj et d’Heptageneiidae sont surtout univoltines, ces deux families contenant plus de la moitie
des cycles univoltins. Les Ephemeridae (toutes les especes ^’Ephemera et d’Hexageniaj forment le groupe principal
d’especes semivoltines. Une longue periode de dormance, a I’etat d’oeuf aussi bien en ete qu’en hiver, est commune chez
les ephemeres: par exemple, environ 25 % des cycles decrits sont des cycles estivaux avec une longue periode de dormance
hivernale.
La duree du stage nymphal (ou intervalle de production des cohortes), une importante donnee requise pour
I’estimation de la production secondaire, a ete calculee pour des especes appartenant a tous les types de cycle
biologiques. Les durees moyennes du stage nymphal des ephemeres de I’hemisphere nord sont respectivement 1 / mois
pour les especes univoltines hivernales (sans periode de dormance estivale), 7.8 mois pour les especes univoltines
hivernales (avec periode de dormance estivale), 4.1 mois pour les especes univoltines estivales et 23.8 mois pour les
especes semivoltines.
Chaque type de cycle biologique a une saison de reproduction (qui est assimilee a la periode d’ apparition des adultes)
qui lui est caracteristique. Pres de la moitie des especes d’ephemeres de I’hemisphere nord se reproduisent en Juin et
Juillet. Les periodes de reproduction les plus longues se trouvent parmi les especes multivoltines tandis que les periodes
les plus courtes se trouvent chez les especes semivoltines. Les especes univoltines estivales se reproduisent generalement
plus tard que les especes univoltines hivernales. La valeur adaptative de ces diverses longueurs de la periode reproductive
est examinee.
II ne parait pas y avoir de correspondance entre le type de cycle biologique et la taille, ou la position phylogenetique
de chaque espece; mais il semble que les cycles biologiques changent d’une faoon previsible des tropiques aux poles.
Certaines caracteristiques des cycles qui sont rigides, vis-a-vis de leurs pendants plus souples sont examinees ainsi que la
necessite de faire la distinction entre cohortes et generations. Plusieurs domaines de recherche future sont suggeres et
I’importance d’obtenir des donnees sur le voltinisme des especes tropicales est particulierement notee.
Ephemeroptera, ephemeres, cycles biologiques, voltinisme, strategies, reproduction, production, revue.
Table of Contents
INTRODUCTION 17
METHODS 19
Literature Survey 19
Life Cycle Types 21
Explanation of Tabular Summary Tables 23
GENERAL ANALYSIS 23
Baetidae 23
Caenidae 27
Ephemerellidae 29
Ephemeridae 31
Heptageniidae 32
Leptophlebiidae 33
Metretopodidae 34
Life cycles of mayflies (Ephemeroptera)
17
Oligoneuriidae 35
Polymitarcidae 43
Potamanthidae 43
Siphlonuridae 43
Tricorythidae 44
Other Families 44
Synopsis 44
SUMMER CYCLES 45
EGG DORMANCY IN SUMMER 46
LIFE CYCLE TYPES AND PRODUCTION 47
LIFE CYCLE TYPES AND REPRODUCTION 49
LIFE CYCLE TYPES AND OTHER FACTORS 52
DISCUSSION 53
FURTHER RESEARCH 54
ACKNOWLEDGMENTS 55
LITERATURE CITED 55
APPENDIX 77
INTRODUCTION
Ephemeroptera, or mayflies, is a small but important order of aquatic insects. Twenty
families of recent mayflies include slightly more than 200 genera and 2,000 species. Adult
mayflies are unique among insects because there is usually an additional molt once the winged
stage is achieved. Adults (the subimago and imago stages combined) seldom live for more than
three days and do not feed. Most nymphs have, relative to most insect groups, a large and
varied number of instars (from about 10 to 50 depending on the species). Most nymphs live
only in unpolluted lakes and streams, where they sometimes occur in very large numbers (up to
10,000/m2); they can be an important component of secondary production generally and food
for fish specifically. They have also been used as models for a variety of experimental aquatic
studies. Unfortunately, accurate instar analysis of field populations is usually impossible, and
this aspect of their way of life is usually missing in most life cycle studies.
The year 1957 might be considered a benchmark in study of mayfly life cycles; in that year
T.T. Macan (1957b) reported on the life cycles of several British species and for each presented
nymphal size frequency data throughout the life cycle. Since the 1 950’s, mayfly life cycle data
have rapidly accumulated (Fig. 1), many of the studies being modelled after those of Macan. In
the 1960’s and 1970’s, with the emphasis on computer-assisted quantitative studies of
communities, descriptive life cycle studies did not receive high research priority. Although
mayfly life cycle data continued to accumulate rapidly, often these data in reports of benthic
studies were relegated to brief, observations, with no possibility of data interpretation by
interested readers.
Presently, there is renewed interest in life cycles, as aquatic biologists realize the life cycle is
an integral part of whole community processes and therefore often needed for interpreting these
processes. In 1978, a symposium entirely devoted to life cycles and sponsored by the North
American Benthological Society of America was held in Winnipeg, Canada (see Rosenberg
Quaest. Ent., 1982, 18 (1-4)
18
Clifford
Years : 1901 - 1979
Figure 1 . N umber of published reports pertaining to mayfly life cycles, 1901-1 979.
Life cycles of mayflies (Ephemeroptera)
19
1979). Several workers pointed out the practical importance of life cycle data to aquatic
research: sampling procedures for insects (Resh) and non-insects (Malley and Reynolds),
taxonomy (Oliver), bioassays (Buikema and Benfield), pollution (Lehmkuhl) and food quality
(Anderson and Cummins). In the symposium’s summary, Waters called for a data bank of
basic life cycle information and pointed out the importance of including features, e.g. voltinism,
that might be expected under different environmental conditions. One result of this symposium
was a review paper (Waters 1979) on how life cycles influence production estimates of benthic
invertebrates.
Since 1965, my students and I have acquired complete life cycle data for 42 mayfly species
of Alberta, Canada. Data on only about 40% of these cycles have as yet been published.
Initially I had planned a life cycle synopsis of only the Alberta species; but an extensive survey
of mayfly life cycles now seems in order, and the synopsis was expanded to include data about
all mayfly life cycle reports that I could obtain.
A few key features, e.g. life span, growth, reproduction, when quantified in time, will each
express information about the sum total of numerous biological processes and can be
conveniently used to classify an organism’s life cycle. The four criteria of mayfly life cycles
considered in this survey follow those outlined by Tauber and Tauber (1978) for insects
generally: egg dormancy, growth and development, reproduction, and dispersal, although
dispersal was not treated in detail. Criteria were treated mainly from a phenological approach
emphasizing voltinism of the various taxa. “Life cycle” and “life history” are used
synonymously, although to some workers the terms have different meanings; e.g. systematists
might prefer to restrict the term life history to connotations about evolutionary history of
species.
The objectives were to (1) make available references about life cycles of the various mayfly
taxa, (2) group and analyze life cycles using a voltinism classification scheme, and (3) discuss
some of the mayfly life cycle types as they relate to interpretations of aquatic community
processes and as they relate to adaptive strategies of mayflies.
METHODS
Literature Survey
About 400 papers were used in the survey. The survey stopped in October 1979; but I also
included some published accounts of mayfly life cycles appearing later, especially those in the
Proceedings of the Second and the Third International Conferences on Ephemeroptera. The
following information was recorded for each species of each life cycle report: geographical
location including latitude and, if pertinent, altitude, lentic or lotic study, type voltinism (see
below), nymphal periodicity, adult periodicity, and egg stage duration. For many species egg
duration was deduced from nymphal and adult periodicity. The species’ life cycles were then
grouped into one of two categories based on completeness of the study: ( 1 ) life cycle reports
that always included data on voltinism and often information on growth, egg stage and adult
periodicity; (2) reports pertaining only to adult periodicity (mainly emergence studies); this
information was usually only recorded if the adult data were from routine sampling or
observations throughout the adult stage of the population in question. Data of this second
category were used only in the Reproduction Section.
Occasionally life cycles of a particular study were reported more than once. An attempt was
made to use these cycles only once in the analysis (although all the literature citations were
Quaest. Ent., 1982, 18 (1-4)
20
Clifford
Table 1. Total numbers and percentage of species having at least one cycle of the type in
question and total number and percentage of life cycles reported for each type. See Methods for
further explanation.
Totals 297 718
included), but some redundancy undoubtedly exists. Another problem pertains to taxonomic
treatment. Several taxonomic revisions have been published during and after the survey. It was
not possible to incorporate all of these into the survey, but their omission should have little
bearing on the analysis. Except for using the recent Baetis revision of Morihara and
Life cycles of mayflies (Ephemeroptera)
21
McCafferty (1979) and Ephemerella revision of Allen (1980), names follow Edmunds et al.
(1976) for North American species, Puthz (1978) for European species; and for the few species
reported from other areas the nomenclature of the article in question was used.
Life Cycle Types
The classification scheme used here implies a degree of rigidity for mayfly life cycles that
for many species (as will be discussed later) is probably not justified. Nevertheless, some sort of
classification is necessary as a first step in reducing the complexities of mayfly life cycle
features to an orderly system. I chose a voltinism scheme because most mayfly life cycles have
been reported in terms of generations per year. An alternate system is to separate annual cycle
into non-seasonal and seasonal cycles, with the seasonal cycle being further divided into slow
seasonal and fast seasonal cycles (Hynes 1961). One advantage here is that generations and
cohorts do not always have to be distinguished. Unfortunately, not enough mayfly life cycles
have as yet been described in terms of fast and slow cycles to make the scheme applicable for a
global survey.
The major categories (see for example Table 1), univoltine, multivoltine and semivoltine,
need no explanation. The greatest degree of certainty in accurately interpreting the life cycle is
implied by the major voltinism types, and much of the analysis and discussion is based only on
major categories. Each of these three major life cycle types (plus two additional major types,
see below) was separated into more specific life cycle categories based in part on Landa’s
(1968) classification system. These subtypes are more subjective, but compilations based only
on major voltinism features could conceal important life cycle differences. Life cycle types are
defined as follows:
Univoltine.
U? - a seasonal univoltine cycle; but the report does not indicate what type.
Uw - seasonal univoltine winter cycle; the population overwinters in the nymphal
stage.
Us - seasonal univoltine summer cycle; hatching, growth and emergence take place
in summer; the population overwinters in the egg stage.
Us-Uw
- a seasonal univoltine cycle where most of the new generation overwinters in the
egg stage, but a small part of the population overwinters in the nymphal stage.
Multivoltine.
M? - seasonal multivoltine cycle, but the report does not indicate what type; the few
reports of three generations in two years were also placed in this type.
MB - the total number of seasonal bivoltine cycles.
MB? - a seasonal bivoltine cycle, but the report does not indicate whether there are
two summer generations or a summer and a winter generation.
MBss - seasonal bivoltine summer cycle; following summer generations, the population
overwinters in the egg stage.
MBws
- seasonal bivoltine winter-summer cycle. There is an overwintering generation
in the nymphal stage and one summer generation.
Quaest. Ent., 1982, 18 (1-4)
22
Clifford
MP - the total number of seasonal polyvoltine cycles where there are three or more
generations a year, usually consisting of two or more summer generations and
an overwintering generation in the nymphal stage. Throughout, polyvoltine
refers only to three or more generations a year, though polyvoltine usually
means the same thing as multivoltine. Using polyvoltine in this sense is more
convenient than constatly saying “trivoltine or greater” when contrasting these
multivoltine cycles with the bivoltine cycles.
MB-MP
- a seasonal multivoltine cycle where the population of a given study might be
bivoltine or polyvoltine depending on year, local habitat, or in some taxa partly
bivoltine and partly polyvoltine from the same egg cohort.
MNP - non-seasonal polyvoltine cycle. These reports are from tropical and
semitropical regions; for many, polyvoltinism was inferred from emergence data
if there were few observations of other life stages.
Total Uni-Multivoltine.
This major category and the Total Uni-Semivoltine category are the variable
life cycle types and include species whose life cycles (of a given report) vary
between major types of voltinism.
Us-MBss
- a seasonal variable cycle, where the report indicates one or two summer
generations, depending on year, local habitat, or even from a single egg cohort.
Uw-MBws
- a seasonal variable cycle where the report indicates the population typically
has a univoltine winter cycle; but, depending on year, local habitat or even from
a single egg cohort, there occasionally might also be a summer generation.
Total Semivoltine.
2Y - seasonal semivoltine cycle with a generation time of about two years (closer to
two years than one year).
3Y - seasonal semivoltine cycle with a generation time closer to three years than two
years.
2Y-3Y
- seasonal semivoltine cycle where the report indicates one generation in two or
three years, depending on year, local conditions, or even from a single egg
cohort.
Total Uni-Semivoltine
Uw-2Y
- a seasonal variable cycle where the report indicates either a univoltine winter
cycle or a two-year semivoltine cycle, dependingin on year, local habitat or from
a single egg cohort.
Life cycles of mayflies (Ephemeroptera)
23
Explanation of Tabular Summary Tables
Tabular summaries for each family and genus have six columns. Meaning of column
headings and significance of their values should be clear by working through an example, i.e.
Table 2, Baetidae.
Column 1 ( Species No.). - Adequate voltinism data were available for 58 species of
Baetidae; 1 3 species of Baetidae had at least one Us (univoltine summer) cycle, etc. Thus, some
subtotals add to more than total number because some species were reported with more than
one life cycle type.
Column 2 (% Species in Family). - 22.4% of all Baetidae species had at least one Us cycle,
etc. For reasons given for column 1, the subtotal percentages do not necessarily add to 100.
Column 3 (% All Species— with that life cycle type). - Baetid species accounted for 16.3%
of all mayfly species having a Us cycle.
Column 4 (No. of Life Cycles). - The 58 species of Baetidae accounted for a total of 183
reported life cycles; 23 of Baetidae life cycles were reported as Us. Here, subtotals of major
voltinism types add to 183.
Column 5 (% Life Cycles in Family). - 12.6% of all Baetidae life cycles were Us.
Column 6 (% All Life Cycles— of that life cycle type). - Baetidae life cycles accounted for
16.3% of all life cycles reported as Us.
Tabular summaries are given for most families and for species having at least five life cycle
reports. Lists of all species, their literature citations, and “consensus” life cycle types are given
in the appendix.
GENERAL ANALYSIS
The survey consisted of 7 1 8 life cycles for 297 mayfly species (Table 1 ). The totals are 1 ,474
items of life cycle information for 490 species with inclusion of data pertaining strictly to adult
periodicity.
Baetidae
Values for this family constituted 25% of all mayfly life cycles reported. Generally,
members of this widely distributed family exhibit seasonal multivoltine cycles (Table 2). Only a
few baetid species exhibit the most common mayfly life cycle type, the univoltine winter cycle.
Baetis Leach
Substantially more life cycles were reported for Baetis than for any other mayfly genus.
Populations of the 32 species with adequate voltinism data are mainly multivoltine (Table 3).
Baetis species account for most of the exclusively summer (Us, MBss, Us-Mbss) life cycle
types. Baetis populations had few univoltine winter cycles.
Number of life cycle reports for B. rhodani populations was second only to the number
reported for Ephemerella ignita (Table 4) . Throughout its range, B. rhodani populations
appear to have consistently a seasonal bivoltine cycle with an overwintering generation in the
nymphal stage and one summer generation. Sometimes, however, there can be a second
summer generation, for example in the Atlantic Pyrenees of France (Thibault 1971b, Benech
1972, Neveu et al. 1979). In many areas, B. rhodani adults are found continuously throughout
the summer, and several workers reporting a bivoltine cycle suggest the possibility of additional
Quaest. Ent., 1982, 18 (1-4)
24
Clifford
Table 2. Baetidae tabular summary. Number and percentage of species having at least one
cycle of the type in question in family and number and percentage of life cycles reported for
each type. See Methods for further explanation.
Species Life Cycles
No. % Fam. % All No. % Fam. % All
Total Univoltine
Uw
Us
Total
Multivoltine
M?
MB
MB?
MBss
MBws
MP
MB-MP
MNP
Total Uni-
Multivoltine
Uw-MBws
58
183
Totals
Life cycles of mayflies (Ephemeroptera)
25
Table 3. Tabular summaries: Baetis and Ephemerella. Number and percentage of species
having at least one cycle of the type in question in genus and number and percentage of life
cycles reported for each type.
Quaest. Ent., 1982, 18 (1-4)
26
Clifford
Table 4. Number of life cycles of each type reported for (1) Baetis rhodani, (2) B. alpinus, (3)
B. muticus, (4) B. vernus, (5) B. scambus, (6) B. fuscatus, (7) B. tricaudatus, and (8)
Centroptilum luteolum.
summer generations. Humpesch (1979b) distinguished 10 B. rhodani cohorts over 30 months in
an Austrian population; although cohorts should not be confused with generations, this perhaps
suggests a seasonal polyvoltine species, with absolute number of generations varying from year
to year.
Baetis rhodani populations with univoltine winter cycles have been reported from the far
north, above latitude 65 °N (Ulfstrand 1968), but also from upland areas at lower latitudes, i.e.
Germany (lilies 1952) and Poland (Sowa 1965). Most of Baetis species of the far north, e.g. B.
macani macani in Europe and B. macani bundyae (see Appendix for all species not included in
the tables) in North America, exhibit summer cycles, but there are apparently no reports of
exclusively summer cycles for B. rhodani populations at high latitudes. There is one report of a
summer cycle, i.e. two summer generations, for B. rhodani from Austria (Pleskot 1961).
Populations of the Palearctic B. alpinus , B. muticus and B. vernus exhibit one of several life
cycle types, although they are mainly bivoltine with a nymphal winter generation (Table 4); in
contrast B. scambus and B. fuscatus , also Palearctic species, are mainly summer species. Life
cycle reports for the North American B. tricaudatus (including B. vagans ) indicate only
multivoltine cycles, populations having either an MBws bivoltine cycle or a seasonal polyvoltine
Life cycles of mayflies (Ephemeroptera)
27
cycle with one overwintering nymphal generation and two or more (although the exact number
often appears to be speculative) summer generations.
Cloeon Leach
Cloeon species exhibit a variety of life cycle types; and, in contrast to Baetis populations, the
life cycle types are fairly evenly distributed between univoltine winter cycles, bivoltine cycles,
seasonal polyvoltine cycles, and the variable uni-multivoltine type. Few exclusively summer
cycles were reported for Cloeon.
Two species, C. dipterum and C. simile , account for most of Cloeon life cycles (Table 7).
The mainly Palearctic C. dipterum (it has been reported once from North America, Traver
1962) has no uniform life cycle type; in fact, populations apparently are very flexible in their
life cycles, often exhibiting univoltine winter cycles, occasionally bivoltine cycles, and
sometimes a single population is partly univoltine and partly bivoltine (a uni-multivoltine life
cycle type). Cloeon dipterum populations are ovoviviparous, at least in part of the species’
range, but how this might be related to the flexible voltinism of this species is obscure.
Certainly, ovoviviparity would account for the absence of exclusively summer cycles in C.
dipterum. The non-ovoviviparous C. simile has been reported as having a summer cycle; but
generally, as for C. dipterum , populations appear to be quite flexible, having either univoltine
or multivoltine cycles, presumably depending on environmental factors.
Other Baetid Genera
The Centroptilum species with adequate voltinism data are mainly multivoltine, with a
relatively large proportion of the species having at least one bivoltine summer-summer cycle.
The Palearctic C. luteolum, the most reported Centroptilum species, apparently always
features an overwintering nymphal population, most of the life cycle being reported as the
bivoltine winter-summer cycle (Table 4). However, there is some dispute, at least for certain
areas, whether the cycle is bivoltine or univoltine with an extended period of emergence (see
Bretschko 1965, Macan and Maudsley 1968, Lavandier and Dumas 1971, and Brittain 1974).
Populations of Pseudocloeon are mainly multivoltine, with a large proportion of seasonal
polyvoltine cycles. The two species of Procloeon with voltinism data, P. bifidum (Table 7) and
P. ornatum , are mainly summer species, with two summer generations. The remaining baetid
genera for which there were voltinism data, Callibaetis , Baetiella and Baetodes (see appendix),
exhibit multivoltine cycles.
In short, members of this important mayfly family exhibit a variety of life cycle types, with
a tendency for multivoltine cycles and also flexibility in life cycle types. Species of Baetidae,
especially Baetis species, are consistently important in numerical abundance studies (Clifford
1980), and perhaps this dominance in numerical abundance is accounted for in part by the
plasticity in their life cycle features.
Caenidae
Life cycle reports for Caenidae made up about 5% of all mayfly life cycle reports. At the
family level, there was no consistent life cycle type, about half the cycles being univoltine and
half being multivoltine (Table 5).
Most life cycles of Caenidae were those of Caenis species. About half the Caenis cycles are
univoltine, mainly univoltine winter, and about half are multivoltine, mainly the bivoltine
Quaest. Ent., 1982, 18 (1-4)
28
Clifford
Table 5. Caenidae and Oligoneuriidae tabular summaries.
Life cycles of mayflies (Ephemeroptera)
29
Table 6. Ephemerellidae tabular summary.
winter-summer cycle. Even at the species level, i.e. C. horaria , both univoltine and multivoltine
cycles have been reported (Table 7). In short, as was found for many of the baetids, life cycle
types of many Caenis populations seem quite flexible. In contrast, Brachycercus populations
appear to exhibit mainly univoltine summer cycles; species of Brachycercus accounted for four
of six univoltine summer cycles of Caenidae.
Ephemerellidae
Generally, ephemerellid species exhibit univoltine cycles, with more winter cycles than
summer cycles (Table 6). Collectively, ephemerellid and heptageniid populations account for
about half the univoltine cycles.
Quaest. Ent., 1982, 18 (1-4)
30
Clifford
Ephemerella (sensu stricto , see Allen 1980)
Ephemerella species account for most life cycle reports (Table 3) of Ephemerellidae.
Ephemerella populations exhibit mainly univoltine cycles. The large number of summer cycles
reported is due mainly to numerous life cycles reported for E. ignita, this species accounting for
more than half of all Ephemerella life cycle reports (Table 7). There were more life cycle
reports for E. ignita than any other mayfly species. Populations of this Palearctic species
exhibit mainly univoltine summer cycles, but there were reports of univoltine winter cycles,
bivoltine winter-summer cycles, and even cycles (Us-Uw) where part of the population
overwinters in the egg stage and part in the nymphal stage. Elliott (1978) surveyed many life
cycle reports for E. ignita through 1977; he states the cycle in upland streams of Europe is
mainly the univoltine summer cycle; whereas in various lowland European streams, nymphs
have been found throughout the winter. The complex relationship between water temperature
Life cycles of mayflies (Ephemeroptera)
31
and hatching time of eggs of E. ignita has been studied in the laboratory by Bohle (1972) and
Elliott (1978). In a field study, Neveu et al. (1979) confirmed Elliott’s results. In contrast to E.
ignita , E. aurivillii and E. mucronata populations generally have straight-forward univoltine
winter cycles.
A variation of the univoltine winter cycle occurs when the population spends much of the
summer in some sort of egg dormancy state. As discussed later, this type of univoltine winter
cycle, which might be considered opposite of the univoltine summer cycle (where egg stage
dormancy occurs in winter), is probably commoner than would be indicated by publication
about life cycles. Of 15 species judged to have at least 4 months of egg dormancy in summer,
five were Ephemerella species.
Other ephemerellid genera
Eurylophella and Drunella populations have mainly univoltine winter cycles. However,
Eurylophella funeralis had a 2-year cycle in a northeastern USA stream (Fiance 1978). This
appears to be the only record, based on adequate life cycle data, of an ephemerellid having a
semivoltine cycle. Serratella species have been reported exhibiting either univoltine winter or
univoltine summer cycles. Torleya major ( — T. belgica) populations were exclusively univoltine
winter (Table 7). The one life cycle reported for the tropical Teloganodes indicates a
non-seasonal life cycle with emergence throughout the year.
Ephemeridae
This is the major family of semivoltine mayflies, accounting for 58% of the species having
exclusively semivoltine cycles and 82% of all the exclusively semivoltine cycles. Species of
Ephemera and Hexagenia account for all the ephemerid life cycles reported (Table 8).
Ephemera Walsh
Most Ephemera life cycles were reported as semivoltine; however four of the six species
exhibit at least one univoltine cycle. Two species, E. danica and E. simulans, account for most
of Ephemera life cycle reports (Table 14). Populations of the Palearctic E. danica were
reported as having univoltine winter cycles, semivoltine cycles, and sometimes with both
univoltine and semivoltine cycles (Uw-2Y). Although other workers have alluded to a 3-year
cycle for ephemerids, or other mayflies, e.g. Palingenia , the studies of lilies (1952) and
Svensson (1977) are the most convincing of a definite three-year cycle. Svensson found for an
E. danica population in Sweden that part of the population required two years to complete the
cycle, whereas the other part took three years; at any one time as many as five generations were
present. In contrast, Thibault (1971b), for an E. danica population of the French Pyrenees,
found that from a single egg cohort, depending on hatching time, part of the population
required one year to complete the cycle and part required two years. Whelan (1980) also
reported a flexible, one- or two-year cycle for E. danica in Ireland. And Landa (1968) reported
E. danica to have two generations in three years in central Europe. Clearly, this thoroughly
studied species has a very flexible life cycle.
Life cycle reports for E. simulans indicate populations having either univoltine winter cycles
or two-year cycles. There were neither variable cycles (i.e. both univoltine and semivoltine
cycles from the same population) nor three-year cycles reported for E. simulans.
Quaest. Ent., 1982, 18 (1-4)
32
Clifford
Hexagenia Walsh
Relatively fewer univoltine cycles were reported for populations of the Nearctic Hexagenia
than for Ephemera (Table 8). Two species, H. limbata and H. bilineata , account for most of
the Hexagenia life cycle reports (Table 14). Populations of H. limbata (not including the
subspecies occulata and viridescens) were reported as having univoltine cycles, semivoltine
cycles, and occasionally within a single population, depending on hatching time of the single
egg cohort, part of the population completed the life cycle in one year and part required two
years. Possibly there is a tendency for two year cycles for northern populations and one year
cycles for southern populations (Swanson 1967); but other workers, e.g. Rutter and Wissing
(1975), suggest that given the proper local conditions, especially water temperature, probably
most H. limbata populations can exhibit either one- or two-year cycles or sometimes both
cycles. In most taxa with both a one-year and two-year component to the cycle, most of the
population exhibits the two-year cycle (Hudson and Swanson 1972, Rutter and Wissing 1975).
Flannagan (1979) reported two life cycle types for H. limbata populations of Lake Winnipeg,
Canada: one type featured a 14-month life span alternating with a 22-month life span; and the
other was the typical two-year cycle. Obviously H. limbata populations have flexible life cycles,
although three-year cycles have not been reported. In fact, Hunt (1953), for a Michigan, USA,
population, obtained evidence based on water temperature data, that the maximum life span
would probably have to be less than three years.
Hexagenia bilineata is mainly univoltine, though Fremling (1973a) obtained evidence for a
quick summer generation as well as a winter generation. Certainly laboratory studies of
nymphal growth suggest that rapid generations are possible for H. bilineata (Thomforde and
Fremling 1968). In contrast, part of a population in a Nebraska, USA, reservoir required two
years to complete the cycle (Hudson and Swanson 1972). Hence there can be an extraordinary
variation in the life span of this mayfly, from about four to 24 months.
Heptageniidae
Complete voltinism data were available for more heptageniid species than for species of any
other family. Heptageniid species exhibit mainly univoltine cycles (Table 9). Of 85 species for
which there were voltinism data, more than 90% had at least one univoltine cycle. Because of
the large number of species, it is difficult to generalize further at the family level.
Rhithrogena Eaton
Unlike most other major mayfly families, no single genus of Heptageniidae accounted for a
disproportional large number of life cycle reports. Rhithrogena and Ecdyonurus populations
comprise the largest number reports of life cycles for heptageniids. Rhithrogena species are
mainly univoltine, all species for which there were adequate voltinism data having at least one
univoltine cycle, most being univoltine winter cycles. Sowa (1975a) found a R. loyalae
population of the Carpathian Mountains, Poland, to be either univoltine or semivoltine
depending upon water temperature. Rhithrogena semicolorata was the most reported
heptageniid species (Table 10). Populations of the Palearctic species exhibit almost exclusively
univoltine winter cycles; although Thibault (1971b) found a partial second generation (MBws)
for R. semicolorata (and Rithrogena iridina picteti ) in some streams in the French Pyrenees.
Neveu et al. (1979) also reported a bivoltine winter-summer cycle for R. diaphana in the
French Pyrenees.
Life cycles of mayflies (Ephemeroptera)
33
Ecdyonurus Eaton
In contrast to Rhithrogena, Ecdyonurus populations, although mainly exhibiting univoltine
cycles, were reported having a relatively large number of bivoltine (MBws) cycles. Life cycle
reports for E. venosus indicate populations having either univoltine or multivoltine cycles, but
apparently not exclusively summer cycles (Table 10). Ecdyonurus torrentis and E. lateralis
( = Heptagenia lateralis) are almost exclusively univoltine winter species. Ecdyonurus dispar
and E. submontanus have often been reported having summer cycles.
Wise (1980) summarizes ecdyonurid life cycle reports of Great Britain. Humpesch (1980)
gives data on hatching time of five Ecdyonurus species and comments on how such information
is essential for distinguishing cohorts and hence accurately interpreting the life cycle. For
example his data indicate no evidence of an obligatory egg diapause in E. dispar , despite
populations usually being reported as having summer cycles.
Heptagenia Walsh
Heptagenia populations exhibit a variety of life cycle types, most being univoltine winter
cycles. Landa (1968) reported H. sulphurea (Table 10) populations of central Europe to have a
summer generation in some years, and Ulfstrand (1968) reported populations of H. dalecarlica
in Swedish Lapland (66°N) to be either univoltine or semivoltine, depending on locality. He
also reported that, depending on hatching time, part of a H. sulphurea population could be
univoltine and the other part semivoltine, with the semivoltine component of the population
spending a year in the egg stage.
Other genera of Heptageniidae
Populations of both Epeorus and Stenonema exhibit mainly univoltine winter cycles,
although several Epeorus species and Stenonema tripunctatum (Table 10) were also reported
having bivoltine (MBws) cycles. Lehmkuhl (1968) and Richardson and Tarter (1976)
summarize life cycle data for some North American Epeorus and Stenonema species
respectively.
Populations of Stenacron interpunctatum (including all subspecies) exhibit several life cycle
types and give no indication of a characteristic type (Table 10). Of all subspecies, S.
interpunctatum canadense was the only one exhibiting univoltine cycles. McCafferty and Huff
(1978) report a complex life cycle for S. interpunctatum , with part of the population
contributing to three generations in two years, and the other part having one generation a year.
McCafferty and Huff suggest potential for genetic mixing between generations because of
differential growth among generations. The great variability in life cycles of S. interpunctatum
might indicate two or more valid species within this group.
Cinygmula, Cinygma and Ironodes nitidus{ = Epeorus nitidus) populations exhibit
exclusively univoltine winter cycles. Arthroplea , Macdunnoa , Anepeorus, and Pseudiron
populations apparently have only univoltine summer cycles. Afronurus and Thalerosphyrus are
tropical mayflies, and the few available data, mainly adult periodicity records, indicate
non-seasonal and presumably polyvoltine cycles.
Leptophlebiidae
Many leptophlebiids are tropical and this accounts for relatively large number of
non-seasonal cycles reported for this family (Table 1 1). There are few variable-type life cycles
Quaest. Ent., 1982, 18 (1-4)
34
Clifford
among the leptophlebiids. At the family level, a majority of life cycles are univoltine, due
mainly to a few abundant temperate species.
Paraleptophlebia Lestage and Leptophlebia Westwood
Generally, Paraleptophlebia populations are characterized as univoltine, having either
winter or summer cycles. All Paraleptophlebia species have at least one univoltine cycle.
Populations of the abundant Palearctic P. submarginata exhibit only univoltine winter cycles
(Table 12). In contrast, the life cycle of the Nearctic P. debilis appears more flexible (Table
12). In a brown-water stream of Alberta, Canada, P. debilis invariably exhibits univoltine
summer cycles, the nymphs appearing, growing, and emerging in about three months, June
through August (Clifford 1969). Further south, in western Oregon, USA, the nymphal
population is in the stream from about March through October (Lehmkuhl and Anderson
1971). And in New York State, USA, Gordon (1933) believed there were three generations in
two years; this implies a nymphal population during some or all the winter months.
Life cycle data indicate Leptophlebia to be almost exclusively a univoltine taxon, but these
data represent only a few abundant temperate species. Populations of the Palearctic L.
vespertina and L. marginata have only univoltine winter cycles (Table 12). The eight cycles for
L. vespertina populations represent one of the largest number of life cycles reported for a single
species in which there was no deviation from a single life cycle type. Even at high latitudes, i.e.
61°N, populations of both L. vespertina and L. marginata are univoltine with invariably an
overwintering nymphal population (Brittain 1978). The Nearctic L. cupida occurs over much
of North America, from at least 35°N to 60°N, and has been studied over several years in
Alberta, Canada (Clifford et al. 1979). Life cycle features of L. cupida have been reported
from other North American areas as well. There is no indication of L. cupida populations
deviating from a univoltine winter cycle, although emergence is progressively later at higher
latitudes in North America (Clifford et al. 1979).
Habrophlebia Eaton, Habroleptoides Schoenemund and Choroterpes Eaton
Populations of Habrophlebia , mostly H. lauta , have mainly univoltine winter cycles (Table
12). Habroleptoides modesta is apparently an entirely univoltine winter species. In contrast,
reports for Choroterpes species indicate both univoltine and multivoltine cycles, the univoltine
cycles being exclusively summer cycles. In Texas, USA, C. mexicanus is a seasonal polyvoltine
species, having three overlapping generations a year (McClure and Stewart 1976).
Other genera of Leptophlebiidae
Atalophlebiodes (= Deleatidium) sp. or spp. from New Zealand apparently can have either
univoltine winter or bivoltine cycles (Winterbourn 1974, 1978). The one life cycle report for
Traverella americana indicates a univoltine summer cycle for this Nearctic species. The other
leptophlebiids for which there were some life cycle data are tropical mayflies; and the cycles,
based mainly on adult periodicity data, appear to be non-seasonal.
Metretopodidae
Few life cycle data are available for the two genera of this small family. For the Nearctic
Siphloplecton , a life cycle study of a S. basale population from Alberta, Canada, indicates a
straightforward univoltine winter cycle (Clifford 1976). Populations of S. speciosum in Florida,
Life cycles of mayflies (Ephemeroptera)
35
USA are apparently non-seasonal (Berner 1950), although emergence data of Pescador and
Peters (1974) might indicate a seasonal cycle for S. speciosum populations in certain areas of
northern Florida. A recent life cycle study of a Metretopus borealis population in a northern
Alberta, Canada, stream indicates a univoltine winter cycle (unpublished data of Daniel Soluk,
University of Alberta). A Metretopus sp. population, probably M. borealis , also had a
univoltine winter cycle in Swedish Lapland (Ulfstrand 1969).
Oligoneuriidae
Most members of this family exhibit summer cycles (Table 5). All Oligoneuriella species,
mainly O. rhenana , have univoltine summer cycles. The two Lachlania species also have
summer cycles. In contrast, the life cycle report for Elassoneuria insulicola indicates a seasonal
univoltine winter cycle in a high altitude stream of Madagascar (Ramanankasina 1973).
Quaest. Ent., 1982, 18 (1-4)
36
Clifford
Table 8. Tabular summaries: Ephemera and Hexagenia.
Species Life Cycles
No. % Gen. % All No. % Gen. % All
Ephemera
4 23
Totals
Life cycles of mayflies (Ephemeroptera)
37
Table 9. Heptageniidae tabular summary.
Quaest. Ent., 1982, 18 (1-4)
38
Clifford
Table 10. Number of life cycles of each type reported for (1) Rhithrogena semicolorata , (2)
Ecdyonurus venosus, (3) E. torrentis, (4) E. lateralis (= Heptagenia lateralis ), (5)
Heptagenia sulphurea, (6) Stenonema tripunctatum, and (7) Stenacron interpunctatum.
Life cycles of mayflies (Ephemeroptera)
39
Table 1 1. Leoptophlebiidae and Polymitarcidae tabular summaries.
Quaest. Ent., 1982, 18 (1-4)
40
Clifford
Table 12. Number of life cycles of each type reported for (1) Paraleptophlebia submarginata,
(2) P. debilis, (3) Leptophlebia vespertina, (4) L. marginata, (5) Habrophlebia lauta, and (6)
Habroleptoides modest a.
1 2 3 4 5 6
Total Univoltine
Uw
Us
Total
Multivoltine
M?
MB
MBws
Totals
7 3 8
7-8
1
2
1
1
1
7 5 8
5 7 5
5 6 5
1
Life cycles of mayflies (Ephemeroptera)
41
Table 13. Siphlonuridae and Tricorythidae tabular summaries.
Quaest. Ent., 1982, 18 (1-4)
42
Clifford
Table 14. Number of life cycles of each type reported for (1) Ephemera danica, (2) E.
simulans , (3) Hexagenia limbata, (4) H. bilineata, (5) Povilla adusta , (6) Siphlonurus
lacustris, and (7) Ameletus inopinatus.
1 2 3 4 5 6 7
Life cycles of mayflies (Ephemeroptera)
43
Polymitarcidae
Polymitarcids exhibit several types of life cycles (Table 11). The two major Ephoron
species, the Palearctic E. virgo and the Nearctic E. album , each exhibit both univoltine winter
cycles and univoltine summer cycles. Scott et al. (1959) report Tortopus incertus populations
in Georgia, USA, to have a two-year cycle, although recently Berner (see McCafferty 1975)
suggests several broods in a year. Clearly more life cycle information is needed for this
burrowing mayfly.
Relatively extensive data for the tropical Povilla adusta indicate a non-seasonal polyvoltine
cycle (Table 14). In some areas of Africa, P. adusta adults exhibit a lunar cycle of emergence
(Hartland-Rowe 1955), but in other areas they do not (Hartland-Rowe 1958, Petr 1970).
Generation time has been estimated for several P. adusta populations and ranges from three to
possibly six months (Corbet 1957, Corbet et al. 1974, and Bidwell 1979).
Potamanthidae
Life cycle data are available for two potamanthid genera. Potamanthus populations, both
Palearctic and Nearctic species, have univoltine winter cycles; and one species, P. myops,
usually requries two years to complete the cycle in Michigan, USA (Bartholomae and Meier
1977). One report for the tropical Potamanthodes indicates a non-seasonal cycle.
Siphlonuridae
Siphlornurus Eaton, Ameletus Eaton and Isonychia Eaton
Members of this primitive and widely distributed family exhibit mainly univoltine cycles,
with a relatively large number of univoltine summer cycles (Table 13). About half of all
siphlonurid species for which there were voltinism data have at least one summer cycle.
Siphlornurus populations are characterized as having summer cycles, usually univoltine but
sometimes bivoltine. The lentic S. lacustris accounts for all univoltine winter cycles of
Siphlornurus , but some S. lacustris populations exhibit summer cycles (Table 14). Although
not indicated in the tabular summary, Bretschko (1966) and Landa (1968, 1969) suggest some
populations might occasionally have a bivoltine (MBws) cycle. Ameletus populations are also
univoltine, but with relatively more winter cycles than summer cycles. The nine life cycle
reports for the Palearctic A. inopinatus indicate an entirely univoltine winter species (Table
14). The few life cycle data for Isonychia populations give no indication of a characteristic
cycle; however, when more information is available, perhaps summer cycles will be the
characteristic type for temperate Isonychia species. Seasonal cycles for tropical Isonychia
species are not indicated.
Other Siphlonurid genera
The few life cycle data for other siphlonurids indicate summer cycles for Siphlonisca,
Parameletus and Aneletris populations. Populations of the Australian Oniscigaster wakefieldi ,
Quaest. Ent., 1982, 18 (1-4)
44
Clifford
I
perhaps one of the most primitive of recent mayflies (Edmunds 1957b), have univoltine winter
cycles (McLean 1970). Coloburiscus humeralis, another Australian species, is also a univoltine
winter species (Wisely 1965).
Tricorythidae
Except for adult records indicating non-seasonal cycles (MNP) for Tricorythus maculatus
and T. tinctus in Uganda (Tjonneland 1960), I could find complete life cycle data only for the
Nearctic-Neotropical Tricorythodes (Table 13). Generally, temperate populations of
Tricorythodes are characterized as multivoltine. Two species, T. atratus and T. minutus, have
been studied in detail. Tricorythodes atratus apparently is a summer species, with one or more
summer generations (e.g. Hall et al. 1975). In contrast, T. minutus populations appear more
flexible in their voltinism. Newell and Minshall (1978) reported T. minutus populations to have
several generations (MP) a year (with the adults on-the-wing during every month except
February) in an Idaho, USA, stream having a constant 18°C water temperature. In contrast, in
a typical stream (normal seasonal water temperature fluctuations) of the same area, the
population was bivoltine (MBws). Flight data of Tricorythodes sp. in Florida give no indication
of a seasonal cycle.
Other Families
The other families are either monogeneric or voltinism data are available for only one genus.
All members of the northern, Holarctic Ametropus (Ametropodidae) have univoltine winter
cycles. Voltinism data, available for five species, account for six univoltine cycles; A. neavei , A.
albrighti, and A. ammophilus are Nearctic; A. eatoni and A.fragilis are Palearctic. The three
species of the Nearctic Baetisca (Baetiscidae) for which there were adequate voltinism data, B.
bajkovi, B. obesa and B. rogersi, have univoltine winter cycles; the seasonal cycle is even
maintained by B. rogersi populations in Florida (Pescador and Peters 1974).
Dolania americana (Behningiidae), a burrowing mayfly, has a two-year cycle in the
southeastern USA, although most of the first year is spent in the egg stage (Harvey et al.
1980). Ramanankasina (1973) reported a Proboscidoplocia sikorai (Euthypolciidae)
population to have a univoltine winter cycle in a high elevation (ca. 1450 m) river of
Madagascar. Neoephemeropsis (Neoephemeridae) was apparently non-seasonal in Malaysia.
Palingenia (Palingeniidae) has been stated to have a three-year cycle, but this is not based on a
complete life cycle study. There is very little life cycle information available for Prosopistoma
(Prosopistomatidae). Alouf (1977) presents a growth histogram for P. oronti of Lebanon and
indicates the population probably has a univoltine cycle. There apparently are no voltinism data
for the single siphlaenigmatid species.
Synopsis
Because few reports were available from the tropics, percentages for the total mayfly fauna
(Table 1) do not reflect the true proportions of global mayfly life cycle types; they probably
give a more accurate picture of Northern Hemisphere life cycle types. For the entire fauna, the
univoltine cycle is commonest, with about three-fourth of all species for which voltinism data
are available having at least one univoltine cycle (Table 1). The most reported single life cycle
Life cycles of mayflies (Ephemeroptera)
45
type is the univoltine winter cycle. Multivoltine cycles, including the non-seasonal cycles,
account for about a third of all cycles. The semivoltine cycles and the cycles judged variable
(i.e. uni-multivoltine and uni-semivoltine) each account for less than 5% of the life cycles
reported.
The small number of variable-type cycles reported does not indicate extent of flexibility in
mayfly life cycles, because variable-type cycles include only cycles where voltinism of a given
species for a particular study was variable, depending on year, local habitat, or from the same
egg cohort. In addition, for many species (indeed, most species having several life cycle reports)
different studies indicated different types of voltinism. Even allowing for some erroneous
interpretations, there is much evidence for life cycle flexibility in many, although not all,
mayfly species. Members of Baetidae mainly Baetis and Cloeon, and Ephemeridae, mainly
Hexagenia, account for more than half of the variable life cycle reports. Within Baetidae, the
variability is mainly bivoltine (MBws)-univoltine winter cycles; within Ephemeridae,
semivoltine (2Y)-univoltine winter cycles.
Members of Baetidae, Leptophlebiidae, and Heptageniidae, account for most non-seasonal
cycles. Mayflies with non-seasonal cycles are tropical or semi-tropical. A few mayflies from
temperate regions, during certain years, were reported emerging every month of the year,
especially a few species from the French Pyrenees, western Oregon, and England. But other life
cycle features, especially nymphal growth, indicate seasonality for these mayflies. There were
no indications of seasonal cycles for mayflies placed in the non-seasonal polyvoltine type; but
many of these life cycles were based on yearly adult records, with few observations of other life
stages.
Certainly, much is to be learned about tropical mayfly life cycles, Povilla adusta being the
only tropical mayfly thoroughly studied. Generally, non-seasonality can be equated with
multivoltinism, such as done in this report, but non-seasonality, as indicated by emergence
throughout the year, would not seem to be invariably associated with multivoltine cycles.
Thorough studies of all life cycle stages of tropical mayflies, especially nymphal growth, would
be valuable and might reveal a surprisingly large number of seasonal cycles in the tropics.
SUMMER CYCLES
About a third of all mayfly species of this report exhibit at least one exclusively summer
cycle (i.e. Us, MBss, Us-MBss). The summer cycle features a long winter period of egg
dormancy, resulting in absence of the nymphal population from aquatic habitats for usually
seven or more months of the year. Such variation in periodicity, when it occurs in large
numbers of organisms, has important implications for design and interpretation of a variety of
studies pertaining to whole community processes. For example, nymphal periodicity can be a
very important parameter in estimating secondary production of aquatic invertebrates. It would
therefore seem instructive to examine summer cycles of mayflies in more detail.
Species of Baetidae (mainly Baetis and Ephemerellidae (mainly Ephemerella) account for
almost half of the exclusively summer life cycles, with Ephemerella ignita populations
accounting for almost a third of these cycles. At least 50% of the species in Oligoneuriidae,
Siphlonuridae and Tricorythidae have at least one summer cycle.
The summer cycle in mayflies seems to be mainly facultative, because most of the
thoroughly studied species exhibiting a preponderance of summer cycles, e.g. B. scambus, E.
ignita , P. debilis, also occasionally have an overwintering nymphal population. It would be
Quaest. Ent., 1982, 18 (1-4)
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Clifford
instructive to know whether the summer cycle is obligate or facultative for Oligoneuriella
rhenana, one species with several life cycle reports but none indicating an overwintering
nymphal generation.
There is evidence (Bohle 1972, Elliott 1978, and Humpesch 1980) that water temperature is
the primary factor in controlling hatching time of mayfly eggs and therefore determining
whether a summer cycle will take place. The water temperature-hatching time relationship is
not necessarily a simple one. For example, in warm lowland streams of Europe, E. ignita
nymphs are sometimes found throughout the year (in contrast to the typical summer cycle in
colder upland streams). Elliott’s (1978) combined laboratory-field study showed that these
higher water temperatures in lowland streams in autumn and winter shorten hatching time of
E. ignita eggs; whereas in the same stream, temperatures above ca. 14.5°C in spring and
summer lengthen hatching time. The overall effect of higher temperatures in lowland streams is
therefore to extend hatching time over several months.
Adaptive significance of the summer cycle in insects generally is considered to revolve
around the long egg stage, namely an adaptive response to uncertainties of harsh winter
conditions, what might be considered an adaptation to high probability events. However, many
mayfly populations exhibit summer cycles in aquatic environments where physical conditions
appear no more stressful in winter than in summer. Of course, at high altitudes or latitudes, the
possibility of stressful winter conditions, especially water freezing into the substrate, is more
obvious. In such localities, a relatively large number of mayfly populations exhibit summer
cycles (Clifford 1969, Andersen et al. 1978).
In localities where winter conditions do not appear especially stressful, adaptive significance
of the summer cycle might relate more to the short nymphal stage in summer than to the long
egg stage of winter. For example, a brief nymphal stage might minimize predation. Also,
because nymphs of summer species are usually exposed to a relatively greater autochthonous
food supply than are overwintering nymphs, one might suspect a correlation between herbivory
and summer species, in contrast to detritivory and winter species. As discussed in the
Reproductive Section, univoltine summer mayflies usually emerge and hence reproduce later in
the year than do univoltine winter species. But the adaptive significance of this temporal
isolation in the reproductive period of the two types seems obscure, since there would appear to
be very little competitive interaction for these non-feeding, short-lived adults.
EGG DORMANCY IN SUMMER
The life cycle scheme used here does not distinguish what might be considered the opposite
of the summer cycle, namely a cycle featuring a long egg stage (probably a true diapause) in
summer instead of winter. Recently Masaki (1980) summarized much of what is known about
summer diapause in temperate insects. The phenomenon is common and widespread, especially
pupal and adult summer diapause for terrestrial insects of several holometabolous orders, but
not so common for aquatic hemimetabolous insects; for example, Masaki gives only one citation
(summer egg-stage diapause) for Ephemeroptera.
I arbitrarily considered summer egg-stage dormancy to exist in Northern Hemisphere
mayflies with univoltine cycles if between April and October there were at least four months
between emergence and appearance of the new generation. Only reports where there appeared
to be routine sampling throughout the summer were considered. The four-month period would
seem a conservative criterion, because three and even two months in the egg stage would
Life cycles of mayflies (Ephemeroptera)
47
probably indicate dormancy in the sense of delayed hatching of some sort. However, it is
sometimes difficult to find very small nymphs by the usual sampling methods, especially if a
fine mesh net is not used, and one relatively long interval between the summer samples could
lead to an erroneous interpretation of the new generation’s appearance; it would therefore seem
best to be conservative in this regard.
Based on the four-month criterion, the following mayflies qualify as exhibiting summer
egg-stage dormancy. BAETIDAE: Centroptilum elsa (Lehmkuhl 1973a).
EPHEMERELLIDAE: Ephemerella dorothea (Sweeney and Vannote 1978), E. notata
(Landa 1968), E. mucronata (Landa 1968), E. subvaria (Sweeney and Vannote 1978).
Serratella deficiens (Coleman and Hynes 1970), Eurylophella versimilis (Sweeney and
Vannote 1978); CAENIDAE: Caenis rivulorum (Thibault 1971b). HEPTAGENIIDAE:
Rhithrogena morrisoni (Lehmkuhl 1970), R. hercynia (Landa 1968), R. germanica (Landa
1968), Heptagenia fuscogrisea (Landa 1968), Ecdyonurus quadrilineatus (Landa 1968).
LEPTOPHLEBIIDAE: Leptophlebia marginata (Landa 1968). SIPHLONURIDAE:
Sip hlornurus occidentalis (Lehmkuhl 1973a).
These 15 life cycles make up only a small fraction of the total 296 univoltine winter cycles
reported. However, when these summer dormancy cycles are considered as a proportion of total
winter cycles where the presence or absence of egg-stage dormancy could be deduced (175),
then the 15 cycles represent about 10% of the univoltine winter cycles; and the percentage
would be 16% if the criterion for summer dormancy was three months instead of four. Probably
even this is an underestimate of summer egg-stage dormancy, a phenomenon not very amenable
to field analysis. Also, there are instances where part of the population’s egg cohort undergoes
what would qualify as summer egg-stage dormancy; for example, the eggs of Leptophlebia
cupida in a brown-water stream of Alberta, Canada, start hatching in July, but continue to
hatch for four or five months after emergence of the old generation is completed; this has been
verified both in the field (Clifford 1969) and in the laboratory (Clifford et al. 1979).
The obvious adaptive value of summer egg-stage dormancy, be it a facultative or obligatory
phenomenon (Pleskot 1962 suggests it is facultative), would seem to involve tiding the
population over unfavorable summer conditions. The 15 life cycles were reported from only 5
localities; but of these, in only one, western Oregon (Lehmkuhl 1969), were harsh physical
conditions in summer, namely high summer temperatures and very low flow, reported.
Regardless of adaptive value of summer egg-stage dormancy in mayflies (and, as for winter
egg-stage dormancy, one could suggest several possibilities), the phenomenon appears common
enough in mayflies to merit much more study. Any phenomenon that removes the obvious
aquatic stage from streams or lakes for long periods has important implications in the analysis
of aquatic communities.
LIFE CYCLE TYPES AND PRODUCTION
The Hynes method (Hynes and Coleman 1968), or size-frequency method, in which the
secondary production estimate is based on average cohorts instead of true cohorts, is one of the
most widely used methods to estimate production of aquatic invertebrates, especially insects.
The method was originally used to estimate yearly production of the entire benthic fauna, but it
appears to be most accurate when applied to single species or groups of species with similar life
cycle features (Benke 1979, Waters 1979).
—
Quaest. Ent., 1982, 18 (1-4)
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Clifford
The yearly production estimate by the Hynes method does not account for populations with
more than or less than one generation a year. To correct for this, the usual procedure, first
proposed by Hamilton (1969), is to multiply the yearly Hynes estimate by the number of
generations per year of the population in question, e.g. two for bivoltine populations and 0.5 for
two-year semivoltine populations. Benke (1979) pointed out if the population spends a
considerable part of the year in a non-larval stage an error in the production estimate might
result by using these correction factors for bivoltine and semivoltine populations or by using no
correction factor for all types of univoltine populations. He proposed a correction factor based
on cohort production interval (CPI). This is essentially the yearly nymphal periodicity of the
population in days (or months).
I recorded nymphal periodicity (in months) for species of all life cycle types and therefore
could consider how these correction factors might influence production estimates of mayflies
exhibiting various life cycle types. The usual procedure is to apply no correction value to the
Hynes estimate when calculating production of organisms with univoltine cycles; and this
would seem valid for most univoltine winter mayflies, because the average correction factor of
the 175 univoltine winter cycles was very close to 1, namely 1.09 (x CPI = 11.0 months,
S.E. = 0.14, R = 7-15 months). Exceptions are the univoltine winter mayflies with a lengthy
egg-stage dormancy in summer (see preceding section). For these 15 univoltine winter cycles,
the average correction factor would be 1.54 (x CPI = 7.8 months, S.E. = 0.14, R = 7-8
months), and if not applied to the Hynes estimate would result in yearly production of these
univoltine winter populations being underestimated.
Not applying a correction factor to univoltine summer mayflies, assuming production
estimate is based on data collected throughout the year, would result in serious errors of
production estimates. The mean CPI of the 102 univoltine summer cycles was 4.1 months
(S.E. = 0.13, R = 2-7 months); the resulting correction factor is 2.92 and if not applied to the
Hynes estimate would result in large underestimates of production. But, these culled, average
values should be considered a tenuous substitute for empirically derived values, because the
range of CPI values for summer mayflies was two to seven months (with a tendency for shorter
CPI’s at higher latitudes and altitudes).
A feature of most univoltine summer cycles of mayflies is that nymphal growth is often easy
to interpret accurately, because hatching is synchronous and the nymphal stage is quite brief. If
sampling intervals are short, the population can usually be followed as a single cohort, and the
Hynes estimate without the correction factor or one of the definitive cohort-based methods
(Waters 1979 discusses these) would appear sufficient for a realistic estimate of production.
For the 69 bivoltine winter-summer cycles, average CPI was 9.5 months (S.E. = 0.21,
R = 7-12 months) for the winter generation and 4.1 (S.E. = 0.17, R = 2-9 months) for the
summer generation; the average CPI for the two generations would be 6.8. But this value is not
valid for calculating a yearly correction factor for bivoltine populations, because the average
CPI of both generations can not be more than six months, i.e. the average individual of one
generation must reach adulthood, mate, and lay eggs before the average individual of the next
generation begins larval development (Benke, A.D., Georgia Institute of Technology, personal
communication). Benke (pers. comm.) suggests the most accurate way to treat multivoltine
populations is to apply the Hynes estimate to each generation separately and then add the
separate values together to obtain annual production (see Menzie C.A. 1980. A note on the
Hynes method of estimating secondary production. Limnology and Oceanography 25:770-773).
Life cycles of mayflies (Ephemeroptera)
49
A modification pertinent for bivoltine winter-summer mayflies might be to calculate a
“weighed” average for the two generations, because individuals of the summer generation
usually are much smaller than those of the winter generation for a given developmental age,
perhaps by as much as 20%.
For the 10 semivoltine cycles (disregarding the three-year cycles) where total life span was
reported in months instead of years, the mean CPI was 23.8 (S.E. = 0.51) and the resulting
correction factor of 0.504 would of course indicate that the usual correction factor of 0.5 would
be valid for most two-year semivoltine mayflies. But even here, there is no substitute for
empirically derived values, as the CPI’s ranged from 21 to 27 months.
The average CPI’s and resulting correction factors are for life cycle types, not for species,
and they will usually vary much less for life cycle type than for individual species. There would
appear to be a much greater danger in using the average CPI values of this report for individual
species, especially wide-ranging species, than for life cycle types, because of the life cycle
flexibility exhibited by many species. To use an extreme example, the CPI of Hexagenia
bilineata ranged from 4.3 to 24.0 months.
LIFE CYCLE TYPES AND REPRODUCTION
Because the adult stages of mayflies are short, seldom more than three days, adult records
can generally be equated with reproduction period. Adult periodicity for mayflies is usually
derived from emergence data. However, it would seem possible, especially for mayflies with
long emergence periods, to have synchrony within the period, resulting in an “effective” adult
period, accounting for most of the successful reproduction, and a total adult period, which
might be longer than the reproductive period. I know of no studies that have investigated this,
and for purposes here, the premise is that the total adult period is equated with the total
reproductive period.
I recorded for all Northern Hemisphere mayflies having seasonal cycles the months of
emergence (or other adult periodicity data) for species of each life cycle type. Numerous other
studies report complete emergence data but not species’ voltinism (see Methods); these were
treated separately. Reproductive periods for the entire Northern Hemisphere fauna and for
each of the major life cycle types are shown in Figure 2. Reproductive periods of important
species, each representing one of the life cycle types, are shown in Figure 3. With the exception
of Leptophlebia vespertina, each of these species was occasionally reported as having other
major life cycle types as well (see General Analysis), but only adult data associated with the
species’ life cycle type in question were used in compiling the line graphs.
Each major voltinism type appears to have a characteristic seasonal reproductive period
(Fig. 2A, B). The line graph indicating the total reproductive period of all Northern
Hemisphere mayflies very much resembles a yearly water temperature curve for
north-temperate regions (Fig. 2C). Almost half of mayfly reproduction in the Northern
Hemisphere takes place during June and July; obviously when considered on a hemisphere
basis, the word mayfly is a misnomer. The multivoltine mayflies have the most extensive
reproductive period (Fig. 2A). (The commonest multivoltine cycle is the bivoltine
winter-summer cycle, and the two minor peaks probably reflect the maximum reproductive
period for each of these generations). The semivoltine mayflies appear to have the most
restrictive total reproductive period (Fig. 2A). Populations of univoltine summer mayflies
generally reproduce later than do the univoltine winter mayflies (Fig. 2B). The total
reproductive periods of individual species, each exhibiting a major life cycle type, were quite
similar to those of the life cycle types (Fig. 3).
Quaest. Ent., 1982, 18 (1-4)
% Total Months of Each Type
50
Clifford
Months
Figure 2. Percentage monthly reproductive period (equated with adult periodicity) for Northern Hemisphere mayflies of
each life cycle type (based on complete voltinism data) and all Northern Hemisphere mayflies, regardless of life cycle type
(based on complete voltinism data and also data of studies pertaining exclusively to adult periodicity). Total months are
the total number of months that all species of a particular life cycle type were reported as adults; e.g. in January, there
were 14 reports (14/632 = 2.2%) of multivoltine species being reported in the adult stage.
% Total Months of Each Species
Life cycles of mayflies (Ephemeroptera)
51
Months
Figure 3. Percentage monthly reproductive period for four species (throughout their range), each representing one life
cycle type.
Quaest. Ent., 1982, 18 (1-4)
52
Clifford
Using words similar to those used by Shapiro (1975) in discussing voltinism of butterflies, it
would appear the semivoltine cycles of mayflies are associated with reproductive periods being
shut down for the adverse season considerably before the average phenological properties of
Northern Hemisphere environments would dictate they would have to be. In contrast, the
reproductive period of multivoltine mayflies suggests reproduction continuing until
deteriorating environmental conditions intervene.
It seems unusual that mayflies with the longest generation time, the semivoltine species,
should have, when considered for all Northern Hemisphere localities, the shortest reproductive
period, while mayflies with the shortest generation time, the multivoltine species, have the
longest reproductive period. The adaptive strategy of the semivoltine cycle might be to
minimize predation on adults, whereas the multivoltine cycle would minimize predation on
nymphs. Edmunds and Edmunds (1980) discuss how adult periodicity patterns of mayflies
might have been influenced by selective pressures of predators. Mackey (1978) discusses this
specifically for Caenis species, one, C. robusta , having a long emergence period and another, C.
macrura, having a short emergence period.
LIFE CYCLE TYPES AND OTHER FACTORS
There were no detectable trends between life cycle types and the various suggested
phylogenetic schemes of extant myflies (e.g. Demoulin 1958, Edmunds 1972). No clear
relationship between life cycle type and body size of individual species was noted. Many very
small mayflies species, e.g. some baetids, caenids, tricorythodids, are multivoltine, and almost
all the semivoltine species are relatively large mayflies, e.g. ephemerids; but numerous other
large mayflies are not semivoltine. For mayflies of intermediate size no relationship between
body size and life cycle type was evident. Total life spans (from hatching of eggs until death of
the adult) vary from 3 years for Ephemera danica (Svensson 1977) to about 30 days, reported
for the non-seasonal Callibaetis floridanus (Trost and Berner 1963) and the seasonal
Centroptilum pennulatum (Macan 1978). Assuming a minimum egg incubation of 15-30 days
for mayflies with very short life spans, in theroy, at least, it would seem possible to have a
maximum of six or more generations per year. No study, even for non-seasonal populations, has
clearly demonstrated more than six generations a year, which appears to be maximum for a
population of the non-seasonal Povilla adustasta, although its populations usually had only four
generations a year (Corbet et al. 1974). There have been suggestions of as many as six
generations for seasonal mayflies; but for these, cohorts could not definitely be distinguished
from generations.
For each life cycle study, latitude of the area in question was recorded, the hope being these
latitudinal values when compared to life cycle types of that area would be amenable to analysis.
However, variations in local conditions, e.g. continental vs. marine climates, altitudinal
influences, etc., made such an analysis too difficult to interpret on a global basis. Nevertheless,
mayfly life cycles ought to change in a predictable way going from tropics to arctic (Clifford et
al. 1973).
In the tropics, mayflies exhibit mainly non-seasonal multivoltine cycles. In moderate
temperate regions, the univoltine winter cycle is most common; however, all major life cycle
types can be locally important. The univoltine winter cycle was the most reported of any life
cycle type and probably in part reflects most life cycle studies in moderate temperate regions.
In cold temperate and subarctic regions (disregarding areas influenced by marine climates)
Life cycles of mayflies (Ephemeroptera)
53
there are fewer multivoltine cycles, relatively more semivoltine cycles, and a tendency for
univoltine winter cycles to give way to the univoltine summer cycle. In the low and high arctic
of continental climates, where mayflies species are few, the univoltine summer cycle
predominates. Several terrestrial insects and even a few aquatic insects can have
extraordinarily long semivoltine cycles in the arctic, while the same species are univoltine at
lower latitudes. Semivoltinism is not a feature of arctic mayflies (the major semivoltine group,
the ephermerids, are not found in the high arctic). Possibly, however, future studies, especially
of the egg stage, may reveal semivoltine cycles for some of the arctic baetids and heptageniids,
which, based on nymphal growth, have usually been reported as univoltine summer mayflies.
For example, Ulfstrand (1968) reported Heptagenia sulphurea to be partly semivoltine at 65°,
with one year probably spent in the egg stage.
DISCUSSION
The survey clearly indicates extensive life cycle flexibility for many mayflies. Admittedly,
most species were reported as having only one life cycle type, but there were usually only one or
two complete life cycle studies for most species. The more thoroughly a species is studied
throughout its range, the more flexible its life cycle usually appears. In fact, the variability that
most species show in their life cycle types appears to be directly correlated with the number of
studies reported for that species. Several workers, e.g. Ulfstrand (1968), Lehmkuhl (1973a),
Flowers and Hilsenhoff (1978), Whelan (1980), have commented on plasticity of mayfly life
cycles for certain species, but most workers probably feel there is an inherent rigidity in the life
cycle of most species— at least to where the type voltinism does not change.
The bet-hedging and risk-spreading strategies of a flexible life cycle seem so obvious (e.g.
partitioning of different foods among different sizes of nymphs, advantages of several life stages
being present if catastrophic events occur, etc.) that one might question how temperate
mayflies with seemingly rigid life cycles manage not only to survive, given the uncertainties of
the environment, but to be very abundant in certain areas. Populations of Ameletus inopinatus
and temperate Leptophlebia are examples of this. In Alberta, Leptophlebia cupida is a very
abundant mayfly; the species apprently always has a univoltine winter cycle, from year to year
and from place to place. Life cycle data for L. cupida from other areas of North America also
indicate a rigid univoltine winter cycle. And there are numerous life cycle reports for the
Palearctic L. vespertina and L. marginata, all indicating a univoltine winter cycle. Clearly, the
life cycle strategies of these successful Leptophlebia taxa must involve something other than
flexibility in voltinsim.
It is important to distinguish cohorts from generations. In many populations, the generation
is essentially a single cohort; but in other populations there can be several groups of nymphs,
each about the same age, growing at the same rate and transforming at the same time. Not
accounting for cohorts can result in erroneous interpretations of life cycles; e.g. an apparent
polyvoltine species where in fact there is a single generation of several cohorts. Where there
were not sufficient data to distinguish cohorts from generations, some of the life cycles reported
as univoltine winter cycle with a long emergence period could just as well have been interpreted
as bivoltine winter-summer cycles; alternately, some of the bivoltine winter-summer cycles
could have been interpreted as univoltine winter cycles, with the eggs hatching in pulses and
resulting in cohorts. One criterion often used to distinguish the two generations of the bivoltine
cycle is that the adults of the summer generation are often much smaller than the winter
Quaest. Ent., 1982,18 (1-4)
54
Clifford
generation adults. However, adults of univoltine mayflies transforming during the latter part of
a long emergence season can also be much smaller than adults emerging early.
It is not always clear what is meant by a cohort. There are different gradations of cohorts.
Ideally, nymphs of the same cohort will all transform at about the same time. Cohorts of this
type might be so separated in time that it is questionable whether genetic mixing between
cohorts of the same generation takes place. Sometimes there is a semblance of nymphal cohorts
within the population, but the nymphs of each cohort tend to accumulate in a single
potential-emerging-stage and then emerge as a single cohort (Clifford 1970a). In other species,
the distinction between cohorts and generations is completely blurred; the population’s eggs
hatch at different rates, nymphs grow at different rates, and adults emerge and hence
reproduce throughout the summer. An arbitrarily designated cohort in this type population
might, at least in theory, contain nymphs of different generations. In short, the concept of
distinguishing cohorts from generations is an important one; but, for mayflies (a group of
insects that usually has a large and sometimes variable number of molts, which will further
compound cohort analysis) in practice it would often seem very difficult, certainly without
supplementary laboratory observations. Pritchard (1978) recently proposed a cohort-splitting
hypothesis for the holometabolous cranefly Tipula. Future modifications of this hypothesis may
result in valuable information useful in analysing cohorts of mayflies.
FURTHER RESEARCH
Little life cycle information is available for several taxa frequently reported as locally
abundant in standing crop studies, occasionally used as experimental animals, or occupying an
interesting phylogenetic position. For the Northern Hemisphere fauna, these include:
Callibaetis and Heterocloeon (Baetidae); Dannella (Ephemerellidae); Pentagenia and
Lithobrancha (Ephemeridae); Cinygma, Anepeorus , Pseudiron, and Spinadis
(Heptageniidae); Traverella and Habroleptoides (Leptophlebiidae); Metretopus
(Metretopodidae); Neoephemera (Neoephemeridae); Homoeoneuria (Oligoneuriidae);
Palingenia (Palingeniidae); Tortopus and Campsuarus (Polymitarcidae); Prosopistoma
(Prosopistomatidae); Isonychia (Siphlonuridae) and Leptohyphes (Tricorythidae). Much more
life cycle information is also needed for all taxa of tropical mayflies, basing type voltinism on
egg stage and nymphal growth data, as well as on adult periodicity.
A small number of genera in a few families usually account for most of the reports
pertaining to number of animals per unit area of substrate (numerical abundance studies). For
example, for the Holarctic region, species of Baetis, Ephemerella , Rhithrogena , Hexagenia ,
Stenonema-Stenacron, Caenis and Paraleptophlebia made up about 80% of the numerical
abundance values reported in the literature (Clifford 1980). Concentrating life cycle studies on
the abundant species of these genera might be a mGre feasible objective, in the sense of
obtaining data about important mayfly contributors to the community, than attempting to work
out the life cycle of every mayfly species in the community. In North America, Hexagenia
populations are often an important biomass component of the community. Although there are
considerable life cycle data for Hexagenia (though sometimes only reported at the generic
level), more information on how the life cycle of the important Hexagenia species vary from
region to region and year to year would be valuable.
Studies on how the various mayfly life cycle types relate to different methods of estimating
secondary production would seem especially pertinent. Related to this is ability to distinguish
Life cycles of mayflies (Ephemeroptera)
55
cohorts from generations. This is often not easy, but perhaps new methods will be developed to
solve this problem (see, for example, the model relating egg hatching time to temperature
presented by Elliott and Humpesch 1980).
The mayfly’s egg stage is the least understood of the major life stages. Hatching time and
other egg stage phenomena, e.g. predation on eggs (especially in cycles featuring a long egg
stage), are not easy to study in the field. But a thorough understanding of the egg stage
specifically and the role water temperature plays generally in influencing all life stages are the
keys to better interpreting mayfly life cycles.
ACKNOWLEDGMENTS
I am grateful to Dr. John E. Brittain, University of Olso for reading a draft of the
manuscript and providing helpful suggestions and to Dr. Arthur C. Benke, Georgia Institute of
Technology, for helpful suggestions on the Production section. I thank Mrs. Robert Baker for
technical assistance. Support was provided by the National Research Council of Canada.
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APPENDIX-Species literature citations (by number) and consensus life cycles.1
Ametropodidae
Ametropus albrighti Traver 8 Uw
A. amnophilus Allen and Edmunds 8 Uw
A. eatoni Brodskij 777 Uw
A.fragilis Albarda 196 Uw
A. neavei McDunnough 66,208 Uw
Baetidae
Baetiella sp. 277 MBws
B. japonica Imanishi 124 MB?
Baetis sp. 51,166,277,306,37 1 MBws
B. alpinus (Pictet) 158, 160,161, 180, 196, 321, 323,330, 341 see text
B. amplus (Traver) 51,250
B. atrebatinus Eaton 143,796 MBws
B. australis Traver 27
B. beskidensis Sowa 321 MBss
B. bicaudatus Dodds 191, 279 MBws
B. brunneicolor McDunnough 24,217 Us-MBss
B. buceratus Eaton 796,199,200,327 MBws
B. ephippiatus Traver 27
1 Italicized citations indicate some voltinism data available; others refer to strictly adult
periodicity, see Methods.
Quaest. Ent., 1982, 18 (1-4)
Clifford
B.flavistriga McDunnough 24,217,22(5,250,289,327 MBss
B.frondalis McDunnough 24 MBss
B.fuscatus (L.) (=B. bioculatus) 1 16,1 17,143,79(5,799,200,
26 1 , 295,296,321,323,354,355,368 MBss
B. gemellus Eaton 1 17,261,540 MP
B. gracilis Bogoescu and Tabacaru 321 Us
B. hageni Eaton 24,(59,216 MBws
B. intercalaris McDunnough 24,27,217 Us
B. jesmondensis McDunnough 191
B. lapponicus (Bengtsson) 10,354,355 Us
B. lutheri MUller-Liebenau 161 ,321 ,323 no consensus
B. macani macani Kimmins 10,41,42,43,44,45,355 Us
B. macani bundyae Lehmkuhl 210 Us
B. macdunnoughi Ide 24,69, 250 MBws
B. melanonyx (Pictet) 261,527 Us
B. meridionalis Ikonomov 261
B. muticus (L.) ( = B . pumilus ) 70,103,1 16,1 17,121,143,796,799,200,225,257,310,527,
323,330,339,340,354,355,368 MBws
B. niger (L.) 10, 99, 128, 143, 153, 796,314, 327„346 MBws
B. ochris Burks 289
B. persecuta McDunnough 737,191 MBws
B. propinquus (Walsh) 24,17,57,217,234 MBss
B. pygmaeus (Hagen) 24,27,217,270,284,289 MBws
B. rhodani (Pictet) \0, 18, 20, 35, 37 ,43,44,99,102,103,1 16,121,733,739,143,153,767,
164,173,174,196,199,200,202,228,231,252,273,295,296,310,
314,317,. 320,321,. 323, 330,340,344,345,346,354, 355 see text
B. scambus Eaton 1 1 ,99, 103,\2\, 143,164,196,321 ,323,340,368 see text
B. sinaicus (Bogoescu) 327,323 MBss
B. soror Ulmer 337 MBws
B. subalpinus Bengtsson 21%, 29 5, 29 6, 3 5 4, 355 no consensus
B. tricaudatus Dodds ( = B . vagans ) 24,32,60,69,73,7 5,169,209,216,2X1 ,249,262,359,360
see text
B. vardarensis Ikonomov 327,323 MBws
B. vernus (Curtis) ( = 77. tenax ) 70,1 1 6, 1 2 1 , 1 43, 1 53,7 64, 1 74,7 75, 1 84,7 96, 1 99,267 ,
295,296,310,323,346 see text
B. yamotoensis Gose 1 25
Baetodes sp. 93 MNP
Callibaetis sp. 154,269
C. coloradensis Banks 32,60,137 M?
C. ferrugineus (Walsh) 140,217
C.floridanus Banks 27,81 ,351 MNP
C. praetiosus Banks 27, 1 1 5 MNP
Centroptilum sp. 32,51,60,737,167 Us
C. album McDunnough 217
C. bellum McDunnough 51
C. corbeti Kimmins 347 MNP
Life cycles of mayflies (Ephemeroptera)
79
C. convexum Ide 2 1 7
C. elsa Traver 209 Uw
C. fragile McDunnough 226
C. luteolum (Muller) 28,55,47,1 16,1 17, 128, 143, 176, 184, 196,203, 232, 234,260, 278, 310,
311,31 4,327 ,323,346,355,353 MBws
C. notabile Kimmins 347 MNP
C. pennulatum (Eaton) 1 43,7 95,232,260,3 1 1 ,327 no consensus
C. pulchrum Eaton 327 MBss
C. quaesitum McDunnough 51
C. rivulare Traver 51
C. rufustrigatum McDunnough 217,250,289,336 Us
C. simile McDunnough 299
C. viridocularis Berner 27 MNP
C. walshi McDunnough 25 MBss
Cloeon sp. 217
C. dentatum Kimmins 347 MNP
C. dipterum (L.) 35,47,47,52,53,73,95,725,143,153,775,184,737,795,200,223,237,258,
313,321,346 see text
C. rubropictum McDunnough 27,40,51,226,289 MNP
C. samaeleni Gillies 40
C. simile Eaton 35,47,128,143,153,795,237,258,327,346,355 see text
C. simplex McDunnough 140
C. triangulifer McDunnough 773,327 MP
Dactylobaetis sp. 93
Heterocloeon sp. 93
Paracloeodes sp. 93
Procloeon bifidum (Bengtsson) \2S, 176, 196, 232,3 21 ,322 see text
P. ornatum Tshernova 327,322 MBss
Pseudocloeon sp. 51,137,140
P. alachua Berner 27 MNP
P. anoka Daggy 217
P. bimaculatum Berner 27 MNP
P. Carolina Banks 250
P. dubium (Walsh) 57,226
P. inexpectatum Tshernova 327 MBss
P. myrsum Burks 27 1
P. parvulum McDunnough 27 MNP
P. punctiventris McDunnough 27,271 MNP
Baetiscidae
Baetisca bajkovi Neave 208 Uw
B. becki Schneider and Berner 283
B. laurentina McDunnough 217
B. obesa (Say) 57,207,217,226,234 Uw
B. rogersi Berner 23,234 Uw
Quaest. Ent., 1982, 18 (1-4)
80
Clifford
Behningiidae
Behningia ulmeri Lestage 182
Dolania americana Edmunds and Traver 749,288 2Y
Caenidae
Brachycercus sp. 16 Us
B.flavenus Traver 140
B. harisellus Curtis 72,178,193,796,227 Us
B. lacustris (Needham) 217,226 Us
B. maculatus Berner 21,284 MNP
B. minutus Tshernova 196,321 U?
B. pallidus Tshernova 196,321 U?
B. prudens (McDunnough) 208 Us
Caenis sp. 31,166
C. arnica Hagen 5 1
C. beskidensis Sowa 321,323 Us
C. delicata Traver 238
C. dimulata Walker 27,57,775,284 MNP
C. forcipata McDunnough 106,217,265 Uw
C. hilaris (Say) 27,284 MNP
C. horaria (L.) 47,128,143,153,776,178,796,226,252,258, 278,314,346 see text
C.jocosa McDunnough 217
C. Burmeister 796 Us
C. luctuosa Burmeister ( = C. moesta) 22,47,143,796,260, 278,309 no consensus
C. macrura Stephens 7 76,143,178,796,200,236,227,240 MBws
C. noturna Bengtsson 278
C. pseudorivulorum Keffermuller 196,321 Us-MBss
C. rivulorum Eaton 77,702,178,184,237,272,227,240 Uw
C. robusta Eaton 24,178,796,236 no consensus
C. simulans McDunnough 32,722,727,140,217,226,289,206,336 no censensus
Tasmanocoenis tillyardi (Lestage) 227 MBws
Ephemerellidae
Attenella margarita (Needham) 4
Danella simplex (McDunnough) 5,217,246,284,329
Drunella coloradensis (Dodds) 6,98,122,207,277 Uw
D. cornuta (Morgan) 217,254
D. doddsi (Needham) 6,122,191,207,277 Uw
D. flavilinea (McDunnough) 6,137,148 Us
D. grandis grandis (Eaton) 6
D. grandis ingens (McDunnough) 6,371 Uw
D. lata (Morgan) 217
D. spinifera (Needham) 6,16,137 Uw
D. walkeri (Eaton) 217
Ephemerella aurivillii Bengtsson 14,202,218,220,321 ,254,355 Uw
E. basalis Imanishi 14,125,272,277,227 Uw
Life cycles of mayflies (Ephemeroptera)
81
E. catawba Traver 51
E.-choctawhatchee Berner 284
E. dentata Bajkova 218 Uw
E. dorothea Needham 51,217 ,336,357 Uw
E. excrucians Walsh <59,2 1 7 Uw
E. gracilis Tshernova 218
E. ignita (Poda) 1 1 ,1 2,1 3,14,17,46,50,18,82,84,95,96,99,100, 102, 103,1 75,117 ,
121, 129, 143, 164, 173, 178, 185, 196,199, 200, 201, 203, 218, 220, 227,
228, 231, 233, 237, 239, 260,272,273,282,290,295,296, 3 1 0,3/ 2,3 1 4,
321,323,330,239,340,364,368 see text
E. inermis Eaton 7,16,122,7 37,148,37 1 Uw
E. infrequens McDunnough 7
E. invaria (Walker) 140,217
E. kozhovi Bajkova 14 ,218 Us
E. lacustris Allen and Edmunds 7
E. latipes Tshernova 218 Uw
E. lenoki Tshernova 218
E. lepnenae Tshernova 218 Uw
E. levanidovae Tshernova 218 Uw
E. longicaudata Tshernova 125
E. mesoleuca (Brauer) 196,321 Us
E. mucronata Bengtsson (=E. krieghoffi ) 196,218,220,3X0,321 ,354,355 Uw
E. needhami McDunnough 217
E. nigra Ueno 125,352 U?
E. notata Eaton 1 16,143,196,321 ,368 Uw
E. rufa Imanishi 125 MBws
E. rotunda Morgan 217
E. setigera Bajkova 218
E. subvaria McDunnough 54,2X1 ,265,336,362 Uw
E. ts her novae Bajkova 218
E. triacantha Tshernova 218,321 Us
E. trispina Ueno 1 25,277 ,331 ,352 Uw
E. yoshinoensis Gose 125 ,277,352 Us
E. zapekinae Bajkova 14,218
Eurylophella sp. 51
E. bicolor (Clemens) 51,226 Uw
E.funeralis (McDunnough) 104,1 36,336 Uw
E. lutulenta (Clemens) 51,2X1,226 Uw
E. minimella (McDunnough) 69
E. prudentalis (McDunnough) 51
E. temporalis (McDunnough) 106,140,217,226 Uw
E. trilineata (Berner) 27,284
E. versimilis (McDunnough) 336 Uw
Serratella deficiens (Morgan) 69,2X1 ,336 Uw
S.frisoni (McDunnough) 271
S. serrata (Morgan) 254 ,336 Us
Quaest. Ent., 1982, 18 (1-4)
82
Clifford
S. sordida (McDunnough) 289
S. tibialis (McDunnough) 737,191,577 Us
Teloganodes sp. 31 MNP
Timpanoga hecuba pacifica (Allen and Edmunds) 3
Torleya major (Klapalek) (=T. belgica) 1 29 ,196,260,296,321 ,323,339,340 Uw
Ephemeridae
Ephemera sp. 31
E. amurensis Navas 218 Uw
E. blanda Traver 51
E. danica Muller 1 17,143,773,776,178,796,237,280,281,295,296,310,327,323,332,
333,339,340,363 see text
E. formosana Ulmer 218
E. guttulata Pictet 183,243,299
E. japonica McLachlan 352 Uw
E. lineata Eaton 181,188,796,327 2Y
E. sachalinensis Matsumura 218
E. simulans Walker 27 ,33,49,57 ,69, 106,1 14,737,140,168,203,217,226,243,249,
263,289,304,324 see text
E. strigata Eaton 2 1 8
E. varia Eaton 299
E. vulgata L. 143,153,178,737,188,796, 260,278,314,327,346 2Y
Hexagenia sp. 51,77,1 1 1,225
H. atrocaudata McDunnough 217
H. bilineata (Say) 27,57,703,109,1 10,772,757,224,243,287,304,343 Uw
H. limbata Serville ( + 7/. limbata limbata) 32,76,77,93,705,106,109,140,755,756,
757,763,217,226,243,264,265,289,303,
325,334,312 see text
H. limbata occulata Walker 51,140,264,304,325 2Y
H. limbata viridescens (Walker) 140
H. munda Eaton 27,243
H. munda affiliata McDunnough 140
H. munda elegans Traver 27,51
//. munda marilandica Traver 27,51
H. munda orlando Traver 27
H. rigida McDunnough 705,140,217,243,264 2Y
Litobrancha recurvata (Morgan) 93,217,242,243,255,256,257,266,326
Pentagenia vittigera (Walsh) 109,1 13,243,289
Euthyplociidae
Campylocia sp. 93
Euthyplocia hecuba (Hagen) 93
Proboscidoplocia sikorai Vayssiere 302 Uw
Heptageniidae
Afronurus ugandanus Kimmins 347 MNP
Life cycles of mayflies (Ephemeroptera)
83
Anepeorus sp. 28
A. rusticus McDunnough 208 Us
Arthroplea bipunctata McDunnough 95,707,140 Us
A. congener Bengtsson 755,796,355 Us
Cinygma sp. 277 Uw
C. integrum Eaton 191,275 Uw
Cinygmula altaica Tshernova 218 Uw
C. grandifolia Tshernova 218 Uw
C. hirasana Imanishi 218
C. malaisei Ulmer 279 Uw
C. mimus (Eaton) 98,137 Uw
C. ramaleyi (Dodds) 148,301 Uw
C. reticulata McDunnough 191,274 Uw
Ecdyonurus affinis Eaton 196,321 no consensus
E. angelieri Thomas 203,339,340 Uw
E. aurantiacus (Burmeister) ( = E . fluminum) 796,260,527 Us
E. carpathicus Sowa 527 Uw
E. dispar (Curtis) 143, 162,196,234,. 321 ,323,368 Us
E.fascioculatus Sowa 527 Us
E.forcipula (Pictet) 180,796 Uw
E. insigmis (Eaton) 143,796,200,527,525 Us
E. joernensis Bengtsson 355
E. kibunensis Imanishi 352
E. lateralis (Curtis) ( =Heptagenia lateralis ) 1 39,\43, 164,178, 196,228,231 ,310,
527,525 Uw
E. macani Thomas and Sowa 527 Uw
E. picteti Meyer-Diir. 159,196 Uw
E. quadrilineatus Landa 196,198,321 Uw
E. starmachi Sowa 527 Uw
E. subalpinus Klapalek 180,796,527 no consensus
E. submontanus Landa 196,198,321 Us
E. tobironis Takahashi 218
E. torrentis Kimmins 121,7 38,143,196,321 ,323,368 Uw
E. venosus (Fabricius) 99,705,1 17,143,764,180,796,257,260,505,527,339,540,565
see text
E. yoshidae Takahashi 125,352 Uw
Epeorus sp. 3 1 ,5 1 ,7 57, 1 9 1 ,208 no consensus
E. aesculus Ueno 218,337
E. albertae McDunnough 90,204
E. deceptivus (McDunnough) 90,204,301 Uw
E.fragilis (Morgan) 241,254
E. ikanonis Takahashi 125,277 Uw
E. latifolium Ueno 725,218,277,552 MBws
E. longimanus (Eaton) 90,745,179,204,301,577 Uw
E. maculatus Tshernova 218
Quaest. Ent., 1982, 18 (1-4)
84
Clifford
E. pleuralis (Banks) 241,257 Uw
E. rubidus (Traver) 51
E. sylvicola Pictet ( = E. assimilis) 180,795,260,527,525 Uw
E. torrentium Eaton 339 ,545 MBws
E. uenoi Matsomura 1 25,277,352 no consensus
E. vitreus (Walker) 757,777,217,254,299 Uw
Heptagenia abnormis Tshernova 218
H. aphrodite McDunnough 51,140
H. arsenjivi Tshernova 218
H. coerulans Rostock 196,321 Uw
H. dalecarlica Bengtsson 354, 355 Uw-2Y
H. diabasia Burks 107 Us-Uw
H. elegantula (Eaton) 93
H.flava Rostock 178,795,275,527 Uw
H. flavescens (Walsh) 56,757,289 Uw
H.fuscogrisea (Retzius) 79,47,143,153,178,795,355 Uw
H. hebe McDunnough 757,217 Us-Uw
H. inconspicua McDunnough 289
H. juno McDunnough 299
H. kibunensis Imanishi 218
H. longicauda (Stephens) 143,527 Uw
H. lucidipennis (Clemens) 56,757,225 Us
H. maculipennis Walsh 56,289
H. pulla (Clemens) 56,107,140,171,217
H. Soldatova Tshernova 218
H. sulphurea (Muller) 143,178,795,200,257, 231 ,321 ,323,354,355
H. umbricata McDunnough 289
H. yoshidae Takahashi 218
Ironodes nitidus (Eaton) 93,191,254 Uw
Macdunnoa nipawinia Lehmkuhl 272 Us
Pseudiron centralis McDunnough 208 Us
Rhithrogena sp. 757,191,275 Uw
R. alpestris Eaton 795 Us
R. arnica Traver 299
R. brunnea (Hagen) 122
R. diaphana Navas 7 96,273,321 ,323 Us
R. ferruginea Navas 527,525 Uw
R. germanica Eaton ( =R . haarupi) 143,795,527 ,525,555, Uw
R. gorganica Klapalek 527 Uw
R. hageni Eaton 557 Uw
R. hercynia Landa 196,198,321 323 Uw
R. hybrida Eaton 196,321 Uw
R. impersonata (McDunnough) 757,217 Uw
R. iridina iridina Kolenati 527 Us-Uw
R. iridina picteti Sowa 339,545 MBws
R. japonica Ueno 125,277 Uw
see text
Life cycles of mayflies (Ephemeroptera)
85
R. jejuna Eaton 56,707,217 Uw
R. loyale Navas 79(5,527,341 no consensus
R. minus Eaton 267
R. morrisoni (Banks) 206 Uw
R. pellucida Daggy 707,217 Us
R. sanguinea Ide 217
R. semicolorata (Curtis) 75,4(5,9(5,705,1 17,121,755,180,796,225,229,257, 527,525,
330,339,540,565 see text
R. undulata (Banks) 707 Uw
R. unicolor Tshernova 275 Uw
R. virilis McDunnough 148,371 Uw
Spinadis sp. 107
S. wallacei Edmunds and Jensen 92
Stenacron interpunctatum canadense (Walker) 32,56,69,102,707 Uw
S. interpunctatum frontale (Banks) 27,140,217
S. interpunctatum heterotarsale (McDunnough) 140,217,299
S. interpunctatum (Say) (+S. interpunctatum interpunctatum) 51,55,707,
137,2X1,226,
244,284,289,370
see text
Stenonema sp. 51
S. bipunctatum (McDunnough) 707,140 Uw
S. exiguum Traver 27,107,284 no consensus
S. femoratum (Say) 226,289 MP
S.fuscum (Clemens) 707,140,217 Uw
S. integrum (McDunnough) 51,707,217 Uw
S. ithaca (Clemens and Leonard) 217
S. luteum (Clemens) 56,217
S. mediopunctatum (McDunnough) 707 Uw
S. nepotellum McDunnough 140,217
S. pudicum (Hagen) 51
S. pulchellum (Walsh) 707,217 Uw
S. rubromaculatum (Clemens) 56,289
S. rubrum (McDunnough) 707,140,217 Uw
S. smithae Traver 27,284 MNP
S. terminatum (Walsh) 707 Uw
S. tripunctatum (Banks) 55,56,707 ,77 7 ,217 ,226,305 see text
S. vicarium (Walker) 69,70,217 ,305 Uw
Thalerosphyrus sp. 31 MNP
Leptophlebiidae
Adenophlebiodes decoratus Kimmins 347 MNP
Atalophlebiodes sp. ( = Deleatidium) 366,367 Uw-MBws
Calliarcys humilus Eaton 85,286
Choroterpes sp. 57,144,145 MNP
Quaest. Ent., 1982, 18 (1-4)
Clifford
C. albiannulata McDunnough 203 Us
C. basalis (Banks) 51,226 Us
C. bugandensis (Kimmins) 347 MNP
C. crocatus Allen 1
C. curtis (Kimmins) 347,348 MNP
C. elegans (Barnard) 286
C. hubbelli Berner 27
C. mexicanus Allen 1 ,245 MP
C. nigrescens Barnard 286
C. /wctefi (Eaton) 121,196,321 Us
C. trifurcata Ueno 725 MBws
Cryptopenella facialis Gillies 1 19,286
Dipterophlebiodes sp. 31 MNP
Gilliesia hindustanica (Gillies) 1 19,286
Habrophlebia sp. 51
H.fusca (Curtis) 1 17,793,794,796,237,310 Uw
H. lauta McLauchlan 97,1 \1 ,193,194,196,228,260,294,295,296,310,321 ,323,339,
340 Uw
H. vibrans Needham 51
Habrophlebiodes sp. 31 MNP
H. americana (Banks) 51
H. brunneipennis Berner 27
H. gilliese Peters 286
H. prominens Ulmer 285,286,356
Habroleptoides sp. 31,286
H. modesta (Hagen) 1 17 ,1 29,196,260,294,295,296,314,321 ,323,339,340 Uw
Isca purpurea Gillies 1 19,286
Leptophlebia austrina (Traver) 51
L. cupida (Say) 29,32,57 ,60,62,64,67 ,68,69,137 ,141,142,7 50, 151, 152,169,
170,217,226,237,254,255,264,289,325,535,349 Uw
L. intermedia (Traver) 27
L. marginata (L.) 79,39,47,43,44,45,128,143,178,184,796,227,230,237,258,278,
289,319,342,355 Uw
L. nebulosa (Walker) 57,217
L. pacifica (McDunnough) 209 M?
L. vespertina (L.) 23,39,47,43,44,45,143,153,178,736,737,796,237,253,258,278,319,
321,355 Uw
Paraleptophlebia sp. 51,154
P. adoptiva (McDunnough) 74,1 23,172,211,299 Uw
P. bicornuta (McDunnough) 191,275 Us
P. bradleyi (Needham) 27
P. chocolata Imanishi 125,218 no consensus
P. cincta (Retzius) 143,178,796,310,327 no consensus
P. debilis (Walker) 32,60,723,737,191,275,217,328 see text
P. gregalis (Eaton) 191
P. guttata McDunnough 51,123,289,299
Life cycles of mayflies (Ephemeroptera)
87
P. heteronea (McDunnough) 191
P. lunata Tshernova 218
P. moerens (McDunnough) 69, 123,250 Uw
P. mollis (Eaton) ^9,75,74, 1 23,7 72, 1 90,2 1 7 Uw
P. praepedita (Eaton) 32,57,217,254
P. spinosa Ueno 125,352 U?
P. stradii Eaton 128,355
P. strigula (McDunnough) 217
P. submarginata (Stephens) 103,1 29,\43, 118,196, 203, 228,231 ,260,327 ,339,340 Uw
P. temporalis (McDunnough) 215 Uw
P. volitans (McDunnough) 27,172,284 MNP
P. werneri Ulmer 118,196 Us
Simothraulus seminiger Ulmer 286,356
Thraulodes sp. 28 MNP
Thraulus sp. 31
T. bellus Eaton 85,286,358
T. fasciatus (Kimmins) 286,287,347
T. torrentis (Gillies) 286,287
Traverella sp. 93
T. albertana (McDunnough) 28 ,208 Us
Metretopodidae
Metretopus sp. 355 Uw
M. borealis Eaton 93,128,178
Siphloplecton basale (Walker) 32,57,63,137,152,217,226 Uw
S. speciosum Traver 27,284
Neoephemeridae
Neoephemera sp. 27
N. bicolor McDunnough 28
TV. purpurea (Traver) 28
TV. youngi Berner 284
Neoephemeropsis sp. 3 1
Oligoneuriidae
Elassoneuria insulicola Demoulin 302 Uw
Homoeoneuria dolani Edmunds, Berner and Traver 93
Lachlania sp. 93 Us
L. dencyanna Koss 189
L. powelli Edmunds 86
L. saskatchewanensis Ide 208 Us
Oligoneuriella baskale Soldan and Landa 318 Us
O. mikulskii Sowa 321 Us
O. mongolica Soldan and Landa 31 8 Us
O. pallida Hagen 321 Us
O. rhenana (Imhoff) 196,292,293,321 ,323 Us
Quaest. Ent., 1982, 18 (1-4)
88
Clifford
0. zanga Soldan and Landa 318 Us
Polymitarcidae
Ephoron album (Say) 55,93,94,792,205,243,289 Us
E. leukon Williamson 169 Uw
E. virgo (Oliver) 1 96,2 18,527 no consensus
Povilla adusta Navas 30,71,72,146,147,291,347 MNP
Tortopus incertus (Traver) 27,28,243,575 2Y
Potamanthidae
Potamanthodes sp. 31
Potamanthus sp. 222
P. distinctus Traver 243,289
P. luteus (L.) 196 ,218,276,527 Uw
P. myops (Walsh) 75,243,289 2Y
Prosopistomatidae
Prosopistoma africanum Gillies 130
P. oronti Alouf 9 Uw
Siphlonuridae
Ameletus sp. 51,255 Uw
A. celeroides McDunnough 26 Uw
A. connectina McDunnough 191
A. costalis Matsumura 725,352 U?
A. inopinatus Eaton 70,47,45,44,48,97,720,121,143,180,184,795,202,257,527,554,355
Uw
A. ludens Needham 59 ,336 Uw
A. montanus Imanishi 725,218 U?
A. oregonensis Eaton 32
A. similior McDunnough 93
A. sparsatus McDunnough 757 Uw
A. validus McDunnough 93
A. vancouverensis McDunnough 191
A. velox Dodds 26 Uw
Aneletris eximia Edmunds 27 7 Us
Coloburiscus humeralis (Walker) 369 Uw
Edmundsius agilis Day 28,93
Isonychia sp. 31,284
/. bicolor (Walker) 57,58,217,254,289,555,336 MBws
/. harperi Traver 217
/. pictipes Traver 27 MNP
1. sadleri Traver 217,299
I. sicca (Walsh) 208 Us
I. thalia Traver 51
Nesameletus sp. 367
Life cycles of mayflies (Ephemeroptera)
89
Oniscigaster wakefieldi McLachlan 247,248 Uw
Parameletus sp. 355
P. columbiae McDunnough 87,93 Us
P. minor Bengtsson 218
Siphlonisca aerodromia Needham 28, 93 Us
Siphlornurus aestivalis (Eaton) 5,35,43,44,153,178,796,197,355 Us-MBss
S. alternatus (Say) (not including S. linnaeanus) 16,32,60,1 37,140,217 Us
S. armatus (Eaton) 1 3 1 , 1 43,7 96, 1 97 Us
S. lacustris (Eaton) 36,47,43,44,45,83,131,143, 164,196,197,231 ,321 ,353,355
S. linnaeanus (Eaton) 128,143,153,178,796,797,355 Us
S. marshalli Traver 289
S. mirus Eaton 5 1
S. occidentalis Eaton 191 ,209 Uw
S. quebecensis (Provancher) 217
S. rapidus McDunnough 217
S. zetterstedti Bengtsson 218
Tricorythidae
Tricorythodes sp. 752 Us
T. albilineatus Berner 21,284 MNP
T. allectus Needham 57,226,254
T. atratus McDunnough 733,734,289,336 Us/MBss
T. explicatus (Eaton) 217
T.f ictus Traver 238
T. minutus Traver 16,93,274,27 5,306 no consensus
T. stygiatus McDunnough 217
Tricorythus maculatus Kimmins 347 MNP
T. tinctus Kimmins 347 MNP
see text
Quaest. Ent., 1982, 18 (1-4)
EVOLUTIONARY ASPECTS OF GEOGRAPHICAL VARIATION IN COLOR AND OF
PREY IN THE BEEWOLF SPECIES PHILANTHUS ALBOPILOSUS CRESSON
Gerald J. Hilchie
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Quaestiones Entomologicae
18:91-126 1982
ABSTRACT
Prey selection and aspects of life history were studied in a population of Philanthus
albopilosus Cresson, near Empress Alberta, and compared with published data reported about
other populations. A disproportionately large number of sphecid wasps was used as prey in
comparison with prey used by more southern populations. Females did not hunt at flowers,
but appeared to hunt suitable apocritans found around the Empress dune, or captured male
apocritans which pursued them as potential mates. False burrows appeared to function as
visual aid in orientation to the nest. Relative absence of other species of Philanthus implies
specialization of wasps of P. albopilosus for life on sand dunes.
Differences in prey selected, nest structure and colour of adults suggest geographic
isolation and differentiation during Pleistocene glaciations. The Nebraska Sand Hill region is
proposed as a northern refugium where differentiation of the dark race, P. albopilosus,
occurred during the Wisconsinian glacial stage. A southern refugium in the American
Southwest is proposed for the ancestral stock of the pale race, P. albopilosus manuelito a
newly described subspecies (type locality Monahans, Ward County, Texas).
RESUME
Cette etude porte sur la selection des proies et sur divers autres aspects de I'histoire naturelle d’une population de
Philanthus albopilosus Cresson vivant pres d’Empress en Alberta. Les resultats sont compares a d’autres donnees deja
publiees se rapportant a d’autres populations de cette espece. Le regime de proies de la population etudiee comprenait
une proportion anormalement elevee de guepes sphecides compare au regime de populations plus meidionales. Les
femelles ne chassaient pas sur les fleurs, mais semblaient plutot capturer les hymenopteres apocrites convenables trouves
dans les parages de la dune d’Empress. Elies capturaient aussi les males apocrites qui les poursuivaient en vue d’un
accouplement. Les faux terriers semblaient servir de reperes visuels dans V orientation vers le nid. L’ absence relative
d’especes de Philanthus autres que P. albopilosus implique que les guepes appartenant a cette espece sont specialisees
pour vivre sur les dunes de sable. Des differences dans les proies selectionnees, la structure des nids et la coloration des
adultes suggerent y a eu isolation geographique et differenciation pendant les glaciations du Pleistocene. La region de
Nebraska Sand Hill pourrait avoir servi de refuge septentrional ou la differenciation de la forme foncee, P. albopilosus
albopilosus, aurait eu lieu au cours du stade glaciaire wisconsinien. Le sud-ouest americain pourrait avoir servi de refuge
meridional aux populations ancestrales de la race pale, P. albopilosus manuelito, une sous-espece nouvellement derite
(localite du type: Monahans, comte de Ward, Texas).
TABLE OF CONTENTS
Introduction 92
Materials and Methods 95
Results 98
92
Hilchie
Prey Selection 103
Discussion 105
Geographical Distribution and Variation 111
Evolutionary Considerations 120
Evolution of Colour Pattern and Behavioural Features of P. albopilosus 122
Acknowledgements 123
References 123
INTRODUCTION
Wasps of the genus Philanthus, have aroused new interest of researchers. Classical studies
by Tinbergen (1932, 1935) using P. triangulum focused on orientation by females to their
concealed nests. In North America little work was done on the genus until about the 1950’s.
Early workers recorded habitats and prey of various Philanthus species (Rau and Rau 1918,
Peckham and Peckham 1905). Reinhard (1924) reasoned that because of the observed
specificity of females of P. gibbosus (Fabricius) to a particular group of bees (Halictidae) and
the occurrence of another family Andrenidae represented in the prey records, that these
families should be combined. However most members of the genus Philanthus are not specific
to a few species or families of bees. Other workers (Alcock 1974, Armitage 1965 and Cazier
and Mortenson 1965, Evans 1966b, 1970, 1975, and others) report that bees are preferred prey,
but females of some species ( P . pulcher Dalla Torre, P. crabroniformis Smith, P. zebratus
nitens (Banks), P. pacificus Cresson, and others) take many sphecids. Armitage (1965)
explored aspects of prey quality, Alcock (1974) experimented with visual clues used in hunting
and Evans (1970) investigated some areas of competition between species for prey.
Most studies of beewolves have been ethological. Little information is available about
historical factors which have shaped behaviour. Lorenz (1967) stated that behaviour “...must
never be regarded as a product of change or taken as a matter of course when behaviour is
found to be adapted to a corresponding point of the species environment”. In this study
behavioural and structural features are features used to document relationships of populations
against a background of Pleistocene and Holocene changes in environmental parameters, and to
recognize a new subspecies. Different populations have experienced different selective pressures
and these may be manifested in differences in life history and behaviour. Analysis of colour
variation allows interpretation of dispersal patterns and hypotheses about relations between
populations (Freitag 1965). This permits speculation about evolutionary events which may be
responsible for differentiation of various facets of life history and behaviour.
STUDY SITE
The study site was 1 1 km south of Empress, Alberta, east of the Alberta-Saskatchewan
boundary (50° 51’ 30” north, 109° 59’ 27” west; Figure 1). The dune field straddles the
provincial boundary and active dunes are on both sides (Whitaker and Christiansen, 1972). The
Empress dunes are more or less an eastern continuation of the Middle Sand Hills.
Quaest. Ent., 1982, 18 (1-4)
Philanthus albopilosus Cresson
93
Alberta {Saskatchewan
Figure 1. Map of study area. % marks the study dune. The inset shows the relative position of the study site in North
America
94
Hilchie
The study dune was the crest of a low bluff overlooking the South Saskatchewan River.
Sand from the eastern edge of the dune blows down the bluff into the gallery forest near the
river. The Empress dune field is on a layer of glacial till intermixed with lacustrine deposits. All
of these overlie the Judith River formation1 which is exposed along the river valley. The
undulating terrain of the dunes, variety of substrates and close proximity to the river combine
to create a varied physical habitat permitting many species of insects to occupy a small area.
Climate of the study site is related to continental position, absence of modifying bodies of
water, a mid-northern latitudinal location, and rainshadow influence of the Rocky Mountains
(Coupland 1950). Average annual precipitation is low, 22.55 cm at the town of Empress (1975
Climate of Alberta Report). Evaporation rate from free standing water in the Great Sand Hills
of Saskatchewan to the east is approximately 75 cm during the frost free season (Hulett et al.
1966). The Empress area is the most arid region in Alberta and may be drier than any area in
Saskatchewan. Aridity of the region probably permits continued movement of the sand dune by
limiting vegetation cover.
The Empress dunes appear to have similar plant associations to those recorded by Hulett et
al. (1966) for the Great Sand Hills (about 60 km due east). The dunes are inhabited by several
species of xeric plants ( Oryzopis hymenoides Ricker, Psorealea lanceolata Pursh and
Sporobolus cryptandrus A. Gray) which are commonly found further to the south or on sand
dunes in the Great Basin (Chadwick and Dalke 1965).
Philanthus albopilosus shares the Empress area with five other species of Philanthus.
Philanthus psyche Dunning nests on the sand dunes near nesting areas of P. albopilosus ,
Philanthus solivagus Say, P. ventilabris Fabricius, P. gibbosus (Fabricius), and P. gloriosus
Cresson were found near, but did not nest on the open sand of the sand dune.
Fauna of the Prairie Provinces is of comparatively recent origin. The majority of organisms
currently found within the provincial boundaries entered from refugia in the south following
melting of Wisconsinan Ice. Major glacial advances during the Wisconsinan eliminated all
previous living assemblages. Summaries of events of Wisconsinan age are presented by Flint
(1971), Wright and Frey (1965), and Mitchell (1979).
The Wisconsinan Glacial Stage is postulated to have begun about 55,000 years B.P. and
achieved the maximum southward expansion of the ice sheet at about 17,000 years before
present. It is generally accepted that, with the southward movement of the ice, the faunal zones
were either overrun (Love 1959) in part or were displaced (Wright 1969) to the south. Extent
of displacement is still a matter of much speculation. Martin (1958) argued for a wide band of
tundra and taiga. Ross (1970) shows a boreal type forest abutting with the ice front, but
Frenzel (1973) portrays the tundra merging with the steppe on the Great Plains which splits
the boreal forest into eastern and western components. Many groups of animals dependent on
plant associations may have had their populations split into two main groups (Freitag 1965,
Mengel 1970).
Much of the insect fauna at the study site appears to be dependent on a sandy substrate.
Post-glacial colonization and subsequent development of the fauna is dependent on past events
and conditions.
'Bearpaw formation is present above the Judith River formation and beneath the glacial till a
few km to the east or west. The Bearpaw formation was eroded away by a periglacial river at
the site of the Empress dunes.
Philanthus albopilosus Cresson
95
At the zenith of the Wisconsin stage, a large ice sheet moved southward from the Hudson
Bay region. Soil and rocks were crushed, then ground and transported, eradicating evidence of
preglacial floras and faunas.
During melting of the Wisconsinan ice sheet, a large mass of ice impeded drainage of water
to the north. Various glacial lakes developed. Some were very large, such as glacial Lake
Agassiz (Mayer-Oakes 1967). However, most were smaller transient lakes, which formed in
ice-impeded river drainage basins, giving rise to extensive areas of lacustrine deposits
(Edmunds 1962, Klassen 1972) comprised of clay, silt and sand deposits. Wind, reworking
sandy lake bottoms and deltaic deposits, gave rise to recent dune sands (Mitchell et al. 1974).
Requirements for development of sand dunes are: (1) a source of sand and (2) a wind to
move the sand (Flint 1971). Other factors may modify availability of sand such as geologic
history of the area and climate. Geologic events control presence or absence of sand, whereas
climate controls transport of sand. Xeric environments tend to develop sand dunes more readily
than mesic areas. Sand dunes are in regions where sand is available for transport (e.g. active
dunes occur at Brule Lake, Alberta in the boreal forest, Dowding 1929).
The period of dune formation probably occurred in two major phases. The first began
shortly after recession of the ice, before mineral soils could be stabilized by encroaching pioneer
vegetation (Ashwell 1966). This phase may have lasted for a short period. Subsequently, boreal
type forest may have developed and moved north across the plains (Bryson and Wilson 1967),
Wright 1969). The climate ameliorated, allowing expansion of prairie northward, beyond its
present limits, reaching a maximum about 7,000 years before present. During the
Hypsithermal renewed dune building would have occurred. Increased aridity reduced
vegetation cover allowing increased aeolian erosian. During this warm dry period many of the
xeric-tolerant and sand-adapted insects would have migrated north into new unoccupied
habitats. Along with cooling since the Hyspithermal, Boreal forest has reinvaded much of the
northern prairie regions and continues to slowly encroach (Love 1959).
MATERIALS AND METHODS
Study material included 39 adults of P. albopilosus , collected at the study site, 46 specimens
collected in the south-western United States, and 195 borrowed specimens. The latter are from
the following collections: CAS
CAS California Academy of Sciences, San Franscisco, Calif. 94118
CNC Canadian National Collection, Ottawa, Ontario. K1A 0C6
CSU Colorado State University, Fort Collins, Colorado. 80521
CUM Cornell University, Ithaca, N.Y. 14853
GH G. Hilchie, Edmonton, Alberta. T6E 4B5
MCZ Museum of Comparative Zoology, Cambridge, Mass. 02138
SEM Snow Entomological Museum, Lawrence, Kan. 66045
UAS University of Alberta, Strickland Museum, Edmonton, Alta. T6G 2E3
UCB University of California, Berkeley, California. 94720
UCD University of California, Davis, California. 95616
USNM United States National Museum, Washington, D.C. 20560
Quaest. Ent., 1982, 18 (1-4)
96
Hilchie
Behavioural Study
During the summer of 1977, nesting behaviour of P. albopilosus was studied in detail. More
cursory data from previous years have been incorporated with data from the 1977 season.
Early in the season, nests of P. albopilosus were left undisturbed. Nests were located by
watching a potential nesting area for prey-laden female wasps and locating false burrows.
Nesting areas often had males patrolling and landing on the sand. Each nest was marked with a
stake.
In conjunction with marking nests, individual female wasps were marked with small dabs of
Testors model paint on the scutum. Five colours were used in two positions (right and left side).
Males of P. albopilosus were not studied at the Empress site.
Nests were initially observed while recording prey brought back by females. Toward the end
of July, in an effort to determine the prey species more accurately, many staked nests were
sacrificed in the evening when the female was inside with a day’s capture of prey. Prey and
beewolf were collected and preserved in 75% alcohol solution. Voucher specimens are in the
Strickland Museum (University of Alberta).
Potential prey available at the site was sampled by sweeping vegetation around the dune in
potential hunting areas. Bees and wasps were collected from blossoms of Psoralea and
Solidago. Other dune-inhabiting bees and wasps were collected as they were encountered.
Notes about relative abundance were made for some of the common species.
In 1978, several additional populations of P. albopilosus were observed at Roggen, Colorado
(Evans’ study site) and Little Sahara Recreation Area, Utah. The Roggen dunes were very
similar to the Empress dunes in appearance.
Taxonomic Study
Trends were examined through use of a Compound Character Index (Brown and Wilson
1956), because of lack of a single consistent diagnostic character or group of characters.
Colour is important in species discrimination, and it is sometimes necessary to rely heavily
upon colour (Evans and Matthews 1973). Colour is subject to variation in hue and extent in
many species. Trends may be correlated to environmental gradients, such as moisture ( Bembix
variablis, Evans and Matthews 1973). and thus may be ecophenotypic. When trends in colour
do not follow evident environmental gradients, with markedly contrasting (divergent)
populations occupying neighbouring areas and intermediate masculated forms in peripheral
zones, differences may be taken as a measure of genetic differentiation and thus of taxonomic
significance.
Character states used are as follows.
Females. - Eleven characters were studied (Table 1.) whose states were constant in a given
locality, but varied geographically. Many additional characters such as pubesence length and
density were examined but they varied within localities. Variation in density of pile may result
from age of the wasp and variable abrasion by sand.
1. Frons: An estimate of percentage of area marked with yellow was made. The area
considered was delimited at the apex of the head by a line on the microscope ocular grid
positioned at the base of the median ocelli and extending to contact both compound eyes.
Estimates of coloured area were made to the nearest 5%. Maximum ranking was given a value
of 99%. The border between the clypeus and frons was dark and assumed to be 1%.
2. Clypeus: The clypeus was scored in a similar manner with a maximum score of 99% The
area was delimited by the dorsal suture with the frons and the ventral suture with the labrum.
Compound character index values assigned to selected character states in female wasps of Philanthus albopilosus.
Philanthus albopilosus Cresson
Quaest. Ent., 1982, 18 (1-4)
^Denotes percentage area of light colored maculations. Diagrams denote relative size and shape of maculation,
anterior is towards top of column.
98
Hilchie
3. Vertex of Head: Colour pattern was given index values ranging from 0 for no maculations
to 9 for a solid bar.
4. Scutellum: Character states ranged from entirely black through two spots to a single large
median spot with index values of 0 to 9 respectively.
5. Mesonotum: Three character states were recognized; black, two lateral spots and a single
tranverse band with index values of 0, 5 and 9 respectively.
6. Pronotal Collar: Five character states were recognized: black, small lateral spots, large
oblong lateral spots, broken band and a continuous transverse band which were assigned index
values of 0, 2, 5, 7 and 9 respectively.
7. Scutum: An estimate of percentage colour was made. Range was from 0% to 50%.
Estimates were to the nearest 5%.
8. Propodeum: An estimate of percentage colour was taken, which ranged from 0% to 80%.
Maculations on the dorsal and lateral aspects were used in the estimate. Posterior exposure of
the propodeum was excluded from estimates because it was obscured by the abdomen in many
specimens.
9. First Abdominal Tergum: An estimate of the area of light colour was made from the
dorsal aspect. Values varied from 0% to 95%. Estimates were made to nearest 5%.
10. Abdominal Tergum Two: Colour was estimated as for the first abdominal tergum with
values ranging from 20% to 99%.
11. Abdominal Sternal Spots: These are on the lateral portion of the sternal sclerites, one
spot per sclerite per side. Total number of spots on the right side were recorded and assigned an
index value.
Low values were assigned to black or minimum colour and high values to the maximum
amount of yellow. All character states were given equal weighting.
Males. - Males varied less than females. Five characters were studied (Table 2.).
1. Pronotal Collar: Maculations varied from solid black to a continuous bar of yellow.
2. Scutellum: Only two character states were recognized; presence or absence of two yellow
spots.
3. Colour of Abdomen: Most frequent colour for males is bright lemon yellow, tinted in some
with orange, in others the colour is a rich orange to red. The red or darker colour is assigned an
index value of 0 and yellow the index value of 4.
4. Pronotal Lobe: Presence or absence of colour was noted and given a value of 0 (no colour)
or 4 (colour).
5. Fourth Abdominal Tergum Maculation varied from a single median spot (index value 0),
a series of spots, median and lateral (index value of 2) and a band spanning the width of the
tergite (index value 4).
A score of 0 typified the extreme dark male and a score of 20 the extreme light male.
No taxonomically useful differences were found in the male genitalia. Bohart and Grissell
(1975) and Strandtmann (1946) also report that variability in male genitalia is of little or no
taxomic importance in Philanthus.
RESULTS
Life History
At Empress, P. albopilosus is closely associated with a dune environment. Females typically
nested on margins of broad, flat bare areas of the sand dune. Nests were in a zone of lightly
Philanthus albopilosus Cresson
99
Table 2
Compound character index values assigned to selected
character states in male wasps of
Philanthus albopilosus Cresson.
Diagrams denote relative shape and size of maculation.
drifting sand, away from vegetation.
A nest is of elaborate design (Evans, 1975). The entrance of a new nest is concealed by a
sand closure and a number of false burrows (0 to 6) may be present. They are constructed
immediately following completion of a nest at the time of initial closure, but are not
maintained, and some fill with loose sand in a few hours. From the nest entrance a horizontal or
nearly horizontal vestibule extends for about 25 cm. The vestibule terminates at the beginning
of a vertical gallery. Vertical galleries extend 25 to 30 cm in depth. Provisioned cells were not
located. Prey is stored in the vestibule until a cell is provisioned.
Females occupied their nest for a relatively short time, usually one or two days. Nests were
provisioned from late June through July into early August. Life span for adults of P.
albopilosus is as yet undocumented but females of P. gibbosus ( Reinhard 1924) and P.
crabroniformis (Alcock 1974) live and provision their nests for about three weeks.
Males in the Empress area typically perched on the sand in alert positions. They often flew
after passing insects and frequently grappled with females (prey-burdened) about to enter their
nests. In 1980 copulating wasps were observed resting on the dune surface.
Comparisons with Other Populations
At Roggen, Colorado, male P. albopilosus behaved as reported by Evans and as observed on
the Empress dunes. No females were sighted or nests located. In contrast, 19 males and 5
females were captured at Little Sahara, Utah. No males were seen patrolling and no nests were
found in “suitable” areas. All beewolves were captured foraging on flowers along the' dune
edges in the vegetation belt.
Quaest. Ent., 1982, 18 (1-4)
8
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AREA 1
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MAY
JUNE
JULY
AUGUST
AREA 12
-i — | — i — i — i — i — i — i — rp — i — i — i — i — i — r
SEPTEMBER 1 OCTOBER
Temporal distribution of adult Philanthus albopilosus Areas are those illustrated in Figure 3. Dashed line above
for Area 12 (Empress) indicates collecting dates at the study site.
Philanthus albopilosus Cresson
101
Figure 3. Distribution of Philanthus albopilosus . Numbers shown are used in the text to refer to the localities enclosed by
the dotted line. The dark race P. albopilosus albopilosus occurs northeast of the dashed line, and the light race P. a.
manuelito to the south west. Symbols used: ♦ locality record (specimens seen); A state record (specimens seen); A state
record (published).
Quaest. Ent., 1982, 18 (1-4)
102
Hilchie
Evans (1975) reported that cells at the terminal end of burrow galleries averaged about 2.5
times deeper in Colorado populations than in those studied near Albany, New York. Vertical
gallery depths at Empress compare favourably with those at Albany, New York.
If depth of the burrow reflects past evolutionary interactions and with diminished response
to local conditions, presence of shallow burrows in the northern populations may reflect
adaptation to past climatic stress. Cells closer to the surface ought to experience warming
earlier in the season. This permits a longer period of development, which would allow
colonization of areas with a shorter growing season. The opposite may be true in southern areas:
intense heating of the sand by the sun may be lethal to cells near the surface (Chapman et al.
1926) which would select for deep nests.
Data about periods of adult activity were obtained from locality labels; dates were
consolidated into four day units and illustrated in Figure 2 using the area groups illustrated in
Figure 3. Northern populations (Areas 7 to 12) appeared to have one main generation active
during July, and in some (Areas 7, 8, and 9) there appears to be a smaller second generation in
early September. Among southern populations, collection dates range from May 27 to October
17, indicating the possibility of several generations per year.
Data obtained at Empress (Area 12), provide a more unbiased indication of activity of
adults. Collecting dates and number of beewolves collected are both presented so that negative
evidence is an indicator of when beewolves were not present.
Females at the Empress study site appear to occupy nests for a short duration comparable
with Colorado and New York populations (Evans 1975). Evans reports that of 26 marked nests
in Colorado only a few were active after a few days. In the New York Sample of marked nests
(n = 5), two were active three days later (Table 3).
The Empress population is not distinctly different from the New York population. However
Colorado females dig deeper galleries than observed in the two more northerly populations.
Table 3. Duration of nest occupancy on the Empress dunes in comparisons with Evans’ (1975)
study sites.
Numbers refer to the nests observed. Day zero refers to when the nests were first
observed.
Philanthus albopilosus Cresson
103
PREY SELECTION
Prey specificity in Philanthus is a result of evolutionary interactions and environmental
factors expressed as a mosaic of different wasp populations using different complexes of prey
species (Evans 1966c). Precise entrainment to a particular group may be detrimental because
prey species are subject to fluctuations in relative abundance; a plastic response to a complex of
prey groups may be a more successful evolutionary strategy.
Females of philanthine genera use various apoids and, to a lesser degree, other wasps.
Females of most species of Philanthus show a preference for halictid bees, with a marked
tendency of some species to use sphecids as prey (Armitage 1965, Evans 1970). Females of
Aphilanthops prey exclusively on ants of the genus Formica (Bohart and Grissell 1975).
Remarkably, females of most species of Cerceris and those of all species of Eucerceris prey on
beetles. (Bohart and and Menke 1976). This is exceptional among philanthines, but illustrates
the sorts of differences among groups included in a single subfamily. These last two genera are
closely related to one another, but are quite distantly related to Philanthus (Menke 1976).
The Empress Dunes Populations
Females of P. albopilosus captured a varied group of prey including members of three
families of wasps and three families of bees. Twenty six male and four female wasps used as
prey were members of nine species. Aphilanthops frigidus (F. Smith) males were the most
common wasps used as prey (17 individuals). The 48 male and two female bees captured as
prey were members of seven species. Most of the prey records were divided between males of
two species, Halictus rubicundus (n = 23) and Colletes sp. (n=19). Overall, 37.5% of the prey
were wasps and 92.5% were male apocritans. Prey ranged in length from 5 to 1 1 mm with a few
wasps measuring 14mm and 19.5 mm. The most commonly used prey ranged from 8 to 1 1 mm.
General collecting of wasps was done in the study area to establish a basis for comparing
what prey was taken with what was available. Table 7 contains data about apocritans whose
adults seem to be within the size range preferred by females of P. albopilosus.
Table 4. Provisioning rates for selected females of Philanthus albopilosus .
Time intervals are from when the female left the nest to when she returned with prey.
Quaest. Ent., 1982, 18 (1-4)
104
Hilchie
Table 5. Number of prey per nest.
Rates of provisioning were variable (Table 4). One wasp (#004) returned in five minutes
with prey; another wasp completed the task in 55 minutes. Females captured about five prey for
each cell provisioned (Table 5).
On August 4 I observed a female P. albopilosus attack and capture prey. A beewolf cruising
about 50 to 60 cm above the sand near a colony of Microbembex monodonta (Say) wasps,
dived to the ground, attacked and captured one of the Microbembex. The latter individual was
either flying near or resting on the sand. I did not see it until the beewolf made contact.
Females of Microbembex monodonta nested along the edges of the sand dune.
Throughout the 1977 field season, neither sex of P. albopilosus was observed at flowers. If
female P. albopilosus did hunt at flowers, I would expect that number of female bees taken as
prey would increase or exceed number of males captured. Females of several species of bees of
the correct approximate size to be used as prey were quite common on flowers on or near the
dune.
Comparisons with Other Populations of P. albopilosus
Evans (1975) reported a prey capture pattern similar to the Empress population of P.
albopilosus from Albany, New York. Females in the Albany population captured wasps of two
families (Sphecidae and Eumenidae) and captured bees of three families (Colletidae,
Halictidae and Andrenidae). Prey records of wasps comprised 17.3% of the total catch, and of
male Apocrita, 79.1%
Prey data for the Colorado population of P. albopilosus are different. Number of species
and families of Hymenoptera decline, for 73 prey records include only four species of two
families of bees (Halictidae and Colletidae). A single nest of P. albopilosus in Tuba City,
Arizona contained one halictid and four anthrophorid bees (Evans and Lin 1959). Because few
populations were examined, difference may be due in part to sampling error.
Many authors (Armitage 1965, Evans 1966b, 1970, Reinhard 1924, Tinbergen 1935, and
others) have reported female Philanthus hunting at flowers. Pinned specimens of P.
albopilosus from several localities bear flower capture data. These are as follows: Colorado, on
Tamarix; Arizona, on Poliomintha and on Parvella. There are no data for capture at flowers
for any northern localities.
Philanthus albopilosus Cresson
105
DISCUSSION
Selection of prey. - Data presented show that prey selected by females of P. albopilosus at
Empress is typical for philanthines. A mixture of bees and wasps is taken, and this is consistent
with Evans’ (1975) observations of a colony of P. albopilosus in New York. However,
information of this sort is only the first step toward understanding the general phenomenon of
prey selection for this species.
In an organism specializing on a particular group of prey species, many factors influence
choice. Physical constraints of the predatory apparatus and behavioural restraints control
which type of prey are suitable. Behavioural restraints may also regulate where the organism
hunts, type of prey, and quality of prey acceptable (Armitage 1965, Alcock 1974, Tinbergen
1935). Availability of suitable prey and its relative abundance will affect what is caught by the
predator.
Certain parameters may be useful in predicting whether or not a particular species of bee or
wasp will be exploited. Evans (1970) used length of prey, and despite some disadvantages I use
also body length to estimate prey acceptability. Adult apocritans whose minimum body length
is 5 mm and maximum length is 1 1 mm for stout forms and 20 mm for slender forms, are
regarded as potential prey.
Prey selected by female P. albopilosus on the Empress dunes consists of mostly bees and
sphecid wasps (Table 6). Table 6 lists apocritan species found in the study area whose adults
are of a suitable size to serve as prey for females of P. albopilosus. As an indicator of where
females of P. albopilosus hunt, notes are included about locations and relative abundance of
potential prey.
Proportions of bees and wasps differed between prey used (Table 6) and potential prey
(Table 7) (X2=3.84, p>0.05). Females specialized on a few common species occurring near
dune margins. Surprisingly, dune dwellers are underrepresented among prey used. There is also
heavy bias toward males of the dominant species of prey. Males of many species of bees and
wasps frequent flowers or positions near flowers while seeking potential mates.
Hunting females flew swiftly from the nesting area into vegetation surrounding the dune,
where they were quickly lost from sight. In view of the prey captured, I think that two different
methods of hunting are employed. The first is to attack and capture any apocritan seen, of
appropriate size. This would produce a wide spectrum of prey and would not be biased toward a
particular species or sex. A number of apocritans not commonly encountered in large number
would be used. A second method of prey capture may be to attack and capture male bees or
wasps that trail the female beewolf, as a potential mate. Male bees and wasps which are much
larger or smaller than females of P. albopilosus may detect the error at greater distance; those
whose females are of similar size and colour would likely continue pursuit and may possibly
grapple with the beewolf. The three species of prey captured in the greatest number were males
of Aphilanthops frigidus (F. Smith) Colletes sp. and Halictus rubicundus Christ.. Females of
the three prey species approximate the size and appearance of female P. albopilosus. Neither
the A. frigidus wasps nor the Colletes sp. and H. rubicundus bees were very abundant around
the dunes.
Prey selected by female P. albopilosus (Empress dunes) is consistent with data from a
population examined by Evans (1975) at Albany, New York, but is inconsistent with data from
the population examined by him at Roggen, Colorado. This may suggest a genetic basis, which
is behaviourally manifested in either, 1) site of prey capture, or, 2) acceptability of potential
prey after contact has been made: or, 3) prey selection is the result of opportunism.
Quaest. Ent., 1982, 18 (1-4)
106
Hilchie
Table 6. Prey records for wasps of Philanthus albopilosus Empress study site.
Change in prey selected, may have occurred as a result of populations of P. albopilosus
becoming isolated in different refugia during the Wisconsin glaciation. In a northern refugium
(i.e. Nebraska Sand Hills, see below) changes in faunal composition coincident with changing
climate may have led to a shift in prey selection. Decreasing abundance and number of suitable
species of bees, may have selected populations of P. albopilosus that used increasing
proportions of wasps. Selection of sphecids over other groups of wasps may be in part, due to
prevalence of dune nesting species with similar environmental tolerances, relative abundance
around dunes and a suitable body size.
False burrows.. - A false burrow or accessory burrow is defined by Evans (1964b) as “any
burrow started from the soil surface in close proximity to the true burrow and made by the
same individual”. Evans suggests that these burrows serve to dupe nest parasites into
ovipositing in an innapropriate place or to divert parasites into exploring empty holes where
there are no hosts. Wasps which maintain or repair false burrows dig them early in the history
Table 7. Some taxa of Hymenoptera collected in the Empress area which are available as potential prey* to the beewolf Philanthus albopilosus.
Philanthus albopilosus Cresson
107
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Anthrophoridae
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Philanthus albopilosus Cresson
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Table 8. Occurrence of accessory burrows in the Sphecidae
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Philanthus albopilosus Cresson
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of the nest, usually during or after initial closure. The habit is thought to have evolved from
quarrying soil for a closure of the nest. Maintaining false burrows would simply entail use of
one or a few of the holes as a source of soil for later closures. Maintenance of false burrows may
provide continued protection from nest parasites. Females of P. albopilosus construct accessory
burrows after completion of the nest.
Associated with species which dig false burrows early, but do not maintain them, are various
behavioural features which may reduce the effect of nest parasitism(Table 8). Females of
Bembix provision their nests progressively. If parasites enter the nest the female may be able to
supply enough prey, permitting survival of both larvae and parasites. This tactic may remove
some of the selective advantage of maintaining false burrows.
Females of one species, B. texana Cresson, construct false burrows during final closure.
Debris is removed from the nest and parasites may be swept out during this process. False
burrows prepared at this time may offer protection from invasion by parasites, at least until
environmental effects obliterate all traces of the burrow.
Construction and failure to maintain false burrows in a mass-provisioning species such as P.
albopilosus is unexpected. The association of provisioning and nesting behaviours has changed,
I propose that false burrows made by P. albopilosus serve as visual markers aiding in initial
orientation to the nest in a habitat with an unstable substrate. A large premium would be
placed on learning the correct location of the nest without advertising its true entrance during
the first orientation flight. This would permit the female to take bearings on distant land marks
and visually relate the location of her nest (van Iersel 1964). After orientation, false burrows
would no longer have a function, and so they would not be maintained. A similar land mark
function has been attributed to the mound building activities of Bembix littoralis Turner.
Evans and Matthews (1975) reported B. littoralis females building a mound of soil at the
entrance of their burrow, which may serve as a species specific marker identifying the nest
location when other species of sand wasps were present.
GEOGRAPHICAL DISTRIBUTION AND VARIATION
Known Range and Locality Data,
Philanthus albopilosus is almost transcontinental in distribution with the most eastern
known locality at Albany, New York (Evans 1975) and the western most at Hatton,
Washington. The most northern locality is 175 km north of Empress at Rutland,
Saskatchewan. It is doubtful that the range extends much further north. The southern known
extreme is on Padre Island, Texas, but the range probably extends into central Mexico.
Localities of material studied are listed below. Numbers following locality names indicate
the number of females examined followed by the number of males examined. Acronyms
represent collections from which the material was borrowed.
Material examined:
CANADA. Alberta: Empress (11 km south), 31, 8 (GH); Medicine Hat, 6, 14, (CNC, UAS); Orion, 0, 2, (UAS).
Manitoba: Aweme, 2, 8, (CNC, UCD); Harteny 0, 1, (SEM); Onah, 3, 0, (CNC). Saskatchewan: (see Empress, Alberta)
Elbow, 5, 6, (CNC, UCD); Rutland, 3, 8, (CNC, UCD).
MEXICO. Chihuahua: Samalyuca, 0, 1, (UCD).
UNITED STATES. Arizona: Coconino County: Tuba City, 1, 0, (CUM); Navajo County: Hotenvilla, 1,1, (UCD);
Indian Wells (16 miles south), 1, 0, (UCD); Judito Trading Post, 1, 2, (UCD); Joseph City, 0, 1, (UCB); Kayenta (19
miles southwest), 2, 0, (CAS); Colorado: Alamosa County: Great Sand Dunes National Monument, 0, 2, (CSU); Bent
County: Caddoa (9 km east), 3, 16, (GH); Hasty, 1, 1, (CSU); Prowers County; Carlton, 1, 3, (CSU); Lamar (10 miles
west), 0, 2, (CSU); Weld County: Roggen, 10, 1, (CSU,GH). Idaho (Strandtmann 1946). Iowa: Woodbury County:
Sergeant Bluff, 1, 1, (USNM); Sioux City, 0, 1, (USNM). Illinois: Cook County: Chicago, 2, 5, (MCZ). Kansas: Kearny
112
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County: McKinney Lake, 0, 1, (MCZ); Stafford County: Salt Flats, 2, 4, (CUM, CNC, MCZ). Minnesota: Polk County:
Muskoda, 0, 1, (CUM). Montana: 0, 2. Nebraska: Cumming County: West Point, 1 , 2, (USNM). New Mexico: Dona Ana
County: Las Cruces, 0, 2, (CAS, SEM); Otero County: White Sands National Monument, 0, 5, (MCZ, UCD); San Juan
County: Shiprock (near), 0, 1, (USNM); Socorro County: La Joya Wild Life Preserve, 8, 14, (CSU, MCZ,GH). New
York: Albany County: Albany (Evans 1975); Colonie 3, 0, (CSU). North Dakota: Billings County: Medora, 0, 1, (MCZ);
Ransom County: McLeod, 1, 1, (UCB), Sheldon, 2, 1, (CUM, MCZ); Richland County: Walcott (11 miles west), 1, 0,
(UCD); Williams County: Williston, 1, 2, (MCZ). Ohio (Strandtmann 1946). South Dakota: Fall River County:
Hotsprings, 1, 0, (MCZ). Texas: El Paso County: Fabens, 0, 2, (CAS, SEM); Hartley County: Romero, 2, 4, (SEM,
UCD); Nueces County: Padre Island, 0, 1, (UCD); Ward County: Monahans State Park, 10, 22, (CSU, MCZ). Utah:
Juab County: Eureka, 0, 1, (UCB); Little Sahara Recreation Area, 5, 19, (GH); Utah County: Utah Lake, 0, 1, (CUM):
Millard County: Oak City (18 km north), 0, 1, (GH). Washington: Adams County: Hatton, 0, 1, (USNM).
In total 234 specimens were examined, 100 females and 134 males. Positions of localities are
mapped in Figure 3. Dotted lines define groups of populations which were united and treated as
a single population to increase sample sizes for statistical purposes. Populations were grouped
naturally using existing geographical barriers as boundaries.
Most localities are in semi-arid to arid regions of the Great Plains with clusters in desert
regions. A range extension appears to follow the arid interior into the Great Basin. Three
eastern localities - Chicago, Illinois: Ohio and Albany, New York - are disjunctive. Beewolves
are not found in regions where annual precipitation exceeds 91.5 cm (36 inches). Moisture may
affect presence or absence of suitable habitats rather than having a direct influence on the
species. Activities by man permit colonization of disturbed sites such as sand and gravel pits
and road cuts.
Geographical Variation in Colour Pattern
Interpretation of species limits for many Philanthus wasps is based on maculation and
punctuation patterns in adults. Several closely related species of Philanthus were once thought
to be subspecies of P. politus. Recognition was in part based on coloration of tibiae, femora,
and punctuation and shape of metanotal lamina. In P. zebratus nitens (Banks) several forms
once recognized as distinct species have been united and given subspecies status. Diagnostic
characters were maculations and punctures. In this paper specimens of P. albopilosus are
placed in recognizable groups based on colour patterns.
Results of analysis with a compound character index are shown in Figure 4 for females and
in Figure 5 for males. To give some perspective to the index values used, Figure 6 a, b and d
illustrate some aspects of a dark northern female. The total compound character index value for
the specimen is zero. Figure 7 a, b and d illustrate a southern female which has a total
compound character index value of 93. Figure 8 illustrates the typical frontal maculation of the
head and an abdomen of a northern male. This male received a total index value of four.
Figure 4 illustrates variation in maculation patterns in females. In the north (Areas 7, 8, 9,
10, 11 and 12) index values are concentrated toward the low range, indicating populations
comprised of dark individuals. Most of the specimens received index values below 30. A few
maculate specimens (Figure 7c) were present but index values did not exceed 70 units.
Females from more southern localities (Areas 4,5 and 6) had extreme expansion of the
yellow markings and few specimens were assigned index values less than 70. No populations are
known to have intermediate index values.
Females from mountain regions of the southern United States (Areas 1, 2 and 3) are
intermediate in coloration, with index values between 30 and 70. These groups are the most
geographically distant populations from those in the northern areas.
Number 20
Philanthus albopilosus Cresson
113
Figure 4. Compound character index values for females of Philanthus albopilosus. Area numbers refer to those illustrated in Figure 3.
Wasps
114
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Quaest. Ent., 1982, 18 (1-4)
Figure 5. Compound character index values for males of Philanthus albopilosus. Area numbers refer to those illustrated in Figure 3.
Philanthus albopilosus Cresson
115
Figure 6. Typical and maculated specimens of female Philanthus albopilosus (northern race). Figures 6a, 6b, and 6d,
head, thorax and abdomen of a dark specimen from Empress, Alberta. Figure 6c, head of maculate specimen of the dark
race at Medicine Hat, Alberta.
Males of P. albopilosus also exhibit some trends in colouration (Figure 5) but these are not
as distinctive. Individuals from northern populations (Areas 7 to 12) tended to be darker
(Figure 8). Males from southern and western areas (Areas 1 to 6) tended to be pale. There is
not a discrete break in index values but a trend to shift to darker forms in the northeast and
lighter forms in the southwest.
Index values for each area were averaged, female index values were plotted against male
index values (Figure 9). The points form loose clusters: northern groups separate together, in
contrast to southwestern populations and geographically intermediate populations. The points
illustrate a break in variation, with the largest break occurring between the geographically
intermediate group and the northern group. The phenotypically intermediate groups are the
extreme southwestern populations which may grade into the geographically intermediate
group.
Figure 7. Typical specimens of females of the light race Philanthus albopilosus manuelito . Figures 7a, 7b, and 7d head,
thorax and abdomen of specimen from Monahans, Texas. Figure 7c, head of specimen from Roggen, Colorado.
Quaest. Ent., 1982, 18 (1-4)
Philanthus albopilosus Cresson
117
Figure 8. Head and abdomen of male Philanthus albopilosus wasp from Empress, Alberta showing typical maculation
pattern for most males.
Taxonomic Interpretation
The beewolf species P. albopilosus includes two major groups of populations. Populations of
the dark form appear to form a cohesive geographic unit across the northern United States and
southern Canada (Areas 7, 8, 9, 10, 11 and 12). Slight differences among northern populations
may be due in part to “ founder effect ” (MacArthur and Wilson 1967), associated with
invasion of new areas by few individuals. Physiographic barriers are not evident. However sand
dune habitats are islands in a sea of grassland or forest, unsuitable for P. albopilosus.
Therefore, these latter habitats serve as barriers.
The southern light form is a slightly less coherent grouping. In Areas 2 and 3, an
intermediate colour form is present, which is partly isolated by mountain divide systems from
populations of the extreme yellow form found in Areas 4, 5 and 6.
It appears that there has been a break in gene flow between northern populations and
southern populations (Figure 9) for an extended period of time (probably since the Wisconsin
glaciation). At present, gene flow is interrupted between some populations in the southern race
by physiographic barriers. On the Great Plains gene flow appears restricted between northern
areas (7 and 8) and southern areas (5 and 6). No populations of intermediate coloration were
found between these areas, even though P. albopilosus is highly vagile and should have
dispersed, forming intermediate populations. This pattern suggests character displacement.
Character displacement occurs between ecologically similar sibling species, many pairs of
which exhibit narrowly parapatric distribution. Lack of a physiographic boundary separating
species indicates contact has recently occurred (Vuilleumier 1971). Taxonomists often
118 Hilchie
Figure 9. Compound character index value comparisons of means for colour variation in wasps of Philanthus albopilosus.
Numbers refer to areas illustrated in Figure 3.
experience difficulty deciding between displacement and hybridization (Brown and Wilson
1956). Races of P. albopilosus might be specifically distinct, and thus cryptic species.
The premise that the races have come into recent contact implies a barrier at some phase in
the species history. Populations in Area 4 may be relatively recently derived from yellow
populations further to the north which are in contact with the northern race. Origin from Areas
1, 2 or 3 is less likely because of mountain divide systems. A north-south movement on the
Quaest. Ent., 1982, 18 (1-4)
Philanthus albopilosus Cresson
119
Great Plains is more probable because of relative lack of geographic barriers.
The major isolating event separating the northern from the southern race was probably the
Wisconsinan glaciation. Isolation of the lineages may have ocurred late in the Sangamon
interglacial stage. This would place the dichotomy at about 60,000+ years B.P. Whitehead
(1972) suggested that speciation occurs at a rate of about one dichotomy per 3,000,000 years.
On the basis of this hypothesis, races of P. albopilosus may not have been isolated long enough,
to have evolved into separate species.
Regardless of what actually happened in the past, populations of P. albopilosus may be
grouped into definable races on the basis of phenotypic variation and these groups are restricted
to geographic regions. For a few populations additional evidence is supplied by variation in
aspects of life history and behaviour.
Subspecies of Philanthus albopilsus
In a species divided into various populations which are more or less isolated over time, when
do the variants warrant taxonomic recognition? Many authors have considered the subspecies
concept and each has a slightly different idea of what the limits should be; some are for
recognition, others are against. I prefer the argument given by Willis (1976)“ ... if subspecies
reflect to some degree the actual pattern of variation, as well as being convenient ‘handles’ for
reference, their value seems sufficient to justify their recognition”.
In philanthines, character differences in head shape, punctation, placement of ocellus and
most importantly maculation patterns, provide useful tools for species recognition. In members
of Philanthus albopilosus two major groups can be recognized. In certain regions, some
specimens feature character states not characteristic of the population mean. Most female
specimens (95%) and the majority of males can be assigned to one subspecies group or the other
consistantly on the basis of maculation patterns.
Philanthus albopilosus albopilosus Cresson Philanthus albopilosus , Proc, Ent. Soc. Phila., 5,
91, 1865 (5). P. simillimus Cresson, Proc, Ent. Soc. Phila., 5, 95, 1865 (2). For a complete
synonomy see Strandtmann 1946.
Female. - Length 10 to 13 mm. Black, marked with yellow as follows: clypeus 0 to 50%
yellow, frons 0 to 40% yellow (Fig. 7a, c), vertex at most five yellow spots, antennae scape black
above yellow below, pedicel black, flagellum black above infusciated below, thorax black to
black with maculations (Fig. 6b), coxae black or black with apical spot, femora black apices
yellow, tibiae yellow with black spot, tarsi pale to testaceous, sterna black may have lateral
spots, tergum 1 with two lateral spots, terga 2 and 3 with biemarginate band, tergum 4 with
broken band, tergum 5 with three spots, pygidium black.
Male. - Length 7 to 1 1 mm. (Fig. 8). Yellow, frons yellow, vertex black, scape black above,
pedicel black, flagellum with basal three yellow ventrally dorsally black infuscated toward tip,
thorax black, pubesence white, apices of femora yellow, tibiae yellow, tarsi infuscated
propodeum black, abdominal sterna black, terga 1,2, and 3 with yellow band, terga 4, 5, and 6
with elongate yellow spot; tergum 7 with or without elongate yellow spot.
Type locality. - Illinois.
Range. — CANADA: Alberta, Manitoba, Saskatchewan.
UNITED STATES: Illinois, Iowa, Montana, Nebraska, New York, North Dakota, Ohio, South Dakota.
120
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Philanthus albopilosus manuelito new subspecies
Female. - Length 10 to 13 mm. Yellow, marked with black, clypeus yellow, frons 10 to 20%
black near vertex (Fig. 7a, c), vertex with five yellow spots or solid yellow band, antennae as in
Philanthus albopilosus albopilosus , collar yellow, mesonotum with two longitudinal yellow
bands, scutum yellow, scutellum yellow, proprodeum large lateral yellow spots (Fig. 7b), apices
of coxae yellow, femora 30 to 50% yellow abdominal sterna with lateral spots, with or without
medial spots, terga yellow with black along apical and basal margins (Fig. 7d).
Male. - Length 8 to 1 1 mm. Similar to Philanthus albopilosus albopilosus , most
specimens with additional maculations on scutellum, collar and tubercle of thorax and with or
without additional spot on tergum 7.
Type material. - Holotype; UNITED STATES: Texas, Ward County, Manahans State
Park, 1-2 vi 1974, H. Evans and W. Rubink (female) [USNM]. Paratypes; 9 females, 22 males
with same data.
Range. — MEXICO: Chihuahua
UNITED STATES: Arizona, Colorado, Idaho, Kansas, New Mexico , Texas, Utah.
Derivation of specfic epithet. - Manuelito was a Navaho chief, who with the aid of other
chiefs, persuaded the United States goverment in 1868 to let the Navaho return to their home
lands. The bee wolf Philanthus albopilosus manuelito nests on reservation land.
EVOLUTIONARY CONSIDERATIONS
Evolution of the Geographic Pattern
The occurence of geographical races of P. albopilosus Cresson indicates fragmentation of
the ancestral range of the species, and hence, isolation of populations. At least two major
refugia seem to be indicated, one in northeastern Nebraska, and the other in southwestern
Arizona-New Mexico. Each region contained unique biotic and abiotic features. Divergence in
structure and behaviour are related to historical events which might have been causative.
Because of similarities between subspecies, they are of relatively recent origin. The
Sangamon interglacial stage between Illioinian and Wisconsinan glaciations may serve as a
reference point. Flint (1971) argued that conditions during the Sangamon probably ranged
from arid in the southwest to cool and moist in the east. Thus climate and plant biomes may
have been comparable with today’s. Dune habitats would have been present in many areas
where they are currently found. During the early part of the Sangamon interglacial stage,
populations of P. albopilosus Cresson were probably widespread (Figure 10a).
Toward the end of the Sangamon interglacial (about 70,000 B.P.) the climate began to cool
eventually giving rise to the Wisconsinan glaciation (from 55,000 to 10,000 B.P.). Cool wet
conditions induced reforestation of much of the Great Plains, and of the northern Great Basin
and Mohave Desert (Martin and Mehringer 1965). Increased moisture also caused reduction of
sand dunes as ground cover increased, with consequent widespread extinction of and disruption
of range of P. albopilosus Cresson. During this isolation, morphological and ecological
differentation occurred.
In large dune fields, however, porosity of the sand would locally induce aridity and thus
blowouts may have developed. Thus populations of P. albopilosus Cresson (s.l.) in the large
dune fields may have survived (Figure 10b).
Toward the end of the Wisconsinan glacial stage, climate ameliorated, lifezones move north
and up mountain slopes. Many stabilized dunes would have become active by loss of their plant
Quaest. Ent., 1982, 18 (1-4)
Philanthus albopilosus Cresson
121
Figure 10. Distribution patterns of Philanthus albopilosus through time, a) 70,000 B.P. hypothetical range of ancestral
stock, b) Ca. 17,000 B.P. geographic isolation of ancestral stock at the zenith of the Wisconsin, c) Ca. 7,000 B.P.
expansion of ranges from refugiom, d) recent distribution of (1) Philanthus albopilosus albopilosus (dark race) and P. a.
manuelito (pale race).
cover and being subject to wind erosion. Maximal rates of range extension may have occurred
during the Hypsithermal due to increased dune and blowout development.
In the southwestern United States, arid regions may have persisted during Wisconsinan
glaciation, permitting dune fields to remain active. Sonoran Desert floras were derived
relatively recently (Axelrod, 1979). The great diversity is due to a ‘vacuum’ effect, floristic
elements from surrounding zones which adapted to increasing arididty and temperature,
provide the bulk of taxa. A rise in rainfall of 250 to 300 mm would eliminate the Sonoran
Desert as a regional lowland. This would allow an oak-conifer association to cover much of the
lowland. Axelrod’s data suggest that taxa survived on the driest sites in bordering zones.
A portion of the american southwest is proposed as a refugium for the southern subspecies,
P. albopilosus manuelito . Very few sites suitable for a northern refugium are available east of
the Mississippi River (Thorp et al., 1952). Most of eastern North America was cloaked in a
continuous forest (Ross 1970, Martin 1958, Freitag 1973, and others). On the Great Plains,
several sand dune areas were probably active during portions of the Wisconsinan glaciation.
Smith (1965) demonstrated that there were several periods of dune building on the Nebraska
Sand Hills, based on past movement of stabilized dunes. This large dune area may have served
122
Hilchie
as a refugium for sand dwelling organisms which were able to adapt to a cooler climate. Willis
(1967) proposed survival of at least five species of tiger beetles ( Cicindela : Cicindelidae) in the
vicinity of the Nebraska Sand Hills.
Following Wisconsin glaciation, many sand dunes developed and older ones became
reactivated. Figure 10c illustrates proposed dispersal patterns for P. albopilosus Cresson from
refugia.
Expansion of Philanthus albopilosus manuelito onto the Great Plains should have allowed
rapid colonization of most prairie sand dunes. Northward expansion may have been limited by
presence of Philanthus albopilosus albopilosus acting as a cryptic species, genetically distinct,
yet morphologically similar. Rare occurrence of Philanthus albopilosus manuelito in the Great
basin may be the result of relatively recent post-Pleistocene dispersal.
EVOLUTION OF COLOUR PATTERN AND BEHAVIOURAL FEATURES OF
P. ALBOPILOSUS
Colour pattern features of P. albopilosus
Ancestral P. albopilosus were probably similar to P. a. manuelito . Extensive yellow
contrasted with black, in typical warning maculation pattern, is similar to that seen on other
sand dune sphecids (e.g. Microbembex, P. psyche).
Adaptation to a cooler climate during Wisconsinan glaciation may have been the major
evolutionary reason for differentiation by populations in the northern refugium. Females of
northern, P. a. albopilosus are dark: this may be a functional adaptation for absorbing radiant
heat, which may prolong the length of time per day in which they could hunt. Males may not
require much darkening, as they could restrict their activities to the hot microenvironment of
the sand dune surface.
The very pale form of P. a. manuelito of the great plains is somewhat of a puzzle: Character
displacement was suggested, but does not adequately explain populations of area 4, unless,
there is considerable gene flow between populations on the plains. Mountain barriers would
buffer populations in Areas 2 and 3 from influx of extreme pale forms.
Present distribution of P. a. albopilosus is the result of post-Pleistocene dispersal into new
dune habitats. Eastern and northwestern expansion of range has occurred. Southward
expansion may have been blocked by P. a. manuelito . There may be enough genetic and
behavioural differences reducing the tendency for the two groups to hybridize (cryptic species).
A few northern populations contain well marked individuals. This may be the result of reversed
selection toward a form with typical warning colouration pattern or they may represent
remnants of the ancestral gene pool or limited introgression.
The above discussion is primarily based on variation in the females; trends are present in
males but are less marked.
Behavioural features of P. albopilosus
Evolutionary adaptations exhibited by P. albopilosus are similar to other digger wasps
living in sand dune environments. Parallel evolution in development of false burrows originated
from nest parasite pressures, which are intense in open environments. Orientation to the nest is
difficult for females, as landmarks near the nest may change in an unstable dune environment.
Use of distant landmarks is important, but fixation to these requires seeing the nest site while
orienting and keeping nest parasites out. False burrows may serve both of these functions.
Quaest. Ent., 1982, 18 (1-4)
Philanthus albopilosus Cresson
123
Short occupancy time of the nest as compared to other species of Philanthus may be due to
friable soil. Little time is spent digging in comparison to species which dig in hard soils. Few
cells per nest would be a function of the ephemeral occupancy coupled with slow provisioning.
A relatively long adult life span, nest location in areas devoid of vegetation and anti-parasite
nest building behaviours may reduce selection pressures for the production of large numbers of
offspring (“k” selection).
Reduction in number of generations per year in northern populations is probably a result of
shorter growing seasons either presently or during the species evolutionary history.
Change in prey specificity from the ancestral limits may be a response to reduction in
number and species of bees available during Wisconsinan time. Alternate prey species would be
incorporated from other groups of Apocrita.
These facets of life history serve to characterize P. albopilosus as a whole. This also serves
in part, as a start in characterization of the northern fauna of Philanthus.
ACKNOWLEDGEMENTS
I wish to express my gratitude to the following people for assistance in this study; G. E. Ball,
my thesis supervisor and sponsor who critically and constructively assisted and guided this
manuscript to completion. W. G. Evans and A. L. Steiner for advice and assistance through the
course of the study.
I would like to thank R. M. Bohart for assistance in identification of eumenid and sphecid
wasps. Thanks are due to the curators of museums from which material was borrowed.
Thanks are extended to J. L. and B. F. Carr for my introduction to the wonders of sand dune
communities; F. A. H. Sperling for assistance in drafting illustrations and checking the
manuscript, and J. R. Spence for critically reviewing this paper.
Financial assistance is acknowledged from The National Research Council of Canada grant
A 1399 held by G. E. Ball and the Entomological Society of Canada Postgraduate Award.
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Hilchie
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Quaest. Ent., 1982, 18 (1-4)
RHYSODINI OF THE WORLD
PART III. REVISION OF OMOGLYMMIUS GANGLBAUER (COLEOPTERA:
CARABIDAE OR RHYSODIDAE) AND SUBSTITUTIONS FOR PREOCCUPIED
GENERIC NAMES
Ross T. Bell
Department of Zoology
University of Vermont
Burlington, VERMONT 05405
U.S.A.
Quaestiones Entomologicae
18: 127-259 1982
Joyce R. Bell
24 East Terrace
South Burlington, VERMONT 05401
U.S.A.
ABSTRACT
This paper is third of a series which will constitute a revision of the Rhysodini of the
world. Two replacements are proposed for genera described in Part 1: Kupeus NEW NAME,
for Kupea Bell and Bell 1978, preoccupied by Kupea Philpott 1930, and Tangarona NEW
NAME for Tangaroa Bell and Bell 1978, preoccupied by Tangaroa Lehtinnen 1967. The
remainder of the paper is a revision of Omoglymmius Ganglbauer. Pyxiglymmius Bell and
Bell is reduced to the rank of subgenus. The subgeneric classification is extensively altered:
eleven subgenera are recognized, compared with five in Part I. New subgenera are:
Boreoglymmius NEW SUBGENUS, type- Omoglymmius americanus ( Castelnau ), three spp;
North America, Japan; Laminoglymmius NEW SUBGENUS, type- Omoglymmius insularis
(Grouvelle), seven spp. Malay Peninsula, Sumatra, Borneo, Nicobar Is., New Guinea;
Caeconavitia NEW SUBGENUS, ^pe-Omoglymmius zimmermani Bell and Bell, one species,
Fiji; Indoglymmius NEW SUBGENUS, type- Omoglymmius lineatus ( Grouvelle ), one species
south India; Carinoglymmius NEW SUBGENUS, type- Omoglymmius carinatus ( Grouvelle ),
three spp., Sumatra, Borneo, Andaman Is.
The following new species are described ( type localities indicated): Omoglymmius
(Hemiglymmius) hemipunctatus (SUMATRA (?), Serdang, Lau Raku); O. (H.) occultus
(SUMATRA, only); O. (H.) rimatus (JAVA, Mt. Kawi); O. (H.) inermis (SUMATRA, only);
O. (Pyxiglymmius) cristatus (PHILLIPPINE IS., Luzon, Mt. Makilung); O. (P.) hesperus
(M ENT AWE I Is. Sipora, Sereina); O. (P.) krikkeni (SUMATRA: Alas Valley, Gumpang); O.
(Laminoglymmius) inaequalis (NICOBAR IS.); O. (L.) actae (NEW GUINEA, Maffin Bay);
O. (L.) trisinuatus (SUMATRA (?), Bangung); O. (L.) gorgo (SUMATRA, Siantar); O.
(Navitia) stylatus (NEW HEBRIDES: Malekula); O. (Nitiglymmius) semioculatus
(PHILIPPINE IS., Siargao, Dapa); O. (Orthoglymmius) microtis (RYUKYU IS.: Ishigaki,
Takeda); O. (sensu stricto) imugani (PHILIPPINE IS., Luzon, Imugan; O. (s. str.) politus
(PHILIPPINE IS., Luzon, Mt. Polis; O. (s. str.) crassicornis (PHILIPPINE IS., Negros,
Horns of Negros); O. (s. str.) amplus (SUMATRA, Palembang); O. (s. str.) modiglianii
128
Bell and Bell
( MENTAWEI IS., Si Oban); O. (s. str.) evasus ( PHILIPPINE IS., Mindanao, E. Slope Mt.
McKinley ); O. (s. str.) nemoralis (SARAWAK, Mt. Matang); O. (s. str.) fraudulentus
( SUMATRA , Palembang); O. (s. str.) coelebs ( PHILIPPINE IS., Palawan, Binaluan); O (s.
str.) thoracicus (JAVA, Tangk. Prahoe); O. (s. str.) summissus (SUMATRA, Palembang); O.
(s. str.) semperi (PHILIPPINE IS., only); O. (s. str.) data (PHILIPPINE IS., Luzon, Mt.
Data); O. (s. str.) hiekei (PHILIPPINE IS., Luzon, Dalbalan); O. (s. str.) pectoralis (JAVA:
Mt. Smetou); O. (s. str.) quadruplex (PHILIPPINE IS., Mindanao, Zamboanga, Kabasalan);
O. (s. str.) duplex (PHILIPPINE IS., Luzon, Mt. Makiling); O. (s. str.) bouchardi
(SUMATRA, Palembang); O. (s. str.) consors (SUMATRA, Palembang); O. (s. str.) repetitus
(CELEBES, Tolitoli); O. (s. str.) opticus (LESSER SUNDA IS., Dammar Is.); O. (s. str.)
viduus (KEI IS.); O. (s. str.) continuus (MOLUCCA IS., Sula Is., Mangole); O. (s. str.)
wittmeri (MOLUCCA IS., Sula Is., Mangole); O. (s. str.) vadosus (MOLUCCA IS., Amboina);
O. (s. str.) morditus (MOLUCCA IS., Morotai ); O. (s. str.) nasalis (MOLUCCA IS., Boeroe,
Kajeli);0. (s. str.) bicarinatus (SCHOUTEN IS., Jobi); O. (s. str.) lindrothi (SOLOMON IS.,
Guadalcanal, Kukum); O. (s. str.) modicus (SOLOMON IS., Savo); O. (s. str.) rusticus
(SOLOMON IS., Russell Is., Loani); O. (s. str.) manni (SOLOMON IS., Malaita, Auki); O.
(s. str.) regius (SOLOMON IS., Isabel, Regi); O. (s. str.) gurneyi (SOLOMON IS.,
Bougainville); O. (s. str.) princeps (SOLOMON IS., Bougainville, Kokure, Crown Prince Ra.);
O. (s. str.) renutus (SOLOMON IS., New Georgia, Mavovo); O. (s. str.) scopulinus (SANTA
CRUZ IS., Reef I); O. (s. str.) mycteroides (SOLOMON IS., Kolombangara, Hunda); O. (s.
str.) tabulatus (SOLOMON IS., Boku); O. (s. str.) classicus (ADMIRALTY IS., Los Negros);
O. (s. str.) oroensis (NEW GUINEA, Oro Bay); O. (s. str.) fringillus (NEW GUINEA, Huon
Peninsula, Finschhafen ); O. (s. str.) puncticornis (NEW GUINEA, Fly R., Kiunga); O. (s. str.)
trepidus (NEW GUINEA, Wau, Morobe Dist .); O. (s. str.) patens (NEW GUINEA, Maffin
Bay); O. (s. str.) cavea (NEW GUINEA, Ramoi Is.); O. (s. str.) sectatus (NEW GUINEA, Mt.
Missim); O. (s. str.) ephemeris (NEW GUINEA, Nabire, s. Geelvink Bay); O. (s. str.) follis
(NEW GUINEA, Wau, Morobe Dist.); O. (s. str.) iridescens (NEW GUINEA, Araboebivak);
O. (s. str.) massa (NEW GUINEA, Eliptamin Val); O. (s. str.) biroi (NEW GUINEA,
Sattelberg); O. (s. str.) denticulatus (NEW GUINEA, Wissel Lakes, Enarotadi); O. (s. str.)
auratus (NEW GUINEA, Swart Val.); O. (s. str.) sus (NEW GUINEA, Katau); O. (s. str.)
planiceps (NEW GUINEA, Wau, Morobe Dist.); O. (s. str.) lentus (NEW GUINEA,
Sattelberg); O. (s. str.) asetatus (NEW GUINEA, Madang); O. (s. str.) sedlaceki (NEW
GUINEA, Wau, Morobe Dist.).
O. (Pyxiglymmius) aterrimus (Chevrolat) is synonymous with O. (P.) strabus (Newman).
RESUME
Cet article est la troisieme d'une serie qui constiteront une revue taxonomique des Rhysodini du monde. Deux
nommes remplaces des autres car ils sont preoccupees: Kupeus NOUVEAU NOM pour Kupea Bell et Bell 1978,
preoccupee de Kupea Philpott, 1930, et Tangarona NOUVEAU NOM pour Tangaroa Bell et Bell 1978, preoccupee de
Tangaroa Lehtinnen 1967. Le reste de Particle est une revision taxonomique du genre Omoglymmius Ganglbauer.
Pyxiglymmius Bell et Bell est redui au rang de sous-genre. Nous changeon beaucoup la classification des sous-genres:
nous reconnaissons onze sous-genres en comparaison de cinque sous-genres en Partie I. Les sous-genres nouveaux sont:
Boreoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius americanus (Castelnau), trois spp. Amerique du Nord,
Japon; Laminoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius insularis (Grouvelle), sept, spp., Peninsule de
Malaisie, Sumatra, Borneo, lies Nicobars, Nouvelle Guinee; Caeconavitia NOUVEAU SOUS-GENRE, type-
Omoglymmius zimmermani Bell et Bell, une sp., Fiji: Indoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius
lineatus ( Grouvelle ), une sp.. Inde du sud; Carinoglymmius NOUVEAU SOUS-GENRE, type- Omoglymmius carinatus
( Grouvelle ), trois spp., Sumatra, Borneo, lies Andamans.
On decrit les especes nouvelles qui voice (en indiquant pour chacune la localite du specimen type): Omoglymmius
(Hemiglymmius) hemipunctatus (SUMATRA (?), Serdang, Lau, Raku); O. (H.) occultus (SUMATRA sans localite
Revision of Omoglymmius Ganglbauer
129
specifie); O. (H.) rimatus (JAVA, Mt. Kawi); O. (H.) inermis ( SUMATRA , sans localite specific); O. (Pyxiglymmius)
cristatus (ILES PHILIPPINES, Luzon, Mt. Makiling); O. (P.) hesperus (ILES DE MENTAWEI, Sipora, Sereina ); O.
(P.) krikkeni (SUMATRA, vallee d'Alas, Gumpang ); O. (Laminoglymmius) inaequalis (lies Nicobars ); O. (L.) actae
(NOUVELLE GUINEE, Baie de Maffin); O. (L.) trisinuatus (SUMATRA (?), Bangung); O. (L.) gorgo (SUMATRA,
Siantar ); O. (Navitia) stylatus (NOUVELLE HEBRIDES, Malekula); O. (Nitiglymmius) semioculatus (ILES
PHILIPPINES, Siargao, Dapa); O. (Orthoglymmius) microtis (ILES DE RYUKYU: Ishigaki, Takeda); O. (sensu
stricto) imugani (ILES PHILIPPINES, Luzon, Imugan); O. (s. str.) politus (ILES PHILIPPINES, Luzon, Mt. Polis);
O. (s. str.) crassicornis (ILES PHILIPPINES, Negros, Horns of Negros); O. (s. str.) amplus (SUMATRA, Palembang );
O. (s. str.) modiglianii flLES DE MENTAWEI, Si Oban); O. (s. str.) evasus (ILES PHILIPPINES, Mindanao,
versant est du Mont McKinley); O. (s. str.) nemoralis (SARAWAK, Mt. Matang); O. (s. str.) fraudulentus
(SUMATRA, Palembang); O. (s. str.) coelebs (ILES PHILIPPINES, Palawan, Binaluan); O. (s. str.) thoracicus
(JAVA, Tangk. Prahoe); O. (s. str.) summissus (SUMATRA, Palambang); O. (s. str.) semperi (ILES PHILIPPINES,
san localite specifie); O. (s. str.) data (ILES PHILIPPINES, Luzon, Mt. Data); O. (s. str.) hiekei (ILES
PHILIPPINES, Luzon, Dalbalan); O. (s. str.) pectoralis (JAVA, Mt. Smetou); O. (s. str.) quadruplex (ILES
PHILIPPINES, Mindanao, Zamboanga, Kabasalan); O. (s. str.) duplex (ILES PHILIPPINES, Luzon, Mt. Makiling);
O. (s. str.) bouchardi (SUMATRA, Palembang); O. (s. str.) consors (SUMATRA, Palembang); O. (s. str.) repetitus
(CELEBES, Tolitoli); O. (s. str.) opticus (PETITES ILES DE SONDE, lies de Dammar); O. (s. str.) viduus (ILES
DE KEI); O. (s. str.) continuus (ILES MOLUQUES, lies de Sula, Mangole); O. (s. str.) wittmeri (ILES MOLUQUES,
lies de Sula, Mangole); O. (s. str.) vadosus (ILES MOLUQUES, Amboina); O. (s. str.) morditus (ILES MOLUQUES,
Moratai); O. (s. str.) nasalis flLES MOLUQUES, Boeroe, Kajeli), O. (s. str.) bicarinatus (ILES DE SCHOUTEN,
Jobi); O. (s. str.) lindrothi (ILES DE SOLOMON, Guadalcanal, Kukum); O. (s. str.) modicus (ILES DE SOLOMON,
Savo); O. (s. str.) rusticus (ILES DE SOLOMON, lies de Russell, Loani); O. (s. str.) manni (ILES DE SOLOMON,
Malaita, Auki); O. (s. str.) regius (ILES DE SOLOMON, Isabel, Regi); O. (s. str.) gurneyi (ILES DE SOLOMON;
Bougainville); O. (s. str.) princeps (ILES DE SOLOMON , Bouganville, Kokure, Crown Prince Ra.); O. (s. str.) renutus
(ILES DE SOLOMON , Nouveau Georgia, Mavovo); O. (s. str.) scopulinus (ILES SANTA CRUZ, lie de Reef); O. (s.
str.) mycteroides (ILES DE SOLOMON, Kolombangara, Hunda); O. (s. str.) tabulatus (ILES SOLOMON, Boku); O.
(S. str.) classicus flLES D’ ADMIRALTY, Los Negros); O. (s. str.) oroensis (NOUVELLE GUINEE, Baie d'Oro ); O. (s.
str.) fringillus (NOUVELLE GUINEE, Huon Peninsula, Finschhafen); O. (s. str.) puncticornis (NOUVELLE GUINEE,
Fly R., Kiunga); O. (s. str.) trepidus (NOUVELLE GUINEE, Wau, Morobe Dist .); O. (s. str.) patens (NOUVELLE
GUINEE, Baie de Maffin ); O. (s. str.) cavea (NOUVELLE GUINEE, ‘lie Ramoi); O. (s. str.) sectatus (NOUVELLE
GUINEE, Mt. Missim); O. (s. str.) ephemeris (NOUVELLE GUINEE, Nabire, sud de la Baie de Geelvink); O. (s. str.)
follis (NOUVELLE GUINEE, Wau, Morobe Dist.); O. (s. str.) iridescens (NOUVELLE GUINEE, Araboebivak); O. (s.
str.) massa (NOUVELLE GUINEE, Eliptamin Val.); O. (s. str.) biroi (NOUVELLE GUINEE, Sattelberg); O. (s. str.)
denticulatus (NOUVELLE GUINEE, Lacs de Wissel, Enarotadi); O. (s. str.) auratus (NOUVELLE GUINEE, Swart
Val.); O. (s. str.) sus (NOUVELLE GUINEE, Katau); O. (s. str.) planiceps (NOUVELLE GUINEE, Wau, Morobe
Dist.); O. (s. str.) lentus (NOUVELLE GUINEE, Sattelberg); O. (s. str.) asetatus (NOUVELLE GUINEE, Madang); O.
(s. str.) sedlaceki (NOUVELLE GUINEE, Wau, Morobe Dist.).
Nous decouvrames que O. (Pyxiglymmius) aterrimus (Chevrolat) est un synonyme de O. (P.) strabus NEWMAN
TABLE OF CONTENTS
Introduction 1 30
Sources of Material 1 30
Genus Omoglymmius 131
Subgenus Hemiglymmius (Figs. 1-16) 132
New Subgenus Boreoglymmius (Figs. 17-26) 140
Subgenus Pyxiglymmius (Figs. 27-56) 146
New Subgenus Laminoglymmi us (Figs. 57-74) 1 56
Subgenus Navitia (Figs. 75,76,79,80) 164
New Subgenus Caeconavitia 166
New Subgenus Indoglymmius (Figs. 77,83) 1 67
Subgenus Nitiglymmius (Figs. 78,8 1 ,82) 168
Subgenus Orthoglymmius (Figs. 84-103) 169
New Subgenus Carinoglymmius (Figs. 104-1 12) 176
Subgenus Omoglymmius sensu stricto (Figs. 1 1 3-237) 180
Substitutions for Preoccupied Generic Names 253
Quaest. Ent., 1982, 18 (1-4)
130
Bell and Bell
Aeknowledgements 254
Supplementary References 254
INTRODUCTION
This paper, third of a series of five, comprises a revision of the large and difficult genus
Omoglymmius Ganglbauer. Pyxiglymmius Bell and Bell, formerly regarded by us as a
separate genus, is included as a subgenus. The discovery of many additional species has resulted
in several changes in limits and definitions of subgenera. Two generic names proposed in Part I
of this series are preoccupied, and substitutes are proposed.
SOURCES OF MATERIAL
The following abbreviations designate collections cited in this paper. Names of curators of
respective institutions are in parentheses.
AMS Instituut voor Taxonomische Zoologie, Amsterdam, Netherlands (J. Duffels)
AP U.S. Dept, of Agriculture, Harrisburg, PA (K. Valley)
ARK University of Arkansas, Fayetteville (E. P. Rouse)
AU S. F. Austin State University, Nacogdoches, Texas (W. W. Gibson)
BMNH British Museum, Natural History, London (R. Pope)
BPBM Bernice P. Bishop Museum, Honolulu (G. Samuelson)
BPM Barry P. Moore, Canberra City, Australia
BSL Naturhistorisches Museum, Basel, Switzerland (W. Wittmer)
CAG U.S. Dept, of Agriculture, Sacramento, CA (F. G. Andrews)
CAS California Academy of Sciences, San Francisco (D. H. Kavanaugh)
CMP Carnegie Museum of Natural History, Pittsburgh, PA (G. Wallace)
CNHM Field Museum of Natural History, Chicago, IL (H. Dybas)
CU Cornell University, Ithaca, NY (L. L. Pechuman)
DM Dayton Museum, Ohio (A. J. Koestner)
DY Daniel K. Young, E. Lansing, MI
FLA U.S. Dept, of Agriculture, Gainesville, FL (R. Woodruff)
GEN Museo Civico di Storia Naturale G. Doria , Genoa (R. Poggi)
GLP Gary L. Peters, Corvallis, OR
HL Harry J. Lee, Fairview Park, OH
IO Iowa State University, Ames (R. Miller)
ISNHS Illinois State Natural History Survey, Urbana (M. Sanderson)
IU Indiana University, Bloomington
KS Karl Stephan, Tucson, AZ
KU Kagoshima University, Japan
LA Los Angeles County Natural History Museum, CA (C. L. Hogue)
LEI Rijksmuseum von Natuurlijke Historie, Leiden, Netherlands (J. Krikken)
LEN Academy of Sciences, Leningrad (O. Kryzhanovskij)
LS Louisiana State University, Baton Rouge (J. B. Chapin)
LUN Zoological Institute, Lund, Sweden (R. Danielsson)
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA
Revision of Omoglymmius Ganglbauer
131
(J. H. Lawrence)
MNHB Museum fur Naturkunde der Humboldt-Universitat, Berlin, DDR (F. Hieke)
MNHN Museum National d’Histoire Naturelle, Paris, France (A. Descarpentries)
MN University of Minnesota, St. Paul (P. J. Clausen)
MO University of Missouri, Columbia (W. R. Enns)
MRAC Musee Royal de l’Afrique Centrale, Tervuren, Belgium (P. Basilewsky)
MSU Michigan State University, E. Lansing
NC North Carolina State University, Raleigh (D. A. Young)
NMNH United States National Museum of Natural History, Washington, D. C. (T. L.
Erwin)
NMNZ National Museum of New Zealand, Wellington (R. G. Ordish)
NMW Naturhistorisches Museum Wien, Austria (F. Janczyk)
OK Oklahoma State University, Stillwater (W. A. Drew)
OS Oregon State University, Corvallis (G. L. Peters)
OSFS Oregon State Forest Sciences Collection, Corvallis
OSU Ohio State University, Columbus (C. A. Triplehorn)
PA Academy of Sciences, Philadelphia (D. C. Rentz)
PU Purdue University, Lafayette, IN
RCG R. C. Graves, Bowling Green, OH
SATO Masataka Sato, Nagoya, Japan
SDA U.S. Dept, of Agriculture, Brookings, SD (V. M. Kirk)
SI Southern Illinois University, Carbondale (J. E. McPherson)
TB Thomas Barr, University of Kentucky, Lexington
UCD University of California, Davis
UK University of Kansas, Lawrence (G. W. Byers)
UL University of Louisville, KY (C. V. Covell)
UM University of Michigan, Ann Arbor (I. J. Cantrall)
UN University of Nebraska, Lincoln (B. C. Ratcliffe)
UT Utah State University, Logan (W. J. Hanson)
UW University of Wisconsin, Madison (J. R. Baker)
VP Virginia Polytechnic Institute, Blacksburg (M. Kosztarab)
WR William Rosenberg
WRS Walter R. Suter, Carthage College, Kenosha, WI
WS Washington State University, Pullman (W. J. Turner)
Genus Omoglymmius Ganglbauer 1892
Type species. - Rhysodes germari Ganglbauer 1892
Description. — Paramedian grooves of pronotum deep, distinct; middle and hind tibiae each with a single terminal
spur; antennal stylet, basal setae, midial angles of temporal lobes variously developed.
The description has been altered from that in Part I, p. 72, to reflect incorporation of
Pyxiglymmius as a subgenus. In the key to genera of Omoglymmiina, Part I, pp. 66-67, adults
of Omoglymmius as presently defined, trace to 5'. The key to subgenera presented below
supersedes that in Part I, p. 72.
Quaest. Ent., 1982, 18 (1-4)
132
Bell and Bell
KEY TO SUBGENERA
1
v
2 (1)
2'
3 (20
3'
4 (3)
4/
5 (4)
5'
6 (50
6'
7 (40
7
8 (70
8'
9 (30
9'
10(90
10'
Clypeal setae present Orthoglymmius Bell and Bell, p. 169
Clypeal setae absent
Spur of middle tibia straight; female with deepest lateral pits on Sternum V;
outer carina very narrow, 0.33 or less as wide as paramedian groove
Carinoglymmius new subgenus, p. 176
Spur of middle tibia curved anteriorly; female with deepest pits on Sternum IV;
outer carina in most species subequal in width to paramedian groove, or only
slightly narrower (in O. ( s . str.) solitarius and O. ( Laminoglymmius ) inaequalis
0.33 as wide as paramedian groove)
Apical stylet of antenna absent
Apical stylet present
Basal setae present on outer antennal segments
Basal setae absent
Medial margin of temporal lobe with broad emargination between two widely
separated medial angles Pyxiglymmius Bell and Bell, p. 146
Medial margin of temporal lobe without emargination or with one or two
shallow ones; each temporal lobe with one medial angle or with two or three
close together
Medial margin with translucent area (ill-defined in O. actae ); medial margin
with distinct emarginations (except in O. inaequalis )
Laminoglymmius new subgenus, p. 1 56
Medial margin without translucent area and without emargination, medial
angle single Boreoglymmius new subgenus, p. 140
Medial angles with translucent area Indoglymmius new subgenus, p. 167
Medial angles without translucent area
Eye large, over 0.5 as deep as head; cornea distinctly facetted
Omoglymmius sensu stricto Ganglbauer, p. 180
Eye reduced; less than 0.25 as deep as head; cornea not facetted
Nitiglymmius Bell and Bell, p. 168
Outer antennal segments with basal setae. Hemiglymmius Bell and Bell, p. 132
Basal setae absent
Eyes large, cornea facetted; frontal grooves obsolete
Navitia Bell and Bell, p. 164
Eye reduced, less than 0.25 as deep as head; cornea not facetted; frontal grooves
deep Caeconavitia new subgenus, p. 166
2
3
4
9
5
7
6
8
Subgenus Hemiglymmius Bell and Bell 1978
Type species. - Rhysodes africanus Grouvelle 1892.
Description. — Basal setae on at least antennal Segments IX, X (in most species on more proximal segments as
well); antennal stylet present; clypeal setae absent; eye large, normal, cornea facetted; frontal grooves deep, complete,
dilated in many species; medial angle simple, no translucent area; postorbital tubercle present; marginal groove of
pronotum deep; spur of middle tibia curved anteriorly; punctures of abdominal Sterna lll-V scattered; lateral pits of
abdomen in female on Sternum IV, poorly developed.
Revision of Omoglymmius Ganglbauer
133
This subgenus differs from others that have an antennal stylet and basal setae in having
simple, unmodified medial angles on the temporal lobes.
Limits of this subgenus have been altered by removal of those species which lack the
antennal stylet ( O . americanus, O. hamatus, and O. lewisi ) to Boreoglymmius new subgenus.
Restudy of types has shown that O. oberthueri belongs in Laminoglymmius new subgenus,
while O. nicobarensis belongs in Orthoglymmius Bell and Bell. We also originally listed O.
borneensis among the species of Hemiglymmius. We have not been able to locate the type for
this species, but a detailed study of the original description suggests that the name is based on a
species of Omoglymmius s. str. from Borneo, and not on the species formerly interpreted by us
as being O. borneensis.
Range of the subgenus Hemiglymmius as limited above includes Java, Borneo, and
Sumatra, and a large part of Africa, from Liberia to eastern Zaire. O. germaini , described from
Bolivia, is certainly a member of this subgenus. However, it is very similar to one of the two
species forming Grouvelle’s concept of O. javanicus, so it seems certain that locality data
associated with the types is erroneous.
Phylogeny. - O. africanus, with its narrow, abbreviated median lobe, appears to stand apart
from the Indonesian species. The latter constitute a closely related and poorly known complex
whose interrelationships will not be easy to decipher until both sexes of all species have been
collected.
KEY TO SPECIES
1 Median lobe of head narrow, linear, short, ended opposite anterior margin of
eye; gular tubercle absent; male without ventral tooth on anterior femur
O. africanus (Grouvelle), p. 135
1 ' Median lobe narrow to broad, long, extended at least to level of middle of eye;
gular tubercle present; male with ventral tooth on anterior femur 2
2 (L) Temporal lobe not extensively pollinose, its glabrous portion separated from
antennal lobe only by pollinosity of antennal groove 3
2' Temporal lobe extensively pollinose anteriorly, its glabrous portion broadly
separated from antennal lobe 6
3 (2) Inner carina of pronotum punctate 4
3' Inner carina impunctate O. hemipunctatus new species, p. 137
4 (3) Temporal lobe with 18-20 coarse punctures; Stria IV with five or six setae along
its length O. occultus new species, p. 138
4' Temporal lobe with one to five punctures, or none; Stria IV with one or two
setae near apex 5
5 (4') Pronotum widest anterior to middle; outer carina very broad anteriorly
O. germaini (Grouvelle), p. 137
5' Pronotum broadest at middle; outer carina less broad anteriorly
O. javanicus (Grouvelle), p. 136
6 (2') Median lobe glabrous; antennal lobe bounded posteriorly by prominent scarp
O. rimatus new species, p. 139
6' Median lobe pollinose; antennal lobe not bounded posteriorly by scarp or with
only a vestige of one 7
7 (6') Male with proximal tooth on anterior tibia; pronotum shorter, length/greatest
Quaest. Ent., 1982, 18 (1-4)
134
Bell and Bell
Plate 1. Figures 1-16, Subgenus Hemiglymmius. Figs. 1-8, Head and pronotum, dorsal aspect; Fig. 1, Omoglymmius
(Hemiglymmius) africanus (Grouvelle); Fig. 2, O. (H.) javanicus (Grouvelle); Fig. 3, O. (H.) germaini (Grouvelle); Fig. 4,
O. (H.) hemipunctatus new species; Fig. 5, O. (H.) occultus new species; Fig. 6, O. (H.) ineditus (Dajoz); Fig. 7, O. (H.)
rimatus new species; Fig. 8, O. (H.)inermis new species; Figs. 9-16, Head, lateral aspect; Fig. 9, O. (H.) africanus
(Grouvelle); Fig. 10, O. (H.) javanicus (Grouvelle); Fig. 11,0. (H.) germaini (Grouvelle); Fig. 12, O. (H.) hemipunctatus
new species; Fig. 13, O. (H.) occultus new species; Fig. 14 O. (H.) ineditus (Dajoz); Fig. 15 O. (H.) rimatus new species;
Fig. 16 O. (H.) inermis new species.
Revision of Omoglymmius Ganglbauer
135
width about 1.26; calcars larger O. ineditus (Dajoz), p. 138
T Male without proximal tooth on anterior tibia; pronotum longer, length/greatest
width about 1 / 36; calcars smaller O. inermis new species, p. 140
Omoglymmius ( Hemiglymmius ) africanus (Grouvelle 1892)
Figs. 1, 9
Rhysodes africanus Grouvelle 1892: 299.
Omoglymmius ( Hemiglymmius ) africanus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE sex not recorded, labelled: “Assinie,” no other data
(MNHN). According to the original description there were two specimens in the type series, so
the second specimen, if still extant, is a paralectotype. Assinie was a former French colony,
centering about the town of the same name, now part of the Ivory Coast. A specimen labelled
“Lac Albert (Albrecht), Afrique Allemande” (MNHN), is labelled as a paratype, but is
probably not one in modern terms, it is not mentioned in the original description. The locality is
ambiguous, as Lake Albert was not in German East Africa.
Description. — Length 5. 8-6. 7 mm. Antennal stylet short, acute; basal setae well-developed on Segments IX, X,
sparse on Segments V I- V III; absent from Segments IV, V; Segments I-XI coarsely punctate; median lobe narrow, with
sides parallel, very short, tip even with anterior margin of eye, apex obtuse; median lobe varied from coarsely punctate to
impunctate; temporal lobe with medial angles nearly rectangular, rather widely separated in most specimens (but nearly
contiguous in some); posteriomedial margin oblique between medial and occipital angles; temporal lobe varied from finely
punctate to impunctate; distinct postantennal depression between antennal base and eye, extended medially from lateral
margin, but not to frontal groove; temporal setae absent; postorbit entirely finely pollinose; postorbital tubercle
well-developed, in most specimens with well-developed pilosity conspicuously coarser and longer than pollinosity of
postorbit.
Pronotum relatively short, length/greatest width averaging 1.18; widest at middle; sides curved, convergent both
basally and apically; margin not sinuate anterior to hind angle; median groove relatively broad, about 0.33 as broad as
inner carina at middle; marginal groove dilated, as wide as outer Carina; both carinae narrow, inner slightly narrower than
outer one; inner carina impunctate (in some specimens one or two); outer carina finely punctate.
Elytra relatively long and narrow (though distinctly wider than pronotum); elytral striae broad, rather shallow; strial
punctures large, deep, pilose; intervals convex, very narrow; each interval distinctly narrower than a strial puncture;
pattern of setae varied, when fully developed more extensive than in any other member of subgenus; Stria I with one to
three in apex; Stria II with three in apex; Stria IV with continuous series of six to seven setae; medial side of apical
tubercle with single seta; apical stride with three or four setae; apex of Stria VII with seven to ten setae; some specimens
without certain setae (most frequently, setae absent from middle portion of Stria IV); hind coxa without setae; abdominal
Sternum VI with pair of setae in both sexes; female with shallow, indistinct lateral pits on Sterna IV and V; male without
ventral tooth on anterior femur and without proximal tooth on anterior tibia; middle tibia without trace of calcar; calcar of
hind tribia very small, triangular, acute.
The short, narrow median lobe gives adults of this species a distinctive appearance, in most
specimens enhanced by well-separated medial angles, which make the head appear almost
“split”. Variation is extensive. A few specimens, especially those from the most eastern part of
the range, have the medial angles almost contiguous. Amount of punctation of medial and
temporal lobes is markedly variable, and there is also considerable variation in development of
elytral setae. Neither of these two characters seem to vary geographically. Perhaps more than
one species is represented among African Hemiglymmius , although we can see no clear-cut
pattern in the material available to us.
Range. - This is extensive for a rhysodine, from Liberia on the west to the eastern borders of
Zaire in the east. In addition to the type series, we have seen specimens from the following
localities;CAMEROONS: one male, two females, Lolodorf, coll. G. Schwab (MCZ); one male,
“Neu-Kamerun” No. 3209-48, coll. Tessmann, S.G. (MNHB); EQUATORIAL GUINEA: one female, Nkolentangan,
XI-07.-V-08, coll. G. Tessman, S.G. (MNHB); one male. Is. Fernando Poo, Musola, 500-800 m.s.m., I- 1 1 1- 1 902, coll. L.
Fea (GEN); LIBERIA: four males, three females, Mt. Coffee, Feb-March 1895, O.F. Cook Colin. (NMNH); ZAIRE:
Quaest. Ent., 1982, 18 (1-4)
136
Bell and Bell
(all specimens in MRAC) one male, Haute-Uele: Moto, 1923, coll. L. Burgeon; one female, Kasai: Ngombe,-1921, coll.
Dr. H. Schouteden; one female, Ituri: Yindi, 18-VI-1948, coll. A.E. Bertrand; one female, Lulua: Kapanga, X-1933,
coll. G.F. Overlaet; one female, Mayidi, 1942, coll. Rev. P. Van Eyen; two males, Territoire de Sandoa, 18-XI-1948,
coll. N. Leleup; one male, two females, Tschuapa, Lac Tumba, Mabali, 350 m., 29- 1 X- 1955, coll. N. Leleup, under
bark of dead tree.
Omoglymmius { Hemiglymmius ) javanicus (Grouvelle)
Figs. 2, 10
Rhysodes javanicus Grouvelle 1903: 1 10.
Omoglymmius (Hemiglymmius) javanicus (Grouvelle) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) male labelled: “JAVA, Gounod Gedeh,
Ledru 1898, TYPE” (MNHN); PARALECTOTYPES one male, two females, same labels as
lectotype (MNHN); one female with same label as lectotype and additional label “Rhysodes
javanicus ty. Grouv.” (MNHN); one female, four males with same label as lectotype and
additional labels “R. javanicus ty. Grouv., 1952-Oberthiir Colin.”, “Syn. Rhysod. A Grouvelle
1903” (MNHN); one male, with same label as lectotype and additional labels “ex. coll.
Oberthiir 1904-175, syn, Rhysod. A. Grouvelle, 1903, co-type, javanicus Grouv. ex typis”
(BMNH).
A female specimen, labelled: “Tougou, Java Occid., J.D. Pasteur 3-96, Type, Rhysodes
javanicus Grouv.” (MNHN), is not conspecific with the series from Gounod Gedeh, but is
probably O. ineditus (Dajoz) or O. rimatus n. sp.
Description. — Length 6. 7-8.0 mm. Basal setae well developed on antennal Segments V-X; Segments I- 1 V each
with pollinose band on dorsal surface; Segments V-X coarsely punctate; Segment XI distinctly longer than wide; corpus
cylindrical, narrower than Segment X; stylet short, acute.
Head in most specimens slightly longer than wide (average of 11 specimens width/length 0.963; range 0.909-1.02);
median lobe impunctate, glabrous, elongate oval, its tip bluntly rounded, just anterior to level of posterior margin of eye;
frontal space 1.5 wider than long; medial angles slightly obtuse, contiguous; temporal lobe convex, not at all pollinose;
antennal groove pollinose, continous with pollinose orbital groove; temporal lobe with one to six rather fine punctures, or
none; temporal setae none to four; temporal setae more frequent than in related species; antennal lobe glabrous, distinctly
raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; postorbit
pollinose; postorbital tubercle small about 0.3 of length of eye.
Pronotum relatively broad, length/greatest width 1/19; widest near middle; base, apex distinctly, nearly equally
narrowed; lateral margins slightly curved except oblique near base and apex; margin shallowly sinuate just anterior to hind
angle; paramedian grooves nearly parallel; outer carina widest just anterior to middle; both inner, outer carinae coarsely
punctate.
Elytral striae broad, rather shallow; strial punctures coarse throughout; intervals convex, narrow, each narrower than
one strial puncture; Stria II with one or two setae near apex; Stria IV with one or two near apex; subapical striole with
seta; Stria VI 1 with several setae near apex; abdominal sterna moderately densely punctate; 25-29 punctures in each half
of Sternum V; male with ventral tooth on anterior femur, proximal tooth on anterior tibia; middle calcar small but distinct;
hind calcar slightly larger than middle one.
This species differs sharply from the sympatric O. ineditus in absence of pollinosity from the
temporal lobe; longer, narrower head; and much shallower emargination of head margin
anterior to the eye. It is more similar to O. hemipunctatus, O. occultus , and O. germaini. The
first species lacks punctures on the inner carina of the pronotum, the second has many coarse
punctures on the temporal lobe and a complete series of setae in Stria IV, and the third has the
pronotum inflated anteriorly and the postorbital tubercle much larger.
Range. - O. javanicus occupies the western 2/3 of Java, where it seems to be the commonest
member of the subgenus. The easternmost locality is Kopeng, near Jogjakarta. In addition to
type material we have seen the following specimens; one male, five females, Gunong Tengkoeban Prahoe,
elev. 1000-1400 m., June 20, 1933, coll, van Doesburg (LEI); one male, two females, Kopeng, elev. 1610 m.. May 14,
1933, coll, van Doesburg (LEI); one female, Preanger, no date, coll. P.F.S. Sijthoff (LEI).
Revision of Omoglymmius Ganglbauer
137
Omoglymmius ( Hemiglymmius ) germaini (Grouvelle)
Figs. 3, 1 1
Rhysodes germaini Grouvelle 1903: 109.
Omoglymmius ( Hemiglymmius ) germaini (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) female labelled: “Cochabamba, Bolivia”
(MNHN) PARALECTOTYPE male, same data (MNHN); the paralectotype has a slightly
deformed pronotum, with an irregular interruption of the lateral margin near the apex. This
species is very close to O. javanicus , and might prove to be conspecific with it. It is very likely
that the collecting data are erroneous, and that the true locality is in one of the Greater Sunda
Islands.
Vulcano and Pereira (1975b) illustrated a specimen under the name R. germaini Grouvelle
labelled as coming from Rio Madeira, Brazil. Their specimen lacks postorbital tubercles and
antennal stylets, and is not O. germaini nor a member of Subgenus Hemiglymmius. We have
not studied this specimen, and cannot form a definite opinion about its specific identity nor its
subgeneric placement. We hope that more specimens will be collected. If it is not a mislabelled
specimen from some other part of the world, it represents the first authentic record of Genus
Omoglymmius from South America. In the paper of Vulcano and Pereira two figures were
transposed, so that illustrations of their Omoglymmius appear under the name Clinidium
rojasi Chevrolat.
Description. — Length 7 .0-7 .2 mm. Basal setae well-developed on antennal Segments V-X; Segments I- 1 V with
pollinose band on dorsal surface. Segments V-X coarsely punctate; Segment XI longer than wide, corpus cylindrical,
narrower than Segment X; stylet short, acute.
Head slightly longer than wide; median lobe impunctate, glabrous, elongate oval, its tip bluntly rounded near level of
posterior margin of eye; frontal space 1.5 wider than long; medial angles slightly obtuse, contiguous; temporal lobe convex,
not at all pollinose; antennal groove pollinose, continuous with pollinose orbital groove; temporal lobe with one to six coarse
punctures; temporal seta absent; antennal lobe glabrous, distinctly raised above level of temporal lobe; lateral margin of
head shallowly sinuate between antennal lobe and eye; postorbit finely pollinose; postorbital tubercle 0.3-0. 5 of length of
eye.
Pronotum with length/greatest width 1.21-1.24, widest anterior to middle; base, apex equally narrowed; lateral
margins more markedly curved than in O. javanicus ; margin shallowly sinuate just anterior to hing angle; paramedian
grooves distinctly convergent anteriorly; outer carina broadest at anterior 0.25 of length; both inner, outer carinae coarsely
punctate.
Elytral striae broad, rather shallow; strial punctures coarse throughout; intervals convex, narrow, each narrower than
one strial puncture; Stria II with two setae near apex; Stria IV with two setae near apex; subapical stride with one seta;
Stria VII with several setae near apex; abdominal sterna densely punctate with about 33 punctures in each half of Sternum
V; male with ventral tooth on anterior femur, proximal tooth on anterior tibia; middle calcar more prominent than in O.
javanicus , longer than hind calcar, similar to that of O. javanicus.
We regard this species as only doubtfully distinct from O. javanicus , distinguished by the
pronotum more inflated anteriorly and perhaps by the larger middle calcar.
Omoglymmius (Hemiglymmius) hemipunctatus new species
Figs. 4, 12
Type material. - HOLOTYPE female, labelled: “Corporaal, Serdang, Lau Raku, 4-1910,
cat. no. 6” (LEI).
This locality is ambiguous, but probably refers to the former Province of Serdang in
Sumatra, and not the town by the same name in northwestern Java.
Description. — Length 7.0 mm approx, (specimen broken). Basal setae well-developed on antennal Segments V-X;
Segments I- 1 V each with pollinose band on dorsal surface; Segments II-XI coarsely punctate; Segment XI distinctly longer
than wide, the corpus nearly cylindrical, slightly narrower than Segment X; stylet acute, longer than in O. germaini.
Quaest. Ent., 1982, 18 (1-4)
138
Bell and Bell
Head as long as wide; median lobe glabrous, elongate-oval, its tip bluntly rounded just posterior to middle of eye;
median lobe impunctate; frontal space 1.5 wider than long; medial angles slightly obtuse, narrowly separated; temporal
lobe convex, not at all pollinose, but with pollinose strip in antennal groove and extended into orbital groove; temporal
lobe with 16-18 coarse punctures, but without temporal setae; antennal lobe glabrous, distinctly raised above level of
temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; postorbit pollinose; postorbital
tubercle small, about 0.4 of width of eye.
Pronotum slightly narrower than in O. germaini. length-greatest width 1.23;widest near middle, apex more
narrowed than base; lateral margin distinctly sinuate anterior to hind angle; pronotal carinae convex, nearly equal; inner
carina impunctate, outer carina coarsely punctate.
Elytral striae broad, rather shallow; strial punctures coarse in anterior 80%, becoming distinctly finer near apex;
Stria II with seta at apex; Stria IV with three setae near apex; apical stride with one seta; several setae near apex of
Stria VII; abdominal sterna densely punctate, with about 33 punctures in left half of Sternum V.
Male unknown.
This species is close to O. javanicus, but differs in having the temporal lobes strongly
punctate and the inner pronotal carina entirely impunctate.
Omoglymmius (Hemiglymmius) occultus new species
Figs. 5, 13
Type material. - HOLOTYPE male, labelled: “SUMATRA” (MNHN); PARATYPE
female, same data (MNHN).
Description. — Length 6. 4-7. 7 mm. Basal setae well-developed on antennal Segments V-X; Segments 1-IV each
with pollinose band on dorsal surface; Segments IV-XI coarsely punctate; Segment XI distinctly longer than wide, corpus
slightly narrowed near base, only slightly narrower than Segment X; stylet short, acute.
Head as wide as long; median lobe impunctate, glabrous except for minute pollinosity on clypeus; median lobe elongate
oval, its tip obtuse, even with middle of eye; median space slightly wider than long; medial angles obtuse, nearly
contiguous; temporal lobes convex, not pollinose, but with pollinose strip in antennal groove and extended along orbital
groove; temporal lobe with approximately 15 coarse punctures; temporal setae absent; antennal lobe glabrous, distinctly
raised above level of temporal lobe; lateral margin of head shallowly sinuate between antennal lobe and eye; post-orbit
pollinose; postorbital tubercle larger than in O. javanicus, its width 0.5 of width of eye.
Pronotum relatively narrow, length/greatest width 1.30; widest near middle; apex more markedly narrowed than base;
margin distinctly sinuate anterior to hind angle; pronotal carinae convex, nearly equal in width; inner carina with six to 12
scattered coarse punctures, most of them near base; outer carina densely, coarsely punctate.
Elytral striae broad, rather shallow; strial punctures coarse throughout, not becoming finer near apex; Stria 11 with
one to two setae near apex; Stria IV with five or six setae scattered along its length; apical striole with one seta; several
setae near apex of Stria VII; abdominal sterna coarsely, densely punctate; male with ventral tooth present on anterior
femur, with proximal tooth present on anterior tibia; middle calcar small but distinct; hind calcar equal to middle one.
This species is unique among Indonesian Hemiglymmius in having a complete series of setae
in Stria IV. It also differs from O. hemipunctatus in having punctures on the inner pronotal
carina, and from O. javanicus and O. germaini in having more punctures on the temporal lobe
and in having a larger postorbital tubercle.
Omoglymmius ( Hemiglymmius ) ineditus (Dajoz)
Figs. 6, 14
Rhysodes ineditus Dajoz 1975: 8-9.
Omoglymmius ( Hemiglymmius ) ineditus (Dajoz) Bell and Bell 1978
Type material. - HOLOTYPE sex not listed, labelled: “JAVA:Tji Salimar, Province de
Preang dans le Sud-Ouest de File, altitude 3,000 pieds, septembre 1897” (MNHN).
We have not been able to study this type, which has apparently been misplaced. We
provisionally conclude, on the basis of the description and figure, that it applies to the species
described below.
Revision of Omoglymmius Ganglbauer
139
Description. — Length 5.8-7. 1 mm. Basal setae well-developed on antennal Segments V-X; Segments 1-1 V each
with pollinose dorsal band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly
cylindrical, narrower than Segment X; stylet short, acute.
Head slightly wider than long; greatest width/length 1.000-1.075; mean 1.050; median lobe pollinose, impunctate,
narrow, margins nearly parallel, tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long; medial
angles obtuse, contiguous; temporal lobe convex, anterior half pollinose, posterior half glabrous, shining, with one to four
fine punctures or none; temporal seta absent; antennal lobe pollinose, continuous with anterior part of temporal lobe,
without trace of scarp; lateral margin of head deeply sinuate between antennal lobe and eye, postantennal groove oblique;
postorbit pollinose; postorbital tubercle very small, less distinct than in O. rimatus.
Pronotum rather narrow; length/greatest width 1.26, widest near middle, apex strongly narrowed, base moderately
narrowed; margin scarcely sinuate anterior to hind angle; pronotal carinae convex, outer and inner ones nearly equal in
width at middle; outer carina with 14-18 punctures; inner carina with one to eight coarse punctures or none.
Elytral striae broad, rather shallow, intervals convex, narrow; each interval narrower than one strial puncture; latter
coarse, closely spaced; Stria 11 with one or two setae near apex; Stria IV with single seta at apex; subapical striole with one
seta; Stria VI 1 with several setae near apex; abdominal sterna densely punctate; each half of Sternum V with 38-48
punctures; male with ventral tooth on anterior femur and proximal tooth on anterior tibia; middle calcar triangular, about
0.5 as long as spur; hind calcar triangular, slightly larger than middle one.
This species is marked by the complete absence of posterior scarp of the antennal lobe. The
latter connects to the narrow, convex anterior portion of the temporal lobe, forming an angular
pollinose ridge. The median lobe is pollinose. O. rimatus is similar, but has a glabrous median
lobe and a distinct scarp posterior to the antennal lobe. O. inermis is intermediate in having an
indistinct scarp posterior to the antennal lobe. It has the median lobe pollinose, as in the present
species, but differs from both O. ineditus and O. rimatus in the male lacking the proximal tooth
on the anterior tibia.
Range. - Western Borneo, western Java, and eastern Sumatra. We have studied specimens
from the following localities: BORNEO; one female, Mowong, West Borneo, coll. F. Muir, Sept. 1907 (BMNH);
JAVA: one male, Pangalengen, elev. 4000 feet, 1893, coll. H. Fruhstorfer (MNHN); one male, Java-Orient (MNHN);
SARAWAK: one female, Mount Matang, West Sarawak, coll. G. K. Bryant, January, 1914 (BMNH); SUMATRA: ten
males, four females, Palembang (MNHN); two males, one female, Palembang (BMNH); one female, Palembang (AMS).
Variation. - The two specimens from Borneo lack punctures of the inner carina, and might
not be conspecific with the others, which have four to eight such punctures.
Omoglymmius (Hemiglymmius) rimatus new species
Figs. 7, 15
Type material. - HOLOTYPE male, labelled: “JAVA, Mt. Kawi” (MNHN).
PARATYPES two females (one mounted on same pin as holotype), same label as holotype
(MNHN); one female labelled: “Mt. Kawi, Java, A. Grouvelle” (LEI); two females, labelled:
“Mt. Kawi, Java, Juli 1934, v. Doesburg” (LEI).
Description. — Length 6. 2-8. 8 mm. Basal setae of antenna well-developed on Segments IV-X; Segents I- 1 V each
with dorsal pollinose band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly
cylindrical, narrower than Segment X; stylet short, acute.
Head as long as wide; median lobe glabrous, impunctate or with a few coarse punctures; median lobe narrow, margins
nearly parallel, tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long; medial angles obtuse,
contiguous or slightly separated; temporal lobe convex; its anterior 0.33 pollinose; posterior portion glabrous, shining with
few punctures; temporal seta absent in most specimens, a few specimens with small temporal seta; antennal lobe glabrous,
raised above level of temporal lobe and separated from it by distinct scarp; pollinosity of temporal lobe in most specimens
ended posteriorly, opposite middle of eye; lateral margin of head deeply sinuate between antennal lobe and eye,
postantennal groove oblique; postorbit pollinose; postorbital tubercle about 0.3 as long as eye.
Pronotum with length/greatest width about 1.23; widest near middle; apex strongly narrowed, base moderately
narrower; margin scarcely sinuate anterior to hind angle; pronotal carinae convex, outer and inner ones nearly equal in
width at middle; outer carina with 20-25 punctures; inner carina with 11-14 punctures.
Elytral striae broad, rather shallow; intervals convex, narrow, each narrower than one strial puncture; strial punctures
coarse, close together; Stria 11 with single seta at apex; Stria IV with two setae at apex; subapical striole with single seta;
Stria VII with several setae near apex; abdominal sterna moderately densely punctate: each half of Sternum V with 27-37
Quaest. Ent., 1982, 18 (1-4)
140
Bell and Bell
punctures; male with ventral tooth on anterior femur and proximal tooth on anterior tibia; middle calcar acute, about
O. 5 as long as spur; hind calcar triangular, its proximal margin concave, smaller than middle one.
This species has a distinct postantennal groove and extensive pollinosity on the temporal
lobe as in O. ineditus , but differs sharply in having the median lobe glabrous and the antennal
lobe glabrous, strongly elevated, and separated from the temporal lobe by a distinct scarp.
Range. - West Central Sumatra and eastern Java. This species appears to be allopatric to O.
ineditus, and to occupy a markedly disjunct range. In addition to the type series, we have seen
specimens from the following localities: JAVA; five males, five females, Gg. Moeria, Tjolo, elev. 700-1000 m.,
P. H. v. Doesburg (LEI); one male, two females, Mt. Moeria, 3-4000' (MNHN); three males, two females (on same pin),
Malang (MNHN); SUMATRA: two males, one female, Pajakombo (MNHN).
Variation. - The series of specimens from Gunong Moeria have the pollinosity of the
temporal lobe less extensive than in specimens from other localities.
Omoglymmius ( Hemiglymmius ) inermis new species
Figs. 8,16
Type material. - HOLOTYPE male, labelled: “SUMATRA” (MNHN). PARATYPE
male, labelled: “SUMATRA, don. Grouvelle 1901” (GEN).
Description. — Length 7.0 mm. Basal setae well-developed on antennal Segments V-X; Segments I- 1 V each with
pollinose dorsal band; Segments V-X coarsely punctate; Segment XI distinctly longer than wide, corpus nearly cylindrical,
narrower than Segment X; stylet short, acute.
Head slightly wider than long, slightly more transverse than in preceding species, median lobe pollinose, impunctate,
narrow, its sides nearly parallel, its tip bluntly rounded at level of middle of eye; frontal space 1.5 wider than long;
temporal lobe convex, its anterior half pollinose, a rather broad band of pollinosity extended posteriorly medial to eye;
medial angles obtuse, contiguous; posterior half of temporal lobe otherwise glabrous, shining with few fine punctures;
temporal seta absent; antennal lobe pollinose except for narrow area along lateral margin; antennal lobe continuous with
temporal lobe, not marked off by a well-defined scarp, though fine trace of scarp is present; lateral margin of head
distinctly sinuate between antennal lobe and eye; postorbit pollinose; postorbital tubercle small.
Pronotum more elongate than in O. ineditus , length/greatest width 1.36; widest near middle, apex strongly narrowed,
base more narrowed than in O. ineditus , lateral margin not sinuate anterior to hind angle; median groove narrow; pronotal
carinae convex, nearly equal, punctate; inner carina more sparsely punctate than outer one.
Elytral striae broad, rather shallow; intervals convex, narrow; each interval narrower than a strial puncture; strial
punctures coarse, closely spaced; two setae in apex of Stria II; one in apex of Stria IV; one in apical stride; several in apex
of Stria VII; abdominal sterna coarsely punctate; 41 coarse punctures in left half of Sternum V, additional very minute
punctures present on sterna; male with ventral tooth on anterior femur, but without proximal tooth on anterior tibia;
calcars. very small.
This species is very similar to the two preceding ones, from which it differs in the absence of
the proximal tooth on the anterior tibia of the male. It is also larger than the males of the
sympatric O. ineditus, and in addition, has a more elongate pronotum and smaller calcars. The
female is unknown.
Subgenus Boreoglymmius new subgenus
Type species. - Rhysodes americanus Castelnau 1836
Description. — Basal setae on antennal Segments V-X, antennal stylet absent; outer segments not at all punctate;
Segment I finely pollinose, others glabrous; clypeal setae absent; eye large, normal, deeper than long; cornea facetted;
temporal setae absent; frontal grooves deep, complete; medial angle simple, no translucent area; postorbital tubercle
present or absent; marginal groove of pronotum deep; spur of middle tibia curved anteriorly; punctures of abdominal
Sterna III-V scattered; lateral pits of female varying, either limited to abdominal Sternum IV and shallow, or else deep
and extended over Sterna III and IV; anterior femur of male with ventral tooth; anterior tibia of male without proximal
tooth; hind calcars of male larger than middle ones.
This subgenus resembles Hemiglymmius in most respects, but differs sharply in lacking the
antennal stylet. It is also close to Omoglymmius s. str. from which it differs in presence of basal
Revision of Omoglymmius Ganglbauer
141
setae on antennal segments.
Three allopatric species are known, two from North America, and one from Japan.
Phytogeny. - O. hamatus is structurally as well as geographically intermediate between the
two other species. It resembles O. americanus in the presence of a small, shallow lateral pit
restricted to Sternum IV in the female, and in the absence of a postorbital tubercle. O. lewisi, in
contrast, has a deep lateral pit extended over Sterna 111 and IV, and has a well-developed
postorbital tubercle. On the other hand, O. hamatus and O. lewisi are very similar in the shape
of the temporal lobe and of the hind calcar.
Two possible phylogenies seem worthy of consideration: either O. lewisi and O. hamatus
share an ancestor, and O. americanus is more distantly related, or else O. hamatus and O.
americanus are more closely related than either is to O. lewisi. We believe that the latter
phylogeny is the more likely one. The presence of a postorbital tubercle in O. lewisi is a
similarity to Hemiglymmius. According to this hypothesis, the ancestral species had most of
the characters of O. hamatus except that the postorbital tubercle was present. Perhaps this
species evolved in eastern Asia. Later it invaded western North America, and still later it split
into two daughter species after the range was interrupted at the Bering Strait. The population
that remained in Asia developed enlarged lateral pits on the abdominal sterna of the female,
and became O. lewisi. The population in western North America lost the postorbital tubercle.
This species later split into two: O. hamatus , essentially like the ancestral North American
species, and O. americanus , in eastern North America, which evolved a shorter, wider head,
and which, in the male, had the hind calcars secondarily simplified. According to this view, the
similarites between O. americanus and O. (s. str.j germari of Europe represent evolutionary
convergence.
KEY TO SPECIES
1 Postorbital tubercle present; female with deep lateral pit, extending over
abdominal Sterna III and IV O. lewisi (Nakane), p. 141
Y Postorbital tubercle absent; lateral pit of female confined to Sternum IV, small
and shallow 2
2 (L) Frontal space twice as wide as long; medial angles of temporal lobe narrow,
obtuse, nearly contiguous; posteriomedial margin distinctly concave between
medial and occipital angles O. americanus (Castelnau), p. 144
2' Frontal space at most slightly wider than long; medial angles broadly rounded;
posteriomedial margin not emarginate O. hamatus (Leconte), p. 143
Omoglymmius (Boreoglymmius) lewisi (Nakane) NEW COMBINATION
Figs. 17, 20,21,24
Rhysodes lewisi Nakane 1973: 5 (described in Japanese and designated by the author as a
manuscript name.)
Omoglymmius (Hemiglymmius) lewisi (Nakane) Bell and Bell 1978
Omoglymmius lewisi Nakane 1978: 129-130 (described in English)
Type material. - HOLOTYPE male, Kirishima, Kyushu, Japan, 26- VII- 1 97 1 , T. Nakane
lgt. ALLOTYPE female, Kurama, Kyoto, Honshu, Japan, 22-IV-1952, F. Takahashi lgt.
Quaest. Ent., 1982, 18 (1-4)
142 Bell and Bell
Plate 2. Figures 17-26, subgenus Boreoglymmius. Figs. 17-19, Head and pronotum, dorsal aspect; Fig. 17, Omoglymmius
( Boreoglymmius ) lewisi (Nakane); Fig. 18, O. (B.) hamatus (Leconte); Fig. 19, O. (B.) americanus (Castelnau); Fig. 20,
Metasternum and abdomen, ventral aspect, O. (B.) lewisi (Nakane); Figs. 21-23, Head, lateral aspect; Fig. 21, O. (B.)
lewisi (Nakane); Fig. 22, O. (B.) hamatus (Leconte); Fig. 23, O. (B.) americanus (Castelnau); Figs. 24-26, Hind tibia,
male, apical portion; Fig. 24, O. (B.) lewisi (Nakane); Fig. 25, O. (B.) hamatus (Leconte); Fig. 26, O. (B.) americanus
(Castelnau).
PARATYPES one male, one female, Fujimidai, Shinano, Honshu, Japan, 1 1 -VI- 1 944, S.
Osawa lgt.; one male, Kurama, Kyoto, Honshu, Japan, 1 1 -VII- 1956, T. Nakane lgt.; one male,
Mt. Kooya, Wakayama, Honshu, Japan, 1 1 - VII- 1 948. M. Hayashi lgt. (all KU).
We have not seen the holotype, but have studied excellent drawings kindly furnished to us
by Dr. Nakane.
Description. — Length 6. 5-8.0 mm. Antennal Segment I coarsely punctate; head distinctly longer than wide;
clypeus with few punctures near margin, continuous with median lobe; latter rhomboidal, narrow, pointed posteriorly even
with anterior third of eye; temporal lobe coarsely, very sparsely punctate, punctures very scarce in medial half;
anteriomedial margin oblique, scarcely sinuate; medial angles obtusely rounded, contiguous; frontal space longer than
wide; posteriomedial margin slightly convexly curved; occipital angle distinct; posteriolateral margin oblique between
occipital angle and eye; orbital groove with pollinose strip narrowed posteriorly, extended about to middle of eye;
postorbital tubercle with well-marked dorsal margin, but with ventral margin indistinct; postorbit with few coarse
Revision of Omoglymmius Ganglbauer
143
punctures near eye, not pollinose.
Pronotum moderately elongate, length/greatest width 1.2; widest near middle, narrowed both anteriorly and
posteriorly, margin sinuate anterior to hind angle; median and paramedian grooves relatively narrow, punctate; carinae
broad; marginal groove fine, linear; outer carina punctate throughout; inner carina with few punctures near base.
Elytra long, narrow, intervals narrow, slightly convex; striae relatively shallow; punctures fine; Stria IV with one or
two setae near apex; apical stride with two setae; several setae near apex of Stria VII; male with middle calcar acute;
hind calcar large, almost twice as wide as tip of tibia, its tip acute, its dorsal side with pronounced “shoulder” with
group of large setae (Fig. 24); lateral abdominal pit of female, deep, long, narrow, extended over Sterna Ill-IV (Fig.
20).
This species differs from other members of the subgenus in presence of a postorbital tubercle
and in form of the lateral pit of the female.
Range. - Restricted to Japan. We have studied the following specimens: one male, one female,
Mt. Kooya (Wakayama Prefecture), VII-1 1-1948, coll. Masao Hayasi (SATO); one female, Mie University Forest,
Ichishi-Gun (Mie Prefecture), 24-VI-1956, coll. M. Sato (SATO). In addition, we have seen one male labelled: “JAPAN:
G. Lewis, 1910-320, C. F. Baker Colin. 1927” (NMNH). This specimen is labelled as a “cotype” of
Rhysodes crassiusculus Lewis.. This can be a true cotype if the date on the label represents the
acquisition date and not the collection date. The description of R. crassiusculus Lewis seems to
be a composite, as the description of the head seems to be based on the present species, while
the secondary sexual characters are clearly those of the Japanese species of subgenus
Pyxiglymmius. We follow Nakane in restricting the name R. crassiusculus to the latter
species.
Omoglymmius ( Boreoglymmius ) hamatus (Leconte) NEW COMBINATION
Figs. 18,22, 25
Rhysodes aratus Chevrolat (nec Newman) 1873a: 209.
Rhysodes hamatus Leconte 1875: 163.
Type material. - HOLOTYPE, sex not recorded, labelled: “CALIFORNIA” (MCZ,
6830).
Description. — Length 6. 2-8. 8 mm. Antennal Segment I coarsely punctate in most specimens (few specimens with
reduced punctation), finely pollinose; head longer than wide; clypeus with few punctures near margin; clypeus continuous
with median lobe; latter rhomboidal, narrow, pointed posteriorly, tip even with anterior third of eye; temporal lobe
coarsely, rather sparsely punctate; anteriomedial margin oblique, only shallowly sinuate; medial angles obtusely rounded,
contiguous or narrowly separated; frontal space about as wide as long; posteriomedial margin convexly curved; occipital
angle obtusely rounded; posteriolateral margin oblique to eye; orbital groove short, narrow, its posterior end opposite
middle of eye; postorbit convex, but not in form of distinct tubercle; postorbit finely pollinose.
Pronotum longer and narrower than in related species, length/greatest width 1.33 or greater; both base and apex
distinctly narrowed; lateral margin not sinuate anterior to hind angle, both inner and outer carinae sparsely, rather
coarsely punctate.
Elytra long, parallel-sided; striae rather deep, coarsely punctate; each puncture about as wide as an interval; Stria IV
with one to three setae near apex; apical striole with one seta; Stria VII with several setae near apex; male with hind calcar
much larger than middle calcar, with pronounced “shoulder” on dorsal surface, separated from shaft of tibia by deep
emargination (Fig. 25); female with lateral pit of abdomen on Sternum IV, shallow, ill-defined.
This species is distinguished from O. lewisi by absence of the postorbital tubercle, and form
of the hind calcar of the male and of the lateral abdominal pit of the female. Compared to O.
americanus , this species is much narrower and more elongate, and with better developed
punctation. Available bionomic information was summarized by Bell (1970).
Range. — Widespread in forested regions of California and Oregon. In California, represented by many specimens
from the Sierra Nevada and in the mountains north of Los Angeles, as well as by a few records from the Coast Range
north of San Fancisco. In Oregon, known from many localities in the Cascade Range, and from isolated mountain ranges
in southcentral and northeastern parts of the state, but not recorded from the Coast Range. There are also records from
several localities in the northern third of Idaho. There is a single record from southeastern Washington, and another, quite
isolated one, from the Huachuca Mountains of southeastern Arizona. There are also records without specific localities
from British Columbia, Nevada, and Texas.
Quaest. Ent., 1982, 18 (1-4)
144
Bell and Bell
We have seen specimens from the following localities: ARIZONA: Huachuca Mountains (Cochise County)
(CNHM).
CALIFORNIA: Ahwahnee (CNHM); Alhambra (CAS); Areata (LA); Arrowhead (LA, CAG); Bartlett Springs (CAS);
Barton Flats (San Bernardino Co.) (UCD, OS, CAG); Bass Lake (LA, CAS); Bear Valley (Santa Barbara Co.)(CAS);
Big Bear Lake (San Bernardino Co.), 6700 ft. (LA); Big Meadows (UCD); Big Tree Grove (MCZ); Big Trees (Calaveras
Co.) (UK, CU, AP, CAS); Blodgett Forest (El Dorado Co.), 4400 ft. (LA); Buck’s Lake (Plumas Co.) (UCD); Butte
Co. (CNHM); Calaveras Co. (CU); Calistoga (CAS); Camino (CAG); Camp Connell (Calaveras Co.) (CAG); Carrville
(Trinity Co.), 2500 ft. (LA, CAS); Cascade Lake (El Dorado Co.) (BSR1); Castle Crag (CMP, MCZ); Cedar Pass
(Modoc Co.) (LA, UCD); China Flats (El Dorado Co.) (CAS; Chester (CNHM); Chiquito Creek (Madera Co.), 4100 ft.
(CU, UCD); Cisco (MNHB); Cole (CAS); Coleville (CNHM); Coffey Creek (Trinity Co.) (UCD); Dalton Creek (Fresno
Co.), 4800 ft. (CAS, CU); Dardanelle (Tuolumne Co.) (LA); Dark Creek (San Jacinto Mts.) (LA); East Crag (CNHM);
Desolation Valley, Echo Lake (El Dorado Co.), 7500 ft. (CU, BSRI); El Dorado Co. (CNHM); Facht (Lassen Co.)
(CAS); Fallen Leaf Lake (El Dorado Co.) (CAS); Feather Falls (Butte Co.) (LA); Fish Camp (MCZ); Fresno Flats,
3500 ft. (ISNHS); Georgetown (CAG); Giant Forest, 6400 ft. (CU); Grass Valley (LA); Hackamore (OSFS); Happy
Camp (MCZ); Hat Creek P.O. (Shasta Co.) (LA); Hassock Meadows (Tulare Co.) (CAS); Hayfork (CAS); Hobart
Mills (CAG); Huntington Lake (UCD); Ice House (El Dorado Co.) (CAG); Kaweah (CAS); Kyburz (El Dorado Co.)
(UCD); Lake Almanor (Plumas Co.) (CAG); Lake Tahoe, 6465 ft. (LA, PA); Leland Meadows (Tuolumne Co.) (UCD);
Madera Co. (CAS); Manzanita Creek (Lassen Co.) (UCD); Mariposa Co. (CNHM); Mather (Tuolumne Co.) (OSU,
UCD, WRS); Meadow Creek (Plumas Co.) (CAS); Mendocino Co. (CNHM): Miami (Madera Co.) (CU, AP, LA);
Mineral (UCD); Mokelumne (CAS); Morgan Summit (Tehama Co.) (CAG); Mount Aukum (El Dorado Co.) (LA);
Mount Home (Tulare Co.) (UCD); Northfork (CU); Old Station (Shasta Co.) (LA); Paradise Valley (King’s River),
7000 ft. (CAS); Pasadena (MO); Peanut (Trinity Co.) (LA); Pinecrest (Tuolumne Co.) (UT); Placer Co. (MO, CNHM,
PA); Plumas Co. (CNHM); Pollock Pines (LA); Pyramid Ranger Station (OSFS); Rainbow Falls (Madera Co.) (UCD);
Red’s Meadow (Madera Co.) (LA); Riverton (El Dorado Co.) (CAG); San Bernardino Mts. (MCZ); Sequoia National
Park (CAS, CAG); Shasta Co. (CNHM); Shasta Retreat (Siskiyou Co.) (CAS); Sierraville (CAG); Sissons (Siskiyou
Co.) (MO, CNHM, MCZ); Strawberry (El Dorado Co.) (CAS, LA); Sugar Pine (CAS); Tahquitz Valley (San Jacinto
Mts.) (LA); Tallac (CAS, SD); Trinity Co. (CNHM); Truckee (CAS); Tulare Co. (CNHM, MCZ); Tuolumne Co.
(CNHM); Uncle Tom’s (CAG); Viola (LA, CAS); Virner (CAG); Walker Mine (OSFS); Weaverville (CAS); West Fork
(San Gabriel Canyon) (UT); Whitehall (El Dorado Co.) (LA); Yosemite (UK, LA, CAS); Yuba pass (Sierra Co.) (LA,
CAG).
CANADA: “B.C.” (CMP).
IDAHO: Clearwater (Idaho Co.) (DY); Coeur d’ Alene (CAS); Harvard (Latah Co.) (WS); Moscow (WS); Moscow Mt.
(WS).
NEVADA: “Nevada” (MNHB).
OREGON: Beaver-Sulfur (Jackson Co.) (OS); Brownsboro (OS); Butte Falls (Jackson Co.) (UCS, OS); Cave State Park,
5 mi. nw of Culver, 1775 ft. (UT); Colton (CAS); Deming Creek (near Bly) (MSU); Fish Lake (Jackson Co.) (UCD);
Huckleberry Mt. (Jackson Co.), 5500 ft. (CNHM); Indian Fork Creek (near Sisters) (UT, OS); Josephine Co. (CNHM);
Klamath Co. (CNHM, OS); Klamath Falls (MSU); Lake of the Woods (Klamath Co.), 5300 ft. (CNHM); Mahama
(Marion Co.) (LA); McAllister Soda Springs (Jackson Co.) (CNHM); McDonald Forest (Benton Co.) (OS); Moon
Prairie (OS); Portland (CAS); Ruch (Jackson Co.) (LA); Union (CNHM); Union Creek (Jackson Co.), 3100-3500 ft.
(CNHM); Upper Klamath Lake (MUS); Wildwood Camp (Ochoco National Forest) (OS).
TEXAS: “Tex.” (MN).
WASHINGTON: Dayton(WS).
Omoglymmius (Boreoglymmius) americanus (Castelnau) NEW COMBINATION
Figs. 19, 23,26
Rhysodes americanus Castelnau 1836: 58.
Rhysodes exaratus Serville 1825: 308 (nec Dalman 1823; nec Erichson 1848).
Rhysodes aratus Newman 1838: 664.
Omoglymmius americanus (Castelnau) Bell 1975: 351.
Type material. - HOLOTYPE male, labelled: “America Sept., De Castelnau 2-43”
(MNHN).
Description. — Length 6. 0-7. 9 mm. Segment I of antenna pollinose, impunctate or with very few punctures; head as
long as wide; clypeus impunctate, continuous with median lobe; latter relatively broad, rhomboid, tip pointed opposite
anterior third of eye; temporal lobe very finely punctate, impunctate in appearance at low magnifiction; anteriomedial
margin very deeply sinuate, anterior half longitudinal, posterior half oblique; medial angles narrow, obtusely pointed,
contiguous; frontal space twice as wide as long; posteriomedial margin shallowly emarginate posterior to obtuse but
Revision of Omoglymmius Ganglbauer
145
distinct occipital angle; lateral margin slightly curved between occipital angle and eye; orbital groove relatively
well-developed, extended to posterior margin of eye; postorbit flat, pollinose, without trace of tubercle.
Pronotum relatively short and broad, length/greatest width is 1.09; widest near middle, both base and apex
narrowed; apex less narrowed than in related species; pronotal carinae with very fine punctures, impunctate in
appearance at low magnification; elytra broader and less parallel-sided than in O. hamatus-, strial punctures less coarse
than in O. hamatus , each puncture clearly narrower than an interval; Stria IV with one to four setae near apex; hind
calcar of male acute, triangular, not “shouldered” dorsally (Fig. 26); lateral abdominal pits of female as in O. hamatus.
This species differs from other members of the subgenus in having the hind calcar
triangular. It is easily distinguished from O. hamatus by broad frontal space, deeply
emarginate anteriomedial margin of the temporal lobe, emarginate posteriomedial margin, and
much finer punctures of temporal lobes and pronotal carinae. Available bionomic information
was summarized by Bell (1970).
Range. - Eastern North America, north to central New York, southern Ontario, and the
vicinity of Minneapolis, Minnesota; west to southeastern Nebraska and eastern Texas; south to
the Gulf of Mexico and northern Florida. Two specimens labelled “Arizona” (ISNHS) are
probably mislabelled, as is certain for a specimen labelled “Nederl. Indie, A. Roller, Colin.
Drescher” (AMS). A specimen labelled “Castle Crag, California” (CMP) is also very likely
mislabelled.
Since Bell (1970) we have seen specimens from the following additional localities:
ALABAMA: Auburn (BSRI); Chiat State Park (UW); Mobile (MO).
ARKANSAS: Arkadelphia (CNHM); Bluff City (Ouachita Co.) (ARK); Rosston (Nevada Co.) (ARK).
DELAWARE: “Del.” (MCZ).
FLORIDA: Alachua Co. (UM); Florida Caverns State Park (Jackson Co.) (FLA); Gainesville (CNHM).
ILLINOIS: Algonquin (ISNHS); Carbondale (SI); Crab Orchard Lake (Williamson Co.) (SI); East Dubuque (ISNHS);
Evanston (CNHM); Funk’s Grove (ISNHS); Galena (ISNHS); Harrisburg (ISNHS); Lusk Creek (Pope Co.) (SI);
Moline (MN); Murphysboro (SI); New Lennox (CNHM); Normal (ISNHS); Pine Hills Field Station (Union Co.)
(BSRI): Pulaski (ISNHS, CNHM); Urbana (CNHM), 10); Utica (UVM); White Heath (UM).
INDIANA: Bloomington (IU); Brown County State Park (IU); Crowthersville (PU, FLA); Dubois County (PU);
Indianapolis (FLA); Jennings Co. (PU); Knox Co. (PU); Kosciusko Co. (CU); Lafayette (PU); Lagrange Co. (PU); Posey
Co. (PU); Smith (CNHM); Smith Station (La Porte Co.) (CNHM); Vigo Co. (PU).
IOWA: Ames (IO); Jefferson Co. (UVM).
KANSAS: Douglas Co., 900 ft. (MCZ, UK, CU, MNHN); Riley Co. (CNHM).
KENTUCKY: Anchorage (UL).
LOUISIANA: Baton Rouge (LS); Chastine (Natchitoches Parish) (MO); Magnolia (LS); Natchitoches (MO);
Opelousas (MNHN): Ruston (LS).
MARYLAND: College Park (LA); Elk Neck State Park (UD); Sparrows Point (CNHM).
MICHIGAN: Allegan Co. (MSU); Ann Arbor (UN); Clinton Co. (HL, UVM); Detroit (MO); Frazer (MSU); Grand
Rapids (UN); Kalamazoo (MCZ); Lansing (MSU); Washtenaw Co. (UM).
MINNESOTA: Hennepin Co. (MN); Ramsay Co. (MN).
MISSISSIPPI: Avera (CU); Chocktaw Co. (RCG); Greer (UW); Lafayette Co. (BSRI); New Augusta (CU).
MISSOURI: Ashland (MO); Columbia (MO); Louisiana (MO); McBaine (MO); Meramee Highland (St. Louis Co.)
(MO); Saint Louis Co. (CMP, MO); Sikeston (MO); Warrensburg (CNHM).
NEBRASKA: Ashland (UN); Lincoln (UN).
NEW JERSEY: “N.J.” (MCZ).
NEW YORK: Buffalo (CNHM); Lodus (MCZ); McLean (Tompkins Co.) (CU); McLean Bogs (Cortland Co.) (CU);
Niagara (CNHM).
NORTH CAROLINA: Southern Pines (UI, NMNH).
OHIO: Bowling Green (RCG); Cedar Swamp (Champaign Co.) (FLA); Columbus (MSU); Dayton (DM); Delaware Co.
(WS); Franklin Co. (NC); Montgomery Co. (DM); Oxford (NC); Wood Co. (RCG).
OKLAHOMA: Bradem (OK).
ONTARIO: Essex Co. (KS, RCG); Kingsville (CNHM); Port Alma (WR); Tilbury (UVM, KS).
PENNSYLVANIA: Allegheny (CMP); Harrisburg (AP); Jeanette (CMP); New Cumberland (AP); Norwich (MNHB);
Pittsburgh (CMP).
SOUTH CAROLINA: Clemson (UVM); Florence (SDA); Sumter (UVM); Wedgefield (UVM).
TENNESSEE: Cookeville (TB); Madison Co. (CU, MCZ); Shelby Co. (DY).
TEXAS: Nacogdoches (AU).
VIRGINIA: Blacksburg (VP, NMNH); Great Falls (NMNH); Mount Vernon (NMNH); Warrenton (UK).
WISCONSIN: Broadhead (Green Co.) (UW).
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Bell and Bell
Subgenus Pyxiglymmius Bell and Bell 1978 NEW STATUS
Type species. - Rhysodes strabus Newman 1838.
Description. — Eleventh antennal segment without true stylet, its apex in most species obtuse (apex acute and
resembling a stylet in O. cristatus)\ basal setae on at least antennal Segments VIII-X (on VI-X in most species); clypeal
setae absent; frontal grooves deep, complete; medial margin of temporal lobe with two medial angles well separated from
one another, emargination between them deep except in O. lederi-, medial margin without distinct translucent area (in
some specimens of O. strabus, slight suggestion of translucent area); eye large, deeper than long; cornea facetted;
postorbital tubercle present; gular tubercle prominent except in O. crassiusculus ; punctures of abdominal sterna scattered;
female with prominent lateral pit in Sternum IV; in most species pit complex, with anteriolateral ridge or brace, and with
longitudinal striation; spur of middle tibia curved anteriorly at apex; male with proximal tooth on anterior tibia and ventral
tooth on anterior femur; calcars large, conspicuous, that of middle tibia slender, nearly cylindrical, tibial spur apparently
inserted in its base; hind calcar very large, compressed, triangular, separated from spur by distinct emargination; hind tibia
of male bent near middle, swollen proximad to middle.
In this subgenus there are two medial angles on each temporal lobe. This feature separates it
from all other subgenera except for some species of Laminoglymmius. Members of the latter
subgenus have the medial angles much closer together and, in addition, all species except O.
actae have a conspicuous translucent area in the region of the medial angles.
This taxon was ranked as a genus in Part I. Further study has convinced us that it should be
reduced to subgeneric status. O. rugosus and O. insularis have been removed to
Laminoglymmius New Subgenus.
The range of Pyxiglymmius extends through most of the Oriental Region, from Japan and
Viet Nam to Java, Sumatra, Borneo, and the Andaman and Mentawei Islands. It is also
represented on Luzon in the Philippine Islands.
Phylogeny. - A phylogenetic tree for the subgenus is presented in Diagram 1. We postulate
that the common ancestor of the subgenus, Species 1, had the following characters: median lobe
longer than wide, acutely pointed; medial margin of temporal lobe deeply emarginate; female
with lateral groove on metasternum; lateral abdominal pits of female with small anteriolateral
brace, but without striations.
Six of the nine species clearly form a monophyletic group, descended from Species 3. In the
latter species, the median lobe became markedly transverse, a specialization not seen in any of
the other subgenera of Omoglymmius. In the six species of this group, the postorbit is flat, and
is bounded ventrally by a distinct suborbital tubercle.
The three remaining species, crassiusculus, subcaviceps , and lederi, agree in having the
median lobe elongate (as is usual in Omoglymmius s. lat .), and in having the postorbit strongly
convex and not bounded ventrally by a suborbital tubercle. They may have arisen from a
common ancestor (Species 2), although none of the common characters is clearly a
synapomorphy. It is also possible that they descended separately from Species 1. O.
crassiusculus and O. lederi lack the lateral groove on the metasternum of the female. We
believe this is a secondary loss. It might be a synapomorphy. O. subcaviceps has a
well-developed lateral metasternal groove as in the remainder of the subgenus.
Of the line descended from Species 3, O. cristatus of Luzon possibly stands apart from the
others. The pointed apical antennal segment and reduced metasternal grooves are clearly
specializations, while the absence of striations from the lateral abdominal pit may represent a
character state more primitive than that in the remaining species.
The remaining species were descended from Species 4, in which deep lateral grooves were
retained on the metasternum of the female and the lateral abdominal pits had become
longitudinally striate. We hypothesize two daughter species to Species 4. One of these. Species
5, greatly increased the number of setae on the elytra, giving rise to O. pilosus and O. krikkeni.
Revision of Omoglymmius Ganglbauer
147
KRIKKENI
PILOSUS
HESPERUS
ARMATUS
STRABUS
CRISTATUS
LEDERI
CRASSIUSCULUS
SUBCAVICEPS
Phylogenetic Diagram 1. Reconstructed Phylogeny of species of Pyxiglymmius.
On the other hand, it is possible that supernumerary setae developed independently, correlated
with large size (cf. O. ( Laminoglymmius ) rugosus ). In Species 6, the number of setae on the
elytron did not increase. It gave rise to O. strabus, O. armatus, and the aberrant O. hesperus.
O. strabus has a wide range, including Sumatra, Borneo, the Malaysian Peninsula and Java.
The two remaing species appear to represent offshoots of O. strabus which have speciated on
isolated islands (the Andamans and Nicobars in the case of O. armatus and the Mentawei
Islands for O. hesperus).
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Bell and Bell
Plate 3. Figures 27-40, Subgenus Pyxiglymmius. Figs. 27-32, Head and pronotum, dorsal aspect; Fig. 27, Omoglymmius
(Pyxiglymmius) subcaviceps (Grouvelle); Fig. 28, O. (P.) crassiusculus (Lewis); Fig. 29, O. (P.) lederi (Lewis); Fig. 30, O.
(P.) cristatus new species; Fig. 31,0. (P.) strabus (Newman); Fig. 32, O. (P.) armatus (Arrow); Figs. 33-38, Head, lateral
aspect; Fig. 33, O. (P.) subcaviceps (Grouvelle); Fig. 34, O. (P.) crassiusculus (Lewis); Fig. 35, O. (P.) lederi (Lewis); Fig.
36, O. (P.) cristatus new species; Fig. 37, O. (P.) strabus (Newman); Fig. 38, O. (P.) armatus (Arrow); Fig. 39, Hind tibia,
male; O. (P.) subcaviceps (Grouvelle); Fig. 40, Antennal Segment XI, O. (P.) cristatus, new species;
Revision of Omoglymmius Ganglbauer
149
Plate 4. Figures 41-56, Subgenus Pyxiglymmius. Figs. 41-43, head and pronotum, dorsal aspect; Fig. 41, Omoglymmius
(Pyxiglummius) hesperus new species; Fig. 42, O. (P.) pilosus (Grouvelle); Fig. 43, O. (P.) krikkeni new species; Figs.
44-46, Head, lateral aspect; Fig. 44, O. (P.) hesperus new species; Fig. 45, O. (P.) pilosus (Grouvelle); Fig. 46, O. (P.)
krikkeni new species; Figs. 47-49, O. (P.) pilosus (Grouvelle), male; Fig. 47, Femur, tibia, anterior leg; Fig. 48, Middle
tibia, apical portion; Fig. 49, Femur, tibia, hind leg; Fig. 50, Metasternum and abdomen, ventral aspect, O. (P.) pilosus
(Grouvelle); Fig. 51 Metasternum and abdomen, right half, O. (P.) krikkeni new species; Fig. 52, Right elytron, dorsal
aspect, O. (P.) krikkeni new species; Fig. 53, Elytron, metasternum, abdomen, left lateral aspect, O. (P.) krikkeni new
species; Figs. 54-56, O. (P.) krikkeni new species, male; Fig. 54, Femur, tibia, anterior leg; Fig. 55, Middle tibia; Fig. 56,
Hind tibia.
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Bell and Bell
KEY TO SPECIES
1 Median lobe distinctly longer than wide, its apex opposite middle of eye;
postorbit convex, in form of deep but short, ill-defined postorbital tubercle 2
Y Median lobe transverse, as wide as long, its tip opposite anterior margin of eye;
postorbit flat, bounded ventrally by well-defined sub- or postorbital tubercle 4
2 (1) Medial emargination of temporal lobe relatively shallow, its depth about 0.25 of
its length; basal setae of antenna on Segments VIII-X
O. lederi (Lewis), p. 152
2' Medial emargination of temporal lobe deep, its depth 0.5 or more of its length;
basal setae on Segments V or VI-X 3
3 (2') Temporal lobe broadly rounded posteriorly; postorbital tubercles scarcely visible
in dorsal view; anteriomedial margin of temporal lobe strongly curved
O. subcaviceps (Grouvelle), p. 150
3' Temporal lobe with distinct occipital angle, margin markedly oblique between
occipital angle and eye; orbital tubercles prominent in dorsal view;
anteriomedial margin of temporal lobe oblique . O. crassiusculus (Lewis), p. 151
4 (L) Apex of Segment XI of antenna pointed; metasternum of female without lateral
grooves O. cristatus new species, p. 152
A' Apex of Segment XI obtuse; metasternum of female with deep lateral grooves ... 5
5 (4') Setae absent from Stria II; one or two setae present near apex of Stria IV 6
5' Five or more setae each in Striae II, IV 8
6 (5) Outer carina of pronotum distinctly narrower than paramedian groove; outer
carina nearly straight, of even width O. armatus (Arrow), p. 154
6' Outer carina wider than paramedian groove, widest near middle, tapered both
anteriorly and posteriorly 7
7 (6') Postorbital tubercles large, prominent in dorsal view
O. hesperus new species, p. 1 54
7' Postorbital tubercles small, not prominent in dorsal view
O. strabus(Nev/man), p. 153
8 (5') All elytral striae setose
O. pilosus (Grouvelle), p. 155
8' Stria I and III devoid of setae O.krikkeni new species, p. 156
Omoglymmius (Pyxiglymmius) subcaviceps (Grouvelle) NEW COMBINATION
Figs. 27, 33, 39
Rhysodes subcaviceps Grouvelle 1903: 122.
Pyxiglymmius subcaviceps (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: (VIET NAM) “Tonkin sept. Ha-giang,
Frontiere de Chine, coll. A. Weiss, 1901” (MNHN).
Description. — Length 7-9 mm. Antennal Segment XI 1.5 longer than wide, tip of cone obtuse; basal setae of
antenna sparse on Segment V, well-developed on Segments VI-X; head about as long as wide; median lobe longer than
wide, pointed posteriorly, its lateral angles very obtuse; anteriomedial margin strongly curved; frontal space thus
U-shaped; first medial angles contiguous, separated from second medial angles by rather small but deep emargination,
wich is nearly as deep as long; temporal lobes rounded posteriorly, occipital angles very indistinct; temporal lobe with few
fine, irregularly scattered punctures; postorbit convex, in form of a deep but inconspicuous tubercle.
Revision of Omoglymmius Ganglbauer
151
Pronotum longer than wide, its greatest width less than width across humeral tubercles of elytra; apex of pronotum
strongly narrowed, base moderately narrowed, widest point at middle; lateral margin distinctly sinuate anterior to hind
angle; paramedian groove narrow, about 0.33 as wide as inner carina at middle; paramedian groove distinctly but
inconspicuously punctate; carinae of equal width, both inner and outer ones finely punctate; marginal groove narrow.
Elytra narrow, elongate; apical tubercle very small; Striae III, IV not sinuate posteriorly; strial punctures large,
close together, each about as wide as an interval; male with short lateral groove in posterior part of metasternum (in
female, a much deeper, conspicuous pit); male with proximal tooth of anterior tibia prominent, near middle of length of
tibia; male with acute ventral tooth on anterior femur; middle calcar acute, slightly shorter than spur; calcar of hind leg
very large, its dorsal margin slightly convex, its apex forming a right angle (Fig. 39; female with deep lateral pit in
Sternum IV; margin of elytron angulate opposite Sternum IV.
This species resembles O. crassiusculus in having the postorbit broadly convex, and in
having both pairs of medial angles contiguous. It differs from O. crassiusculus in having the
temporal lobes broadly rounded posteriorly and in having the anteriomedial margin of the
temporal lobe emarginate. The elongate median lobe and the small, contiguous anterior medial
angles will separete it from O. strabus and its relatives.
Range. - Northern Viet Nam. In addition to the holotype, we have seen: a second specimen from
the type locality, collected by Bonifay, 1912 (MNHN) and one male, Hoa Bihn, Tonkin, coll. A. de Cooman, BM
1929-299 (BMNH); two females, N. Viet Nam hills, 50 km. N. Thai-Nguen, 9-1-1964 and 12-1-1964, coll. Kabakov
(LEN); two males, same locality, 300 m. 15-12-1092 and 5-2-1963, coll. Kabakov (LEN); one male, S.W. Kui-Tchau, 300
m„ 19-2-1963, coll. Kabakov (LEN); one male, Mts. S.W. Dong-Hai, 23-XI-1961, coll. Kabakov (LEN).
Omoglymmius (Pyxiglymmius) crassiusculus (Lewis) NEW COMBINATION
Figs. 28, 34
Rhysodes crassiusculus Lewis 1888: 80.
Pyxiglymmius crassiusculus (Lewis) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: “(JAPAN) Kika,
Chizenji, 17-19 April, 1880” (BMNH) PARALECTOTYPES one male, four females, Hakone
(BMNH); one female, Miyanoshita (BMNH); one female, Nikko, Japan, coll. G. Lewis,
1910-320, C. F. Baker Colin. (NMNH); one male, Kiga, Japan, G. Lewis 1910-320; from C. F.
Baker, July 1923 (Cotype No. 26386) (NMNH). Dates on NMNH paralectotypes seem to
refer to the accession date of the Lewis collection, and thus do not preclude them being part of
the original material of Lewis. One specimen labelled as a cotype of R. crassiusculus (NMNH)
actually pertains to Omoglymmius (Boreoglymmius) lewisi Nakane.
Description. — Length 6. 5-8.0 mm. Antennal Segment XI 1 .5 longer than wide, its tip obtuse; basal setae sparse on
Segments V-VI1, numerous on Segments VIII-X; head slightly longer than wide; median lobe longer than wide, pointed
posteriorly; anteriomedial margin of temporal lobe oblique, frontal space thus V-shaped; first medial angles contiguous,
separated from second medial angles by deep emarginations and are longer than in O. subcaviceps; posteriomedial and
posteriolateral margins of temporal lobe oblique, joined at obtuse but distinct occipital angle; temporal lobe with few
irregularly scattered punctures; postorbit convex, in form of deep but inconspicuous postorbital tubercle.
Pronotum narrow, elongate; length/greatest width 1.22; its width conspicuously less than width across humeral
tubercles; apex markedly narrowed; base moderately narrowed; lateral margin distinctly sinuate anterior to hind angles;
paramedian groove narrow, less than 0.25 width of inner carina at middle; paramedian groove distinctly but
inconspicuously punctate; carinae of equal width, both inner and outer ones finely and irregularly punctate; marginal
groove narrow.
Elytra narrow, elongate; apical tubercle more distinct than in O. subcaviceps-. Striae IV, V united posteriorly in a
shallow depression; strial punctures large, close together, each about as wide as an interval; one seta apex of Stria IV, one
in stride, several in VII near apex; metasternum without lateral grooves in either sex; male with proximal tooth of anterior
tibia prominent, near middle of length of tibia; male with low, obtuse tooth on ventral margin of anterior femur; middle
calcar acute, slightly shorter than spur; hind calcar very large, its dorsal margin straight, its apex in form of acute angle;
female with deep, distinct lateral furrow on abdominal Sterna I- 1 1; female with very large lateral pits on Sternum IV,
separated medially by less than width of one of them; lateral pit with small, inconspicuous anteriolateral brace; pit smooth,
without striae or punctures; ventral margin of elytron conspicuously angulate opposite Sternum IV in female, slightly
angulate in male.
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Bell and Bell
The distinct occipital angles give the head of this species a “diamond” shape, separating it
from all other members of the subgenus.
Range. - Japan. In addition to type material, we have seen one specimen from Shuzenji Izu
(CAS).
Omoglymmius (Pyxiglymmius) lederi (Lewis) NEW COMBINATION
Figs. 29, 35
Rhys odes lederi Lewis 1888: 82.
Pyxiglymmius lederi (Lewis) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “Caucasus, coll. Saunders, G. Lewis,
1901-31” (BMNH). This distinctive species has not been collected subsequently in the
Caucasus, and we suspect that the locality label is erroneous. The species probably really comes
from one of the poorly collected countries of the Oriental Region.
Description. — Length 8.9 mm. Antennal Segment XI slightly longer than wide; apex of cone obtuse; basal setae
sparse on Segments Vlll-IV; numerous on Segment X; head as wide as long; median lobe elongate, narrow, pointed
posteriorly; separated from clypeus by shallow transverse impression; anteriomedial margin of temporal lobe oblique; both
first and second posterior medial angles narrowly separated; first medial angle separate from second medial angle by broad
but relatively shallow emargination which is twice as long as deep; occipital angle indistinct, temporal lobe rounded
posteriorly; temporal lobe with a few coarse, irregularly scattered punctures; postorbit convex, in form of deep but
inconspicuous postorbital tubercle.
Pronotum narrow, elongate; length/greatest width 1 /26; apex markedly narrowed; base moderately so; lateral margin
distinctly sinuate anterior to hind angle; paramedian groove narrow, 0.33 as wide as inner carina at middle; carinae of
equal width, both outer and inner carinae finely, irregularly punctate, more densely so than in O. crassiusculus\ marginal
groove narrow.
Elytra narrow, elongate; striae coarsely punctate; no setae in Stria IV, one in apical stride, several in Stria VII near
apex; ventral margin of elytron conspicuously angulate opposite lateral abdominal pit in Sternum IV of female;
metasternum without lateral groove in female; abdominal Sternum I with lateral pit in female; Sternum IV of female with
moderately large lateral pits (pits separated by more than twice the width of one of them); pits of Sternum IV without
anteriolateral braces, irregularly pitted, but without longitudinal stria.
Male unknown.
Omoglymmius ( Pyxiglymmius ) cristatus new species
Figs. 30, 36, 40
Type material. - HOLOTYPE female, labelled: “PHILIPPINES, Mt. Makiling, Laguna
(Province), 1-1 1931, F. C. Hadden Colin. ”(CAS). The type locality is on Luzon.
Description. — Length 8.0 mm. Antennal Segment XI distinctly longer than wide, apex of cone acutely pointed;
basal setae numerous on Segments V-X (Fig. 40); head slightly wider than long; antennal lobes more elevated than in
other members of subgenus; clypeus coarsely punctate, separated from median lobe by shallow transverse impression;
median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique; first medial angles widely
separated; second medial angles contiguous; temporal lobe rounded posteriorly; temporal lobe convex, with about 30 coarse
punctures; postorbit flat dorsad to tubercle; latter located more dorsad than in O. strabus and related species; postorbital
tubercle about 0.33 as deep and 0.33 times as wide as eye.
Pronotum relatively short; length/greatest width 1.16; base of pronotum moderately narrowed; apex very strongly
narrowed, lateral margin slightly sinuate anterior to hind angles; paramedian grooves punctate, rather broad, about 0.66 as
wide as inner carinae at middle; carinae of equal width at middle, both inner and outer ones coarsely punctate; marginal
groove rather fine, about 0.25 as wide as outer carina at middle.
Elytra moderately elongate; apical tubercle small; strial punctures very large, each equal in width to an interval; one
seta present in apex of Stria IV, one in apical stride; several near apex of Stria VII; female with shallow lateral groove in
posterior fourth of lateral margin of metasternum; female with indistinct lateral impression on Sternum I and large, deep
lateral pits on Sternum IV, separated medially by less than twice width of one of them; lateral pit of Sternum IV with
prominent anteriolateral brace and with confused punctuation, but without longitudinal striation. Ventral margin of
female elytron indistinctly angulate opposite Sternum IV.
Revision of Omoglymmius Ganglbauer
153
Male unknown.
This species is similar to O. strabus, from which it differs in having the apex of the antenna
pointed, the head shorter and wider, the antennal lobes more elevated, and the temporal lobes
more coarsely and densely punctate.
Omoglymmius (Pyxiglymmius) strabus (Newman) NEW COMBINATION
Figs. 31, 37
Rhysodes strabus Newman 1838: 663-664.
Rhysodes aterrimus Chevrolat 1873: 209. NEW SYNONYMY
Pyxiglymmius strabus (Newman) Bell and Bell 1978
Type material. - ( R . strabus) HOLOTYPE female, labelled: “Java, Ent. Club, 44-12”
(labelled as type) (BMNH); (R. aterrimus) HOLOTYPE female, labelled: “Malacca, S. S”,
“typus” (NMW).
Description. — Length 7. 5-9.0 mm. Antennal Segment XI slightly wider than long, its tip obtusely rounded; basal
setae numerous on Segments V-X; head slightly wider than long; clypeus with a few coarse punctures or impunctate,
continuous with median lobe; median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique;
first medial angles widely separated; second medial angles contiguous or very narrowly separated; temporal lobe rounded
posteriorly; temporal lobe convex, with one to 12 coarse punctures; postorbit flat dorsad to tubercle; latter located opposite
ventral third of eye; postorbital tubercle about 0.33 as deep and 0.33 as long as eye.
Pronotum rather short, length/greatest width ranging from 1.12-1.19; base of pronotum moderately narrowed; apex
stongly narrowed; lateral margin slightly sinuate anterior to hind angles; paramedian grooves rather broad, punctate,
about 0.66 as wide as inner carina at middle; inner carina slightly narrower than outer carina at middle; both pairs of
carinae coarsely punctate, inner ones more sparsely so than outer ones; marginal groove fine, less than 0.25 as wide as
outer carina at middle.
Elytra moderately elongate; apical tubercle very small; strial punctures large, each as wide as an interval; one seta
present in apex of Stria IV; one in apical striole, several near apex of Stria VI 1; female with broad, distinct groove along
each lateral margin of metasternum; female with indistinct lateral impression on abdominal Sternum I, and large, deep
lateral pits on Sternum IV, which are separated medially by more than twice the width of one of them; lateral pit of
Sternum IV with prominent aneriolateral brace and with longitudinal striation; ventral margin of female elytron not
angulate opposite Sternum IV; male with well-developed proximal tooth at basal third of anterior tibia; male with ventral
tooth on anterior femur; calcar of middle tibia acute, about 0.5 as long as spur; calcar of hind tibia large, triangular, acute,
with dorsal margin curved.
This species may be separated from the sympatric O. pilosus and O. krikkeni by the small
number of setae on the elytra. The type specimens of O. strabus and O. aterrimus at first
glance look like separate species, that of O. strabus having the temporal lobes nearly
impunctate, while that of O. aterrimus is distinctly punctate. However, numerous specimens
from Java show that this is individual variation. O. armatus, O. hesperus , and O. cristatus are
similar to O. strabus , and appear to have developed from populations of the latter species
isolated respectively on the Andaman Islands, the Mentawei Islands, and Luzon. For
differences between O. strabus and each of these species, see discussion of the latter.
Range. - Borneo, Java, Sumatra, and the Malay Peninsula. In addition to the types, we have
Studied the following specimens: BORNEO: one male, one female, R. Kapah, trib. of R. Tinjar, 3-X, 1932, on
bark, felled timber, Oxford Univ. Exp., coll. B. M. Hobby & A. W. Moore, BM 1933-254 (BMNH); JAVA: 14 males, 5
females. Preanger (MNHN); one male, Mt. Tengger (MNHN); two females, without specific locality (LEI); MALAYA:
one male, Perak, coll. Doherty, 58794, Fry. Colin. 1905-100 (BMNH); SUMATRA: one male, Alas Valley, Gumpang,
foothills of Mt. Kemiri, 10-V1-1972, ca 780 m., coll. J. Krikken (LEI); one female, Palembang (MNHN).
Records for O. aterrimus from the Andaman and Nicobar Islands refer to O. armatus.
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Bell and Bell
Omoglymmius (Pyxiglymmius) armatus. (Arrow) NEW COMBINATION
Figs. 32, 38
Rhysodes armatus Arrow 1901: 85.
Pyxiglymmius armatus (Arrow) Bell and Bell 1978
Grouvelle (1903) synonymized the names R. armatus and R. aterrimus Chevrolat 1873.
Bell and Bell (1978) showed this to be an error.
Type material. - LECTOTYPE (here designated) male, labelled: “Andamans
(Roepstorff)” (BMNH). PARALECTOTYPES one female, same data as lectotype (teneral
specimen) (BMNH); one female, labelled: “Nicobars (Roepstorff)” (BMNH).
Description. — Length 6. 1-8.0 mm. Antennal Segment Xi slightly longer than wide, its apex obtuse; basal setae
sparse on Segment V, numerous on Segments VI-X; head slightly wider than long; clypeus impunctate; separated from
median lobe by a more or less distinct transverse impression; median lobe short, strongly transverse, its posterior margin
varying from nearly transverse to distinctly angled at middle (i.e., median lobe triangular to rhomboid); anteriomedial
margin of temporal lobe oblique; first medial angles separated; second medial angles contiguous; temporal lobe rounded
posteriorly; temporal lobe convex, with 10-15 scattered punctures; postorbit flattened dorsad to tubercle; latter more
prominent than in O. strabus, about 0.5 as deep and 0.66 as long as eye.
Pronotum more elongate than in O. strabus , length/greatest width 1.32, widest anterior to middle; in some specimens
distinctly dilated anterior to middle, in others almost parallel-sided; margin various from scarcely to markedly sinuate
anterior to hind angles, latter obtuse; paramedian grooves broad, punctate, nearly straight, as wide as or wider than inner
carina at middle; both pairs of carinae very narrow; outer carina in some specimens slightly narrower than inner one, in
others, opposite; both carinae sparsely, coarsely punctate; marginal groove very deep and dilated, as wide as outer carina or
nearly so.
Elytra moderately elongate; apical tubercle larger than in O. strabus ; strial punctures large, each equal in width to one
interval; one seta in apex of Stria IV, one on apical striole, several near apex of Stria VII. Female with deep lateral grooves
on metasternum; lateral pits of Sternum IV of female separated by 1.25 times width of one of them; lateral pit with
anteriolateral brace and longitudinal striae.
Proximal tooth of anterior tibia present, but various in size and position; ventral tooth on anterior femur present, also
various in size and position; middle calcar approximately equal in length to spur; hind tibia various in degree of swelling in
proximal third, and especially in size of calcar.
The deep, dilated marginal grooves of pronotum are diagnostic, and form a striking contrast
with the very fine grooves of O. strabus. There are considerable differences among specimens
we assign to O. armatus , including: shape of pronotum, size of postorbital tubercles, depth of
lateral metasternal grooves, size of lateral abdominal pits of females; and particularly in size of
the hind calcars of males. It seems possible that there are subspecific or perhaps specific
differences among the populations on the various islands, but available specimens are too few
and too inexactly labelled to allow definite conclusions.
Range. - Andaman and Nicobar Islands. In addition to type material, we have seen one
male from the Andaman Islands (MNHN), and two males and four females from the Nicobar
Islands (MNHN).
Omoglymmius ( Pyxiglymmius ) hesperus new species
Figs. 41, 44
Type material. - HOLOTYPE male, labelled: “Mentawei:Sipora, Sereinu V-VI-94,
Modigliani, 51” (MNHN). This island, also spelled “Sipoera” or “Sipura,” is in the Indian
Ocean, west of Sumatra. PARATYPES two females, three males, labelled: “Mentawei, Si
Oban, IV-VIII, Modigliani 94” (GEN).
Description. — Length 7. 9-9.0 mm. Antennal Segment XI slightly wider than long, its apex obtuse; basal setae
absent from Segment V, sparse on VI, VII, numerous on VII-X; head slightly longer than wide, smaller relative to
pronotum than in other Pyxiglymmius-, clypeus punctate, pollinose, continuous with median lobe; median lobe rhomboid,
wider than long, its posterior angle obtuse; anteriomedial margin of temporal lobe oblique; first medial angles distinctly
Revision of Omoglymmius Ganglbauer
155
though narrowly, separated; second medial angles contiguous; occipital angles very obtuse but distinct; posteriolateral
margin of temporal lobe slightly oblique; each temporal lobe with seven to 10 coarse punctures; postorbit slightly
concave dorsad to postorbital tubercle; latter large, 0.75 as long and 0.66 as deep as eye, prominent in dorsal aspect;
width across postorbital tubercle greater than width across eyes.
Pronotum short, length/greatest width 1.16; its greatest width greater than width of elytra across humeri; widest
point posterior to middle; apex of pronotum very strongly narrowed; base moderately narrowed; lateral margin scarcely
sinuate anterior to hind angle; paramedian grooves broad, punctate, shallower than in other members of subgenus;
paramedian groove equal in width to inner carina at middle; distinctly narrower than outer one at middle; both carinae
coarsely, densely punctate; marginal groove fine, about 0.20 as wide as outer carina at middle.
Elytra relatively short, broad; strial punctures smaller than in related species, each puncture less than half width of
one interval; one seta in apex of Stria IV, one in apical stride; several setae near tip of Stria VII.
Male with proximal tooth of front tibia small, more proximad than in related species; ventral tooth of anterior
femur of male small, at apical third; calcar of middle tibia appearing truncate, 0.5 as long as spur (but calcar in
holotype may be broken or worn), hind tibia only slightly swollen proximad to middle; hind calcar very large, its tip
slightly obtuse.
Female with short, deep lateral groove in metasternum; lateral margin of latter not flared; lateral pits of Sternum
IV rather small, separated by more than 5 times width of one of them; pit with very strong anteriolateral brace, without
longitudinal striation.
This species has quite different body proportions from other members of the subgenus, with
a short, wide pronotum, a very small head, and short elytra. The paramedian grooves are
shallower and more coarsely punctate than in related species, and the postorbital tubercles are
larger than in other Pyxiglymmius, although they are small in comparison to those of
Omoglymmius (Laminoglymmius) gorgo.
Omoglymmius ( Pyxiglymmius ) pilosus (Grouvelle) NEW COMBINATION
Figs. 42, 45, 47-50
Rhysodes pilosus Grouvelle 1903: 123-124.
Pyxiglymmius pilosus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: “Bengkalis-Maindron
1885” (SUMATRA) (MNHN); PARALECTOTYPES one female, same data as lectotype.
These specimens are labelled as “Types” in the Grouvelle Collection (MNHN), but are not
mentioned in the original description. Additional PARALECTOTYPES, SUMATRA; two
males, four females, Palembang (MNHN); one male, Palembang, Fry Colin. 1905-100,
labelled “co-type” (BMNH); one female, Palembang, 1901-267 (BMNH); BORNEO: two
males, Occ. riv. Sambey, pres Ngabang, coll. J. B. Ledru, 1897, labelled “co-type” (BMN).
Since these localities are mentioned in the original description, they must also represent
paralectotypes. Grouvelle also mentions specimens from the rivers Pontianac and Mander and
Guenong-Ampar in Borneo Occ. We have not located these specimens, but they may be among
recently discovered material in the Oberthur Collection in Paris.
Description. — Length 7.8-10.0 mm. Antennal Segment XI slightly longer than wide, its apex obtuse; basal setae
numerous on Segments V-X; head slightly longer than wide; clypeus coarsely punctate; median lobe continuous with
clypeus; median lobe rhomboidal, wider than long, its lateral and posterior angles acute; anteriomedial margin of temporal
lobe oblique; first medial angles broadly separated; emargination between first and second medial angles very deep; second
medial angles narrowly separated; nearly oblique between second medial angle and occipital angle; each temporal lobe
with 4-7 very coarse punctures in lateral half, some of these with very short setae; postorbit flat dorsad to tubercle; latter
prominent, 0.33 as deep and 0.67 as long as eye; width across postorbital tubercle slightly greater than width across
temporal lobes.
Pronotum rather short, length/greatest width 1.13, widest near middle; base moderately and apex strongly narrowed;
lateral margin at most slightly sinuate anterior to hind angle; median and paramedian grooves broad, rather shallow,
prominently punctate; inner carinae slightly narrower than outer carinae at middle; both pairs of carinae very coarsely,
rather densely punctate; marginal groove shallow, rather broad, punctate.
Elytra moderately elongate; apical tubercle very small; strial punctures very large, some of them slightly irregular in
arrangement; all striae with numerous setae; female with deep, broad lateral groove on metasternum, lateral margin of
Quaest. Ent., 1982, 18 (1-4)
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Bell and Bell
groove conspicuously flared, groove continuous with lateral pit on abdominal Sternum 1; female with very large lateral
pits on Sternum IV, separated medially by less than width of one of them (Fig. 50) pit of Sternum IV with
anteriolateral brace and longitudinal striae; medioposterior border of pit straight, oblique; ventral margin of female
elytron angulate opposite abdominal Sternum I; male with shallow lateral pit on Sternum IV and faint suggestion of
lateral furrow on metasternum.
Male with proximal tooth on anterior tibia located slightly dorsad of middle of tibia; male with ventral tooth on
anterior femur located distad to middle (Fig. 47); calcar of middle tibia about 0.67 as long as spur (Fig. 48); calcar of
hind tibia very large, obtusely pointed (Fig. 49).
Presence of setae on all elytral striae will separate this species from all other members of the
subgenus. In this character it agrees with the sympatric Omoglymmius ( Laminoglymmius )
rugosus, which differs in having a setose pronotum and differently shaped temporal lobes.
Range. - Sumatra and Borneo. In addition to type material, we have seen the following
specimens (all from Sumatra): one male, Bedagei, int. Sumatra s O.K., 600 ft., 2 Sem. 89, coll. 1. Z.
Kannegieter (MNHN); one male. Deli, coll. Dr. Busey (AMS); one male, Manna, coll. M. Knappert (LEI); two males,
one female, Roagu, no other data (MNHN); two females, Tandjong, Morawa, Serdang, N. O. Sumatra, coll. Dr. B. Hagen
(LEI); two females, Tebing-Tinggi, coll. Dr. Schultheiss (MNHN).
Omoglymmius (Pyxiglymmius) krikkeni new species
Figs. 43,46,51-56
Type material. - HOLOTYPE male, labelled: “N. SUMATRA, Alas Valley, Gumpang:
Foothills Mt. Kemiri, 3° 47' N, 97° 27' E, 10-VI-1972, ca. 780 m., coll. J. Krikken, no. 10,
from rotten trunk, lowland multistratal evergreen forest” (LEI). PARATYPES one male, eight
females, same data as type (LEI).
Description. — Length 8.3-10.3 mm. Antennal Segment XI slightly wider than long, its apex obtuse; basal setae
numerous on Segments V-X; head slightly longer than wide; clypeus finely punctate; median lobe continuous with clypeus;
median lobe rhomboidal, wider than long; anteriomedial margin of temporal lobe oblique, first medial angles broadly
separated; emargination between first and second medial angles shallower than in O. pilosus\ second medial angles
narrowly separated; margin distinctly rounded between second medial angle and occipital angle; each temporal lobe with
three to seven coarse punctures; postorbit flat dorsad to tubercle, latter prominent 0.33 as deep and 0.67 as long as eye;
width across postorbital tubercles slightly less than width across eyes.
Pronotum rather short, length/greatest width 1.21; widest near middle, base strongly narrowed, apex moderately
narrowed; lateral margin slightly sinuate anterior to hind angles; median and paramedian grooves broad, rather shallow,
punctate; inner carina slightly narrower than outer carina at middle; both pairs of carinae with few, irregularly distributed
punctures, nearly absent from anterior third; marginal groove fine, shallow.
Elytra moderately elongate; Striae 111, IV slightly sinuate anterior to very small apical tubercle; strial punctures very
coarse; elytron with row of setae in Interval III slightly laterad to Stria 11, and another row in Interval V slightly laterad to
Stria IV; two or three setae in Interval VI slightly laterad to Stria V; row of setae in Stria VII; distinct depression in
Intervals IV, V just anterior to apical tubercle (Fig. 52); female with broad, deep lateral groove on metasternum, lateral
margin of groove conspicuously flared (Fig. 53); female with deep lateral pit on abdominal Sternum 1; female with very
large lateral pits on Sternum IV, separated medially by slightly more than the width of one of them; pit of Sternum IV
with anteriolateral brace and longitudinal striae; medioposterior margin of pit curved; ventral margin of elytron of female
expanded, angulate, opposite hind coxa, and slightly angulate opposite lateral pit of Sternum IV (Fig. 51); male with
small, shallow lateral groove on metasternum and slight, irregular lateral pits on Sterna III, IV, V.
Male with proximal tooth of anterior tibia slightly dorsad to middle; male with ventral tooth of anterior femur
rounded, nearer to apex than to middle (Fig. 54); calcar of middle tibia acute, longer than spur (Fig. 55); calcar of hind
tibia very large, its dorsal margin straight, its tip acute (Fig. 56).
This very large species is easily recognized by the pattern of setae on the elytra. We dedicate
this species to its discoverer, Dr. J. Krikken, in appreciation of his help in our studies.
Subgenus Laminoglymmius new subgenus
Type species. - Rhysodes insularis Grouvelle 1903
Description. — Segment XI of antenna obtuse, without stylet; basal setae on distal antennal segments; clypeal setae
absent; medial margin of temporal lobe with pale translucent area (conspicuous in all species except O. actae, in which it is
Revision of Omoglymmius Ganglbauer
157
obsolete); some species with similar translucent area at tip of median lobe; medial margin of temporal lobe various from
entire ( O . inaequalis ) to bilobed or trilobed, with one, two, or three medial angles; most species with postorbital tubercle
conspicuous, on some species located nearly suborbitally; punctures of abdominal sterna scattered, numerous; female
with small, simple lateral pits in Sternum IV, male with shallow lateral pits in Sternum IV; cleaning organ located
entirely distad to base of anterior tarsus; spur of middle tibia curved; males, where known, with hind calcar small, at or
near base of spur.
Most members of this subgenus are recognized by the presence of a translucent medial
margin, with one or two shallow emarginations on the temporal lobe separating two or three
medial angles, which are close together. Two species are atypical: in O. actae , the translucent
area is scarcely developed, though there is a shallow but distinct emargination. In O. inaequalis
the medial margin is not emarginate, though the translucent area is well marked.
Those species with the medial margin of the temporal lobe distinctly emarginate might be
confused with members of the subgenus Pyxiglymmius. In the latter subgenus, the two medial
angles are widely separated from one another, and the emargination between them is deep in all
species except for O. lederi. O. inaequalis , which lacks the medial emargination, might be
mistaken for Subgenus Indoglymmius. The latter has a well-marked translucent area on the
median margin, but differs in lacking basal setae from the antennae.
The subgenus is largely confined to a small part of the Oriental Region, where it is found in
Sumatra, Borneo, the Malay Peninsula, and the Nicobar Islands. O. actae is found in New
Guinea, far from the other species. It is aberrant in the poor development of the translucent
areas, and is perhaps not really related to the other species. Laminoglymmius is especially well
represented in Sumatra, where there are four sympatric species.
Phytogeny. - A possible phylogeny for the subgenus is presented in Diagram 2. We
postulate that: Species 1, the ancestor of the subgenus, had two medial angles on each temporal
lobe, both included in a distinct translucent area; clypeus was continuous with the median lobe,
punctures of pronotum coarse, moderately dense and relatively evenly distributed; and elytral
setae were absent except for the tip of Stria VII and perhaps the apical striole.
We suggest that O. rugosus is a highly modified descendant of Species 1, having developed
the following features: a transverse groove between the clypeus and the median lobe; numerous
setae on the head, pronotum, and elytra; a third opaque medial angle posterior to the second
one and very dense, confluent pronotal punctures.
All other species are probably descended from Species 2. This was similar to Species 1
except that the pronotal punctures became much sparser than is typical for most species of
Omoglymmius. Species 2 was probably very similar to O. oberthueri, which may be regarded
as an almost unmodified descendent. O. actae is a similar species, marked by the virtual
disappearance of the translucent areas of the temporal lobe. These two species show no obvious
evidence of close relationship with each other, or with the four remaining species. The latter
appear to be descended from two daughter species of Species 2 (Species 3 and 4). Species 3 and
4 might have a common ancestor more recent than Species 2. The only evidence for this is the
presence in all four daughter species of a translucent area at the tip of the median lobe.
In Species 3, the reduction of punctation of the pronotum was extreme, O. insularis lacking
punctures entirely and O. inaequalis having only a single puncture on the outer carina. In
Species 4, the punctuation was less reduced, approximately six punctures being retained on
each outer carina, but the pronotum became rounded in outline, the median lobe became
narrow and elongate, with nearly parallel sides, and the postorbital tubercles were enlarged to
the point of being visible in dorsal view.
Species 3 gave rise to O. insularis and O. inaequalis. In O. insularis , the median lobe
became concave, all punctures were lost from the outer carina, and the temporal lobe lost all its
Quaest. Ent., 1982, 18 (1-4)
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Bell and Bell
Phylogenetic Diagram 2. Reconstructed Phylogeny of species of Laminoglymmius.
punctures. O. inaequalis retained a single puncture at the middle of each outer carina, while
the outer carina became very narrow, the postorbital tubercle was displaced ventrally to lie
opposite the lower margin of the eye, and the anteriomedial margin of the temporal lobe
became oblique, so there is only one medial angle.
Species 4 gave rise to O. trisinuatus and O. gorgo. In O. trisinuatus, a third medial angle
was developed, while in O. gorgo , the postorbital tubercles became greatly enlarged.
KEY TO SPECIES
1 Median lobe separated from clypeus by transverse pollinose groove; temporal
Revision of Omoglymmius Ganglbauer
159
Plate 5. Figures 57-74, subgenus Laminoglymmius. Figs. 57-63, Head and pronotum, dorsal aspect; Fig. 57,
Omoglymmius ( Laminoglymmius ) rugosus (Grouvelle); Fig. 58, O. (L.) inaequalis new species; Fig. 59, 0. (L.) actae new
species; Fig. 60, O. (L.) insularis (Grouvelle); Fig. 61, O. (L.) oberthueri (Grouvelle); Fig. 62, O. (L.) trisinuatus new
species; Fig. 63, O. (L.) gorgo new species ; Figs. 64-66, O. (L.) rugosus (Grouvelle), male; Fig. 64, Femur, tibia, anterior
leg; Fig. 65, Middle tibia; Fig. 66, Hind tibia; Fig. 67, Hind tibia, male, O. (L.) actae new species; Figs. 68-74, Head,
lateral aspect; Fig. 68, O. (L.) rugosus (Grouvelle); Fig. 69, O. (L.) inaequalis new species; Fig. 70, O. (L.) actae new
species; Fig. 71, O. (L.) insularis (Grouvelle); Fig. 72, O. (L.) oberthueri (Grouvelle); Fig. 73, O. (L.) trisinuatus new
species; Fig. 74, O. (L.) gorgo new species.
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Bell and Bell
lobes, genae, pronotum, and elytra with numerous setae
O. rugosus (Grouvelle), p. 160
r Median lobe continuous with clypeus; head, pronotum without setae; setae of
elytron limited to a few in apex of Stria VII and none or one on apical tubercle ... 2
2 (1) Outer carina distinctly narrower than inner carina 3
2' Outer carina not narrower than inner carina 4
3 (2) One medial angle on temproal lobe; translucent area present; outer carina 0.25
width of inner carina; outer carina with one puncture near middle, inner carina
impunctate O. inaequalis new species, p. 161
3' Two medial angles on temporal lobe; latter without translucent area; outer
carina 0.5 width of inner carina; outer carina with five punctures, inner carinae
with one or two O. actae new species, p. 161
4 (2') Median lobe flat; outer carina punctuate 5
4' Median lobe concave; pronotum entirely impunctate
O. insularis (Grouvelle), p. 162
5 (4) Inner carina of pronotum with three or four coarse punctures; median lobe of
head rhomboidal O. oberthueri (Grouvelle), p. 162
5' Inner carina impunctate; median lobe of head narrow, its margins parallel or
nearly so 6
6 (5') Temporal lobe with three medial angles; postorbital tubercle relatively small, its
length about 0.5 of diameter of eye O. trisinuatus new species, p. 163
6' Temporal lobe with two medial angles; postorbital tubercle very large, its length
equal to diameter of eye O. gorgo new species, p. 163
Omoglymmius (Laminoglymmius) rugosus (Grouvelle 1903) NEW COMBINATION
Figs. 57, 64-66, 68
Rhysodes rugosus Grouvelle 1903: 122-123.
Pyxiglymmius rugosus (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE sex not recorded, labelled: “SINGAPORE, Raffray”
(MNHN).
Description. — Length 6. 9-8. 5 mm. Segment XI of antenna wider than long, rounded at apex; basal setae present
on Segments V-XI; head relatively short; clypeus punctate, microsculptured, separated from median lobe by deep
transverse pollinose groove; antennal rim very thick, elevated; frontal space minute; temporal lobe with three medial
angles, first and second ones translucent, each narrowly separated from corresponding angle of opposite temporal lobe;
third medial angles opaque, more broadly separated from each other; temporal lobe very convex, shining, with about 15
coarse punctures near lateral margin, most of punctures with stout but very short setae; temporal lobe rounded posteriorly;
postorbital tubercle moderate in size, with deep concavity dorsad to it; postorbital tubercle densely coarsely punctate,
irregular network of ridges between punctures; some punctures with short setae.
Pronotum moderately long; length/greatest width 1.15; widest point distinctly anterior to middle, apex narrowed,
lateral margins oblique, nearly straight posteriorly, in form of long but shallow sinuation anterior to rectangular hind
angle, pronotal grooves broad, rather shallow, ill-defined, finely pollinose, coarsely punctate; carinae very coarsely,
irregularly rugose, punctate; each puncture with seta; inner and outer carinae nearly equal at middle; marginal groove
ill-defined medially, but with its lateral margin raised, in form of fine carina.
Elytral striae very broad, intervals narrow, about 0.5 as wide as striae; intervals convex; strial punctures very irregular,
in places in form of two parallel rows within stria; all striae with many setae; apical tubercle very small; female with
distinct lateral pits on Sternum IV.
Male with sharp ventral tooth at apical third of anterior femur (Fig. 64); female with sinuation in this posititon;
proximal tooth at middle of anterior tibia in male; middle tibia of male with a small, acute calcar (Fig. 65); hind calcar
small, obtuse, distinctly dorsad to level of spur (Fig. 66).
Revision of Omoglymmius Ganglbauer
161
The numerous setae of the pronotal carinae, genae, and temporal lobe separate this large
species from all others. Large size, completely setose striae and partly irregular strial
punctation give a superficial similarity to the sympatric Omoglymmius (Pyxiglymmius)
pilosus, but the form of the temporal lobes and the absence of pronotal setae eliminate the
latter species.
Range. - Singapore, Sumatra and Borneo. In addition to the type, we have studied the
following specimens: BORNEO: one male, one female, Mt. Matang, West Sarawak, coll. G. E. Bryant, 31-1-14
(BMNH); SUMATRA: one male, one female, Palembang (MNHN).
Omoglymmius (Laminoglymmius) inaequalis new species
Figs. 58, 69
Type material. - HOLOTYPE female, labelled: “Nicobars” (MNHN). This specimen was
identified by Grouvelle as Rhysodes nicobarensis Grouvelle (1895b). The latter species, as
shown by the holotype, also in Paris, belongs to Subgenus Orthoglymmius , and, despite the
name, was collected from the Andaman Islands. Our placement of R. nicobarensis in Subgenus
Hemiglymmius (Bell and Bell 1978) was based on a study of O. inaequalis , and not the true R.
nicobarensis.
Description. — Length 6.5 mm. Antennal Segment XI as wide as long, its apex obtusely rounded; basal setae sparse
on Segment V, numerous on Segments VI-X; clypeus impunctate, continuous with median lobe; latter also impunctate,
broad, its tip pointed, translucent; anteriomedial margin of temporal lobe almost straight, small but distinct semicircular
translucent area present anterior to medial angle; only one medial angle present; margin oblique between medial and
occipital angles; temporal lobe convex, shining, with 10-12 coarse punctures present in lateral half; postorbital tubercle
large, 0.5 as deep and 0.66 as long as eye, its dorsal margin only slightly above ventral margin of eye (so it could better be
termed a “suborbital tubercle”); width across potorbital tubercles slightly less than width across temporal lobes.
Pronotum relatively short, length/greatest width about 1.1; widest point slightly anterior to middle, sides strongly
curved, convergent to apex, which is narrow; base relatively broad, lateral margin not sinuate anterior to hind angle, which
is slightly obtuse; paramedian groove deep, broad, more than 0.33 as wide as inner carina at middle; outer carina very
narrow, less than 0.25 as wide as inner carina at middle; outer carina with one minute puncture near middle, this puncture
with minute seta; inner carina entirely impunctate; marginal groove dilated, over 0.5 width of outer carina at middle.
Elytra relatively short, broad; striae very shallow; intervals flat, finely microsculptured, with faint sheen; strial
punctures very small, each about 0.20 as wide as an interval; elytron with several setae in apex of Stria VII; female with
large, deep lateral pit on Sternum IV; female with small ventral tooth on anterior femur; male unknown.
This is the only member of the subgenus in which there is only one medial angle. The very
narrow outer carina with a single minute setiferous puncture is also diagnostic. The
combination of a single medial angle with a well-developed translucent area is suggestive of
Subgenus Indoglymmius. Members of the latter subgenus differ in lacking basal setae on the
antennae, and in having the inner and outer carinae of equal width.
Omoglymmius (Laminoglymmius) actae new species
Figs. 59, 67, 70
Type material. - HOLOTYPE male, labelled: “NEW GUINEA, Maffin Bay, IX- 1944, E.
S. Ross” (CAS).
Description. — Length 5.5 mm. Antennal Segment XI slightly wider than long, its apex rounded; basal setae
numerous on Segments V-X; clypeus continuous with median lobe; latter rhomboidal, with a few coarse punctures; head
distinctly longer than wide; anteriomedial margin of temporal lobe oblique; latter with two minute medial angles which are
located close together; without obvious translucent area near medial angles (under strong light, translucent area is faintly
suggested); temporal lobe convex, shining, with 12-15 coarse scattered punctures; postorbital tubercle small, 0.25 as deep
and 0.33 as long as eye, located slightly above ventral margin of eye; width across postorbital tubercles less than width
across eyes.
Quaest. Ent., 1982, 18 (1-4)
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Bell and Bell
Pronotum moderately elongate, length/ greatest width 1.20; widest near middle, lateral margins curved; apex
strongly narrowed; base moderately narrowed; lateral margin not sinuate anterior to hind angle; latter obtuse;
paramedian grooves deep, rather narrow; outer carina distinctly narrower than inner one, about 0.33 as wide as inner
one at middle; outer carina with four or five scattered coarse punctures, without setae; inner carina with one or two
coarse punctures; marginal groove dilated, about 0.33 as wide as outer carina at middle.
Elytra narrow, convex; elytral striae deep, coarsely punctate, each puncture about 0.6 as wide as in interval;
intervals narrow, convex, shining; several setae present in apex of Stria VII; male with distinct proximal tooth on
anterior tibia and ventral tooth at apical fourth of anterior femur; middle calcar small, triangular; hind calcar obtuse,
triangular, located 0.33 of length of tibia dorsad to spur (Fig. 67); female unknown.
This species is the only member of the subgenus which does not have well marked
translucent areas near the medial angles of the temporal lobes. It is recognized as a member of
Subgenus Laminoglymmius by the double medial angles. This character and the presence of
basal setae on the antenna will separate it from superficially similar species of Omoglymmius s.
str. from New Guinea.
Omoglymmius (Laminoglymmius) insularis (Grouvelle) NEW COMBINATION
Figs. 60,71
Rhysodes insularis Grouvelle 1903: 124-125.
Pyxiglymmius insularis (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “Bengkalis, Maindron 1885” (MNHN).
The type locality is a small coastal island near Sumatra.
Description. — Length 7.0 mm. Segment XI of antenna as wide as long, its apex rounded; basal setae sparse on
Segments V-VI, dense on Segments VII-X; head slightly longer than wide; clypeus continuous with median lobe; clypeus
impunctate; median lobe elongate oval, deeply concave at center, its margins^ raised; tip of median lobe rounded,
translucent; temporal lobe with two medial angles, separated by very shallow emargination; first medial angles separated
from one another by approximately the width of one of them; second medial angles separated by about half the distance;
both angles and adjacent margins in a translucent area; temporal lobe evenly rounded posteriorly, laterally; temporal lobe
entirely smooth, impunctate scarcely overhanging postorbit, latter smooth, glabrous; postorbital tubercle small but
well-defined; gular tubercle slightly prominent.
Pronotum moderately elongate, length/greatest width 1.25; widest anterior to middle, lateral margins curved, apex
strongly narrowed; base moderately narrowed, lateral margin distinctly sinuate anterior to hind angle, which is nearly
rectangular; paramedian grooves deep, rather narrow, outer carina slightly narrower than inner one at middle; both
carinae entirely impunctate, without setae; marginal groove relatively narrow.
Eltyra moderately elongate, interval V slightly elevated anteriorly; striae relatively narrow, each puncture about 0.25
as wide as interval; several setae present in apex of Stria VII; female with deep, round, small lateral pit on abdominal
Sternum IV and with small ventral tooth on anterior femur; male unknown,
The deeply concave, oval median lobe separates this species from all other Rhysodini.
Omoglymmius ( Laminoglymmius ) oberthueri (Grouvelle) NEW COMBINATION
Figs. 61,72
Rhysodes oberthueri Grouvelle 1903: 1 18-119.
Omoglymmius ( Hemiglymmius ) oberthueri (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “Poulo-Pinang, Raffray” (MNHN)
Description. — Length 7.0 mm. Antennal Segment XI as wide as long, its apex rounded; basal setae numerous on
Segments V-X; head slightly longer than wide; clypeus continuous with median lobe; latter rhomboidal, its tip pointed;
clypeus and median lobe impunctate; anteriomedial margin of temporal lobe oblique; two medial angles, separated by
shallow but rather long emargination; each medial angle narrowly separated from that on opposite temporal lobe; medial
angles and emargination within well-marked translucent area; temporal lobe with nine or 10 coarse punctures in
posteriolateral third; posteriolateral margin of temporal lobe evenly rounded; temporal lobe slightly overhanging postorbit;
latter glabrous, impunctate; postorbital tubercle rather large, about 0.5 as long and 0.5 as deep as eye; its dorsal margin on
level with middle of eye.
Revision of Omoglymmius Ganglbauer
163
Pronotum relatively short, broad, length/greatest width 1.13; widest slightly anterior to middle; sides curved, apex
strongly narrowed, base moderately narrowed; sides curved, sides oblique just anterior to obtuse hind angle, paramedian
grooves deep, rather narrow; outer carina scarcely narrower than inner one; outer carina with seven to 1 2 coarse,
scattered punctures; inner carina with three or four coarse punctures near middle of its length, otherwise impunctate.
Elytra relatively short, broad; striae deep; Intervals III, V slightly broader, more elevated than others anteriorly;
strial punctures coarse, each more than half as wide as interval; Stria IV with one seta in apex; apical stride with one
seta, Stria VII with several setae near apex; female with very deep, large lateral pits on Sternum IV; each pit as long as
Sternum IV; pits separated medially by less than twice the width of one of them; female with distinct ventral tooth on
anterior femur; male unknown.
The rhomboid median lobe and the presence of two medial angles make this species most
like O.actae. It differs from the latter species in having a well-marked medial translucent area
on each temporal lobe, as well as in having the medial angles of each lobe more separated from
one another.
Omoglymmius (Laminoglymmius) trisinuatus new species
Figs. 62, 73
Type material. - HOLOTYPE male, labelled: “Bangung” (MNHN). This is possibly
Bangoen (modern spelling “Bangun”). east of Pematangsiantar, in northern Sumatra.
Description. — Length 6.9 mm. Antennal Segment XI wider than long, its apex bluntly rounded; basal setae
numerous on Segments V-X; head longer than wide; clypeus continuous with median lobe; clypeus, median lobe
impunctate; median lobe elongate, narrow, its tip translucent; anteriomedial margin of temporal lobe curved; temporal lobe
with three very small medial angles, separated by very shallow sinuations; first pair well separated; second pair very
narrowly separated; third pair almost contiguous; all angles and adjacent margin translucent; posteriomedial margin
slightly oblique; temporal lobe convex, shining, with three coarse punctures near lateral margin; postorbit pollinose;
postorbital tubercle 0.3 as deep and 0.3 as long as eye, its dorsal margin on level with lower third of eye; postorbital
tubercles visible in dorsal view, width across them slightly less than width across eyes.
Pronotum short, broad, length/greatest width 1.05; widest slightly anterior to middle, apex very strongly narrowed;
base slightly narrowed; lateral margin scarcely sinuate anterior to obtuse hind angles; paramedian grooves deep, about 0.5
as wide as inner carina at middle; outer carina only slightly narrower than inner one at middle; inner carina impunctate;
outer carina with five to seven punctures near its lateral margin; marginal groove narrow.
Elytra relatively short, broad; strial punctures small, well separated from one another, less than 0.3 as wide as an
interval; Interval V slightly elevated anteriorly; elytron with setae near apex of Stria VII, otherwise without setae; male
with faint lateral pit in Sternum IV; female unknown.
Male with small ventral tooth on anterior femur; middle calcar very small, obtuse; hind tibia with small obtuse calcar
at level of spur.
Three medial angles of the temporal lobe separate this species from all others except for O.
rugosus, from which it differs in lacking setae on the temporal lobe and pronotum, as well as
many other characters. Shape and punctation of the pronotum, narrow median lobe, and
secondary sexual characters of the male indicate that its true relationships are with O. gorgo,
which differs in having the postorbital lobes greatly enlarged.
Omoglymmius ( Laminoglymmius ) gorgo new species
Figs. 63, 74
Type material. - HOLOTYPE male, labelled: “SUMATRA: Siantar, NGSSI exp, 1937,
Mann” (NMNH). PARATYPES one male, one female, labelled “Bedagei, int., Sumatra’s
O.K. 600', 2de de Sem, '89, coll. I. Z. Kannegieter” (MNHN).
Description. — Length 6. 0-8.0 mm. Antennal Segment XI as wide as long, its apex rounded; basal setae numerous
on Segments V-X; head longer than wide (if postorbital tubercles are not considered); clypeus continuous with median
lobe; latter very narrow, its tip narrowed, rounded, opaque; clypeus finely punctate; median lobe impunctate; anteriomedial
margin of temporal lobe curved; temporal lobe with two small medial angles, separated by shallow emargination; first
medial angles slightly separated; second medial angles contiguous; first medial angles translucent; second medial angles
less distinctly so; temporal lobe convex, impunctate; postorbit pollinose; postorbital tubercle very large, longer than eye, 0.6
Quaest. Ent., 1982, 18 (1-4)
164
Bell and Bell
as deep as eye, its upper and lower margins parallel, its apex subtruncate; width across postorbital tubercles much
greater than that across eyes.
Pronotum short, relatively broad, length/greatest width 1.04; widest near middle, apex very markedly narrowed;
base moderately narrowed, sides markedly curved; lateral margin distinctly sinuate anterior to obtuse hind angle;
paramedian grooves deep, about 0.5 as wide as inner carina at middle; outer carina only slightly narrower than inner
one at middle; inner carina impunctate; outer carina with five to seven punctures near its lateral margin; marginal
groove narrow.
Elytra relatively short, broad; strial punctures small, well separated from one another, less than 0.3 as wide as one
interval; Interval V slightly elevated anteriorly; several setae near apex of Stria VII, elytron otherwise without setae;
female with small but distinct lateral pit on abdominal Sternum IV; male without lateral pit; male with small ventral
tooth on anterior femur; female with very small tooth in this location; middle calcar scarcely evident; hind calcar very
small but distinct (smaller than in O. trisinuatus).
The grotesque enlargement of the postorbital tubercles easily separates this species from
other Laminoglymmius .
Subgenus Navitia Bell and Bell 1978
Type species. - Rhysodes intrusus Grouvelle 1903
Description. — Basal setae of antennae entirely absent; antennal stylet present, conical, acute; clypeal setae absent;
eye large, normal, cornea facetted; frontal grooves very shallow; frontal space wide, crescentic; medial angles of temporal
lobes simple; translucent areas absent from temporal lobes; one temporal seta on each lobe; postorbital tubercle absent;
gular grooves each containing an enlarged pit.
Pronotal carinae impunctate except for a few punctures at base of outer carina; inner and outer carinae equal in width;
marginal groove of pronotum abbreviated posteriorly, replaced there by row of punctures; apex of elytral Stria VII not
impressed; most posterior puncture of Stria III enlarged; Stria II with one seta at apex in most specimens (absent in some
specimens); Stria IV with three or four setae, distributed along its entire length; subapical stride with 2 setae; Stria VII
with three to five setae near apex; lateral pits present on abdominal Sternum IV in both sexes; spur of middle tibia curved
anteriorly; punctures of abdominal Sterna III-V each aligned in single transverse row.
This small and distinctive subgenus combines the characters of several other subgenera. An
antennal stylet and the simple medial angles of the temporal lobes suggest Hemiglymmius. The
latter subgenus differs in having well-developed basal setae on the antennae and deep frontal
grooves. Alignment of abdominal punctures in a single row on each sternum is suggestive of
Orthoglymmius , but in the latter subgenus, the middle tibial spur is straight, and clypeal setae
are present. The habitus is similar to Omoglymmius s. str ., but the latter subgenus lacks the
antennal stylet and has deeper frontal grooves.
The frontal grooves are shallower than in any other members of Subtribe Omoglymmiina. In
O. intrusus , they are glabrous and scarcely visible, giving a superficial resemblance to Dhysores
and related genera. In Navitia , however, the frontal space is crescentic, while in Subtribe
Dhysorina it is small and round. In O. stylatus, the shallowness of the frontal grooves is less
conspicuous because they are pollinose.
Navitia is known from Fiji and the New Hebrides.
KEY TO SPECIES
1 Marginal groove confined to anterior 0.85 of pronotum, followed posteriorly by
three to four very crowded punctures; pronotal epipleura with line of punctures
ventrad to marginal groove; frontal grooves pollinose
O. stylatus new species, p. 165
V Marginal groove confined to anterior 0.50-0.66 of pronotum, followed
posteriorly by five widely separated punctures; pronotal epipleura without line
of punctures; frontal grooves glabrous, scarcely evident
O. intrusus (Grouvelle), p. 166
Revision of Omoglymmius Ganglbauer
165
Plate 6. Figures 75, 76, 79, 80, Subgenus Navitia-, Figures 77, 83 Subgenus Indoglymmius-, Figures 78, 81,82, Subgenus
Nitiglymmius; Figs. 75-78, Head and pronotum, dorsal aspect; Fig. 75, Omoglymmius (Navitia) stylatus new species;
Fig. 76, O. (N.) intrusus (Grouvelle); Fig. 77, Omoglymmius (Indoglymmius) lineatus (Grouvelle); Fig. 78, O.
( Nitiglymmius ) semioculatus new species; Figs. 79-80, Prothorax, lateral aspect; Fig. 79, O. ( Navitia ) intrusus
(Grouvelle); Fig. 80, O. (TV.) stylatus new species; Fig. 81, Hind tibia, male, O. ( Nitiglymmius ) semioculatus ; new species;
Figs. 82-83, Head, lateral aspect; Fig. 82, O. ( Nitiglymmius ) semioculatus new species; Fig. 83, O. (Indoglymmius)
lineatus (Grouvelle);
Omoglymmius ( Navitia ) stylatus new species
Figs. 75, 80
Type material. - HOLOTYPE male, labelled: “NEW HEBRIBES.Malekula, XII-1929,
coll. L. E. Cheesman, BM 1930-393 (1669)” (BMNH). PARATYPES two females, N. E.
Malekula, 1-1930, coll. L. E. Cheesman, BM-1930-178 (BMNH); one female, Malekula:
Ounua, IV-V-1929, coll. L. E. Cheesman, BM 1929-371 (531) (BMNH); one female,
Aoba:Banka Harijitoa, above Dundy, elev. 1500-2000 ft., IX-8-1958, collector not specified
(CNHM).
Quaest. Ent., 1982, 18 (1-4)
166
Bell and Bell
Description. — Length 5. 0-6. 2 mm. Head cordate, slightly longer than broad; temporal lobes distinctly flattened;
frontal grooves slightly impressed, with fine pollinosity; temporal lobe with few coarse punctures in posteriolateral third,
near temporal setae; orbital groove relatively well-developed, extending posteriorly to or beyond middle of temporal lobe.
Pronotum markedly varied in proportions, widest anterior to middle; lateral margin with distinct sinuation anterior to
hind angles in some specimens, in others sinuation slight or absent; marginal groove of pronotum entire except for basal
0.16, replaced there by very crowded row of three or four punctures; pronotal epipleuron with row of punctures ventrad to
marginal groove (Fig. 80).
Elytron with most posterior puncture of Stria III moderately enlarged; punctures of abdominal Sternum V not
confluent laterally; male with ventral tooth on anterior femur; calcar of middle tibia obsolete; hind calcar small, triangular,
its tip slightly obtuse, distinctly proximad to base of spur.
This species is easily distinguished from O. intrusus by the more complete marginal groove
of the pronotum, line of punctures on the pronotal epipleura, and deeper, pollinose frontal
grooves. Grouvelle (1903) mentioned a specimen tentatively identified as R. intrusus from the
New Hebrides. We have not located this specimen, but we suspect that it really represents O.
stylatus.
Omoglymmius (Navitia) intrusus (Grouvelle)
Figs. 76, 79
Rhysodes intrusus Grouvelle 1903:1 12-113.
Omoglymmius ( Navitia ) intrusus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) female, labelled: FIJI “Viti,” no collector
or date (MNHN). PARALECTOTYPE: Grouvelle (1903) mentioned a second specimen in the
Oberthiir collection. It may still exist among Oberthiir material recently rediscovered in Paris.
Description. — Length 5. 0-6.0 mm. Head broader, more convex than in O. stylatus, temporal lobes not flattened;
frontal grooves scarcely impressed, shining, glabrous; temporal lobe entirely without coarse punctures, but with sparse
minute ones; orbital grooves obsolete.
Pronotum widest near middle; sinuation anterior to hind angle shallow but distinct; marginal groove distinct in anterior
0.50 to 0.67, replaced posteriorly by row of five widely spaced punctures; pronotal epipleura without row of punctures (Fig.
79).
Elytron with most posterior puncture of Stria III greatly enlarged, so that Stria IV also seems to lead to it; punctures
of Sternum V confluent near lateral margin; male unknown.
The frontal grooves are virtually absent in this species. Glabrous frontal grooves, the more
abbreviated marginal groove on the pronotum, absence of punctures from the pronotal
epipleura, and confluent punctures of Sternum V are the most conspicuous differences from O.
stylatus. Shape of the pronotum shows an unusual amount of variation in both species.
O. intrusus is known only from the Fijian islands of Viti Levu and Ova Lau. In addition to
the lectotype, we have examined the following specimens; one female, labelled “Viti Levu, Naivithuia,
Taileru-Fiji, V 1 1 1- 1 6- 1 937, coll. J. M. Valentine” (BPM); one female, labelled “Ova Lau, Fiji, Draiba Trail, V 1 1-38,
rotten log, coll. E. C. Zimmerman” (BPBM).
Subgenus Caeconavitia new subgenus
Type species. - Omoglymmius (Nitiglymmius) zimmermani Bell and Bell 1978:77.
Description. — Basal setae of antenna entirely absent; antennal stylet present, conical, obtuse; clypeal setae absent;
eye small, only slightly visible in dorsal aspect; depth of eye about 0.33 in depth of head; facetting of cornea much reduced,
barely visible at high magnification; frontal grooves deep, narrow; medial angles of temporal lobes simple; translucent
areas absent from temporal lobes; temporal setae absent; postorbital tubercle absent; each gular groove with enlarged pit.
Marginal groove of pronotum entire; elytral striae distinctly impressed; abdominal Sterna III-V each with transverse
row of punctures; Sternum IV with deep and Sternum V with shallow lateral pit in both sexes.
This subgenus contains one species, from Fiji. We originally assigned it to Subgenus
Nitiglymmius. Futher study of the genus has led us to attribute greater importance to presence
Revision of Omoglymmius Ganglbauer
167
or absence of the antennal stylet, and has convinced us that O. zimmermani is more closely
related to Navitia than to the remaining species of Nitiglymmius. Characters shared between
Navitia and Caeconavitia include an antennal stylet, enlarged gular pits, and alignment of
punctures of abdominal sterna into one transverse line per sternum. On the other hand, O.
zimmermani cannot simply be regarded as a species of Navitia with reduced eyes, as it lacks
two specializations typical of Navitia : obsolete frontal grooves and an incomplete marginal
groove on the pronotum. We therefore consider it best to isolate this species in a subgenus of its
own.
Omoglymmius ( Caeconavitia ) zimmermani (Bell and Bell) NEW COMBINATION
Omoglymmius ( Nitiglymmius ) zimmermani Bell and Bell, 1978:77.
Type material. - Part 1:77.
Description. - Part 1:77.
Indoglymmius new subgenus
Type species. - Rhysodes lineatus Grouvelle 1908
Description. — Segment XI of antenna obtuse, without stylet; basal setae of antenna absent; clypeal setae absent;
medial margin of temporal lobe with pale translucent area; medial angles simple; frontal grooves straight and linear;
temporal lobe with line of punctures along orbital groove, otherwise impunctate; temporal setae absent; frontal grooves
linear; frontal space very small; postorbital tubercle absent; median groove of pronotum coarsely conspicuously punctate;
punctures pollinose, groove otherwise glabrous; paramedian grooves coarsely punctate, but inconspicuously so, punctures
partly hidden by pollinosity; pronotal carinae impunctate; borders between carinae and grooves somewhat irregular, with
carinae encroached by punctures of grooves; elytral striae scarcely impressed; Stria IV with one seta at apex; apical stride
with one seta; apex of Stria VII with several setae; abdominal sterna with scattered punctures; both sexes with deep lateral
pits on abdominal Sternum IV, and with shallow ones on Sternum V; ventral tooth on anterior femora in both sexes.
The only species in this subgenus resembles Laminoglymmius in having a conspicuous
translucent area near the medial angle of the temporal lobe, but differs in lacking basal setae
on the antennae. The simple medial angles separate it from all Laminoglymmius except for O.
(L) inaequalis. The very narrow outer pronotal carinae separate the latter species from
Indoglymmius. The translucent area is the most conspicuous character distinguishing
Indoglymmius from Omoglymmius s. str.
Omoglymmius ( Indoglymmius ) lineatus (Grouvelle) NEW COMBINATION
Figs. 77, 83
Rhysodes lineatus Grouvelle 1908:319-320.
Omoglymmius (s. str.) lineatus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: “(INDIA),
Chambaganor, Madura, 48” (MNHN) PARALECTOTYPES 17 specimens, sexes not
recorded, same data as lectotype (MNHN; one male (labelled as “cotype”) labelled: “INDIA,
Shembaganur, Madura, Andrews Bequest BM 1922-22” (BMNH) According to Grouvelle, the
type series was collected by R. P. Dubreuil.
Description. — Length 5. 6-6. 2 mm. Antennal Segment XI conspicuously narrower than Segment X; head slightly
longer than wide; pronotum moderately elongate, length/greatest width 1.28; lateral margin distinctly sinuate anterior to
nearly rectangular hind angle; middle calcar of male minute; hind calcar small, triangular, its apex obtuse.
In addition to the type series, we have seen one male and one female, labelled “Key Inseln,
coll. Reithoffer” (BSL). This refers to the Kei Islands, west of New Guinea. It is highly
Quaest. Ent., 1982, 18 (1-4)
168
Bell and Bell
unlikely that the species occurs both in the Kei Islands and southern India. Probably either the
type series or the Basel specimens have incorrect locality data. Since there is no independent
record from either place, the true provenance of this species must be listed as uncertain.
Subgenus Nitiglymmius Bell and Bell 1978
Type species. - Omoglymmius ( Nitiglymmius ) fulgens Bell and Bell 1978
Description. — Antennal Segment XI without apical stylet; clypeal setae absent; frontal grooves deep but narrow to
linear; eye reduced, in profile view, entirely dorsad to middle of head; ommatidia not distinct; cornea not facetted; cornea
with pinkish tint; silvery-white reflecting disc medial to cornea and at angle to it; marginal grooves of pronotum fine, linear
entire or partly or completely effaced; elytral stria not impressed, but represented by rows of punctures; spur of middle
tibia with tip curved anteriorly; punctures of abdominal sterna scattered or in single transverse row on each segment;
lateral pits present on Sternum IV in both sexes (where known); ventral surface with bluish opalescence (such opalescence
on dorsal surface also in most species); hind wings vestigial.
The description has been modified from that in Part I in accordance with removal of O.
zimmermani to Subgenus Caeconavitia and addition of O. (N.) semioculatus new species from
Philippines. The range of Nitiglymmius as here limited is New Guinea, the Solomon Islands
and Siargao Island in the Philippines.
KEY TO SPECIES (Supersedes that in Part I: 76-77)
1 Elytron with longitudinal scarp present at base of Stria IV; elytral striae slightly
impressed, rather coarsely punctate; eye minute, less than 0.1 of depth of head.
O. semioculatus new species, p. 169
T Elytron without scarp at base of Stria IV; elytral stria not at all impressed, very
finely punctate; eye less minute, more than 0.1 of depth of head 2
2 (T) Pronotum relatively elongate, its margin evenly curved to apex, not constricted
to form “collar.”
O. greensladei Bell and Bell, (part I; 78)
2' Pronotum less elongate, its sides abruptly narrowed anteriorly, in form of
“collar” at apex 3
3 (20 Postorbital tubercles distinct 4
3' Postorbital tubercles absent 5
4 (3) Pronotum distinctly narrowed at apex; lateral margin of pronotum sinuate just
anterior to hind angles; posterior median pit large, as wide as basal impression .
O. hornabrooki Bell and Bell (Part I: 81)
4' Pronotum only slightly narrowed anteriorly, lateral margin not sinuate,
posterior median pit very small, much narrower than basal impression
O. offafinus Bell and Bell (Part I; 81)
5 (30 Marginal groove of pronotum present 6
5' Marginal groove absent O. lustrans Bell and Bell (Part I: 80)
6 (5) Frontal grooves very fine, linear; median lobe long, acutely pointed, extending
posterior to middle of temporal lobes O. fulgens Bell and Bell (Part 1:80)
6' Frontal grooves broader, not linear; median lobe short, obtusely pointed, not
extended to middle of temporal lobes O. toxopei Bell and Bell (Part I: 82)
Revision of Omoglymmius Ganglbauer
169
Omoglymmius ( Nitiglymmius ) semioculatus new species
Figs. 78,81,82
Type material. - HOLOTYPE male, labelled: “PHILIPPINEN: Mindanao, Dapa, leg.
Boettcher, 30-9-16” (NMHB). Despite the label, the locality is not on Mindanao, but on the
small island of Siargao, northeast of it.
Description. — Length 6.2 mm. Form elongate for genus; dorsal surface not opalescent; ventral surface scarcely
opalescent; head elongate, its lateral margin markedly sinuate anterior to eye; median lobe elongate, its apex rather acute,
opposite middle of temporal lobe; frontal grooves long, wider than in other members of subgenus; orbital groove absent;
frontal space larger than in other members of subgenus; medial angles rectangular, nearly contiguous; margin posterior to
medial angle sinuate; one temporal seta; temporal lobe with about six fine punctures, restricted to lateral region; eye
minute, oval, visible only in lateral view; postorbital tubercle absent; mentum punctate; gular region extensively pollinose.
Pronotum elongate; length/greatest width 1.35; anterior end in form of short, ill-defined “collar”; lateral margin
curved into “collar” anteriorly, slightly sinuate just anterior to hind angle; outer carina slightly narrower than inner one,
with about nine punctures scattered along its lateral margin; inner carina impunctate; median groove deep, anterior and
posterior median pits equal, moderately large; lateral margin of paramedian groove distinct, but medial margin sloped
gradually from inner carina; marginal grooves broader than in other members of subgenus, complete; epipleura
impunctate.
Elytra elongate, more parallel-sided than in other members of subgenus; elytron with short, longitudinal scarp at base
of Stria IV; elytral striae shallowly impressed, rather coarsely punctate; Stria IV with one seta near apex; Stria VII with
one seta near apex; elytron in profile with highest point at apical fifth, sloped rather abruptly posterior to this point;
metasternum with entire surface densely, coarsely, shallowly punctate; abdominal sterna similarly punctate; anterior
femur of male without ventral tooth; hind calcar of male large, its apex obtuse, raised above spur, margin between apex
and spur sinuate; margin proximal to calcar distinctly emarginate (Fig. 81).
This species differs from all other members of the subgenus in the scarp at the base of Stria
IV very reduced eyes, extensive pollinosity in the gula impressed, coarsely punctate elytral
striae, posteriorly displaced elytral “hump” and the coarsely punctate metasternum and
abdominal sterna. Form of the hind calcar is also distinctive.
Subgenus Orthoglymmius Bell and Bell 1978
Type species. - Rhysodes sulcicollis Lewis 1888
Description. — Antennal Segment XI without stylet; basal setae of antenna absent; clypeal setae present; frontal
grooves deep, broad; margins of temporal lobe oblique both medial to and lateral to occipital angle, posterior temporal lobe
is pointed; temporal setae absent; eye large, deeper than long; cornea facetted; ommatidia distinct; marginal groove of
pronotum complete; elytral setae relatively well-developed, with complete row on Stria IV, and in some species, with
complete rows in Striae II and VI as well; spur of middle tibia straight or curved; metasternal punctures limited to medial
band and lateral margin; abdominal Sterna 1 1- V each with single transverse row of very coarse punctures, or with
punctures coalesced to form transverse sulci; female with lateral pits better developed on Sternum V than on Sternum IV;
hind coxa with seta; Sternum VI with pair of setae in both sexes; males, as far as known, with ventral tooth on anterior
femur but without proximal tooth on anterior tibia; hind calcar, where known, bisinuate; hind wings and genitalia not
investigated.
The straight spur of the middle tibia and deepest lateral pits on the fifth, rather than the
fourth, abdominal sternum, distinguish this subgenus from Omoglymmius s. str.. Only a few
members of the latter subgenus have the punctures of each abdominal sternum in a single
transverse row, as is consistently true in Orthoglymmius. In addition, Orthoglymmius
specimens have clypeal setae and have the temporal lobes pointed posteriorly. The Subgenus
Orthoglymmius is confined to a part of the Oriental Region, where it is known from the
Himalaya, the Indochina Peninsula, the Ryukyu Islands, and southern Japan. It is apparently
absent from southern India and from Indonesia.
Phylogeny. - This subgenus is perhaps the most inadequately known major group of
Rhysodini. It consists of a complex of very similar and closely related species. Of the eight
species, both sexes are known in only two. Several of the species described by Grouvelle are
Quaest. Ent., 1982, 18 (1-4)
170
Bell and Bell
known only from type material, and are so similar to one another that it would be premature to
reach final conclusions about their validity. The group is so uniform that it is almost a waste of
time to speculate about interrelationships among the species. The group will need a futher
revision when adequate series of specimens are available and the genitalia can be studied.
KEY TO SPECIES
1
V
2 (1)
2'
3 (10
3'
4 (3)
4'
5 (40
5'
6 (50
6'
7 (60
7
Medial angles of temporal lobes well separated, obtuse; posterior margin of
temporal lobe without pilosity 2
Medial angles contiguous or almost so; posterior margin of temporal lobe
fringed with pilosity 3
Postorbital tubercle distinct, 0.33 as long and 0.33 as deep as eye, glabrous
except for tuft of pilosity at tip; tip of median lobe obtuse
O. sulcicollis (Lewis), p. 170
Postorbital tubercle reduced to minute vestige with small pilose tuft at its tip;
median lobe acute at tip O. microtis new species, p. 172
Postorbital tubercle pilose 4
Postorbital tubercle glabrous O.feae (Grouvelle), p. 172
Median groove closed anteriorly O. alticola (Grouvelle), p. 173
Median groove open anteriorly 5
Lateral margin of pronotum obtusely angulate at middle
O. longiceps (Grouvelle), p. 173
Lateral margin of pronotum evenly curved at middle 6
Pronotal margins nearly parallel; lateral margin scarcely sinuate anterior to
hind angle O. cavifrons (Grouvelle), p. 174
Pronotal margins more curved; pronotal base more distinctly narrowed; lateral
margin distinctly sinuate anterior to hind angle 7
Orbital groove relatively wide, extending posteriorly to level of middle of eye;
posteriomedial and posteriolateral margins of temporal lobe both distinctly
sinuate; occipital angle obtusely pointed; postorbital tubercle large, prominent in
dorsal view O. crenatus (Grouvelle), p. 175
Orbital groove narrow, short, ending at anterior 0.33 of eye; posteriomedial and
posteriolateral margins of temporal lobe oblique or very slightly sinuate;
occipital angle obtuse; postorbital tubercle smaller, barely visible in dorsal view.
O. coomani (Arrow), p. 1 76
Omoglymmius (Orthoglymmius) sulcicollis (Lewis)
Figs. 84, 92
Rhysodes sulcicollis Lewis 1 888:8 1 .
Omoglymmius (Orthoglymmius) sulcicollis (Lewis) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “JAPAN: Oyayama, Higo (Island of
Honshu), 1881, coll. G. Lewis 1910-320” (BMNH).
Description. — Length 6.0 mm. Antennal Segment XI as long as wide, its cone well-developed; clypeal setae
present; median lobe short, tapered posteriorly, its apex obtuse, located anterior to widest point of head; anteriomedial
margin of temporal lobe oblique, scarcely sinuate; medial angles obtuse, widely separated from one another; posteriomedial
Revision of Omoglymmius Ganglbauer
171
Plate 7. Figures 84-103, Subgenus Orthoglymmius. Figs. 84-91, Head and pronotum, dorsal aspect; Fig. 84,
Omoglymmius (Orthoglymmius) sulcicollis (Lewis); Fig. 85, O. (O.) microtis new species; Fig. 86, O. (O.) feae
(Grouvelle); Fig. 87, O. (O.) alticola (Grouvelle); Fig. 88, O. (O.) longiceps (Grouvelle); Fig. 89, O. (O.) cavifrons
(Grouvelle); Fig. 90, O. (O.) crenatus (Grouvelle); Fig. 91, O. (O.) coomani (Arrow); Figs. 92-99, Head, lateral aspect;
Fig. 92, O. (O.) sulcicollis (Lewis); Fig. 93, O. (O.) microtis new species; Fig. 94, O. (O.) feae (Grouvelle); Fig. 95, O. (O.)
alticola (Grouvelle); Fig. 96, O. (O.) longiceps (Grouvelle); Fig. 97, O. (O.) cavifrons (Grouvelle); Fig. 98, O. (O.) crenatus
(Grouvelle); Fig. 99, O. (O.) coomani (Arrow); Figs. 100-103, Hind tibia, male, apical portion; Fig. 100, O. (O.) longiceps
(Grouvelle); Fig. 101, O. (O.) cavifrons (Grouvelle); Fig. 102, O. (O.) crenatus (Grouvelle); Fig. 103, O. (O.) coomani
(Arrow);
Quaest. Ent., 1982, 18 (1-4)
172
Bell and Bell
and posteriolateral margins oblique, not margined by pilosity; occipital angle obtuse but distinct; orbital groove narrow,
ended just anterior to middle of eye; postorbital tubercle small, 0.33 as deep and 0.33 as long as eye, with tuft of long
pollinosity.
Pronotum moderately elongate, length/greatest width 1.25; widest at middle, sides strongly curved, narrowed to
apex, slightly narrowed to base; lateral margin with slight sinuosity anterior to hind angle; paramedian grooves
narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra rather short, broad for subgenus; striae subequal, intervals equally convex; base of Interval V not elevated;
Stria I with two to four setae near apex; Stria 11 with two setae near apex (in one specimen these setae are actually
slightly laterad to stria in medial part of Interval III); Stria IV with five or six setae in form of complete row; apical
stride with three setae; Stria VI with two setae near humerus; Stria VII with about seven setae near apex.
Female with shallow depression at middle of metasternum, abdominal Sterna II-V each
with a transverse row of punctures; male unknown.
Widely separated medial angles and absence of marginal pollinosity from the vicinity of the
occipital angles separate this species from all other except O. microtis. The postorbital tubercle
is much better developed than in the latter species.
In addition to the holotype, we have studied one female, from the collection of Dr. Sato,
labelled simply “Japan”.
Omoglymmius (Orthoglymmius) microtis new species
Figs. 85, 93
Type material. - HOLOTYPE female, labelled: “Loochoo, Is. Ishigaki, Takeda,
17-IV-1962, S. Tamai leg.” (SATO). Ishigaki is an island in the Sakashima Retto, the
southernmost group in the Ryukyu Islands.
Description. — Length 5.3 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present;
median lobe more elongate, more pointed than in O. sulcicollis, its tip even with greatest width of head; anteriomedial
margin of temporal lobe shallowly sinuate, nearly oblique; medial angles obtuse, widely separated; posteriomedial,
posteriolateral margins oblique, not margined by pilosity; occipital angle obtuse; orbital groove narrow, ended just anterior
to middle of eye; postorbital tubercle minute, less than 0.2 as long as eye, located immediately ventrad to tuft of short
pilosity.
Pronotum moderately elongate, length/greatest width 1.25; widest at middle, sides curved, narrowed to apex, slightly
narrowed to base, side slightly sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle
of pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra longer and more cylindrical than in O. sulcicollis ; stria subequal, intervals equally convex; base of Interval V
not at all elevated anteriorly; Stria I with three or four setae near apex; Stria II with one to two setae near apex; Stria IV
with six setae forming complete row; apical stride with two setae; Stria VI with two setae near humerus; Stria VII with
about six setae near apex.
Female with shallow depression at middle of metasternum; abdominal Sterna II-V each with transverse row of
punctures; male unknown.
This species differs from all other members of the subgenus in the extreme reduction of the
postorbital tubercle. It is otherwise very similar to O. sulcicollis of Japan. It contrasts sharply
with O. cavifrons of the nearby island of Taiwan in shape of the temporal lobes.
Omoglymmius ( Orthoglymmius ) feae (Grouvelle)
Figs. 86, 94
Rhy socles feae Grouvelle 1895b:76 1-762.
Omoglymmius ( Orthoglymmius ) feae (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “BURMA, Carin Cheba, 900-1100 m,
XII-88, coll. L. Fea” (MNHN). This may be a unique type, although the original dscription
implies the existence of a series of specimens.
Revision of Omoglymmius Ganglbauer
173
Description. — Length 6.9 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present;
median lobe tapered posteriorly, its tip obtuse; anteriomedial margin distinctly curved, emarginate; medial angles acute,
contiguous; posteriomedial and posteriolateral margins oblique, slightly sinuate, margined by pilosity; occipital angle
distinct; orbital groove narrow, ending anterior to middle of eye; postorbital tubercle large, 0.5 as long and 0.66 as deep as
eye, glabrous.
Pronotum moderately long, length/greatest width 1.22; widest near middle, strongly curved and narrowed to apex;
slightly to base; lateral margin slightly sinuate anterior to hind angle; paramedian grooves narrower than carinae at level
of middle of pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra moderately long; striae subequal, intervals equally convex; base of Interval V not at all elevated; Stria 1 with
four setae near apex; Stria II with three setae near apex; Stria IV with seven setae in form of complete row; apical striole
with one seta; Stria VI with two setae near humerus; Stria VII with six or seven setae near apex; female without impression
in metasternum; abdominal Sterna Il-V each with transverse row of punctures.
The single specimen assigned to this species differs from other members of the group in
having the postorbital tubercle glabrous.
Omoglymmius (Orthoglymmius) alticola (Grouvelle)
Figs. 87, 95
Rhysodes alticola Grouvelle 1913: 99-100.
Omoglymmius ( Orthoglymmius ) alticola (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE male, labelled: “Kobo, 400 feet, Abor Exped, 30-XI-ll,
Kemp” (NMHN). The Abor Hills are now in the State of Arunachal Pradesh, in northeastern
India.
Description. — Length 5.5 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present;
median lobe tapered posteriorly, its tip obtuse; anteriomedial margin strongly curved and emarginate; medial angles acute,
contiguous; posteriomedial and posteriolateral margins oblique; margined by pilosity; occipital angle obtuse; orbital groove
narrow, ended anterior to middle of eye; postorbital tubercle large, 0.5 as long and 0.67 as deep as eye; postorbital tubercle
and postorbit pilose.
Pronotum moderately long, length/greatest width 1.25, widest near middle, strongly curved and narrowed to apex,
moderately incurved and narrowed to base; lateral margin scarcely sinuate anterior to hind angle; paramedian grooves
narrower than outer carinae near middle; inner carinae relatively narrow, about 0.67 as wide as outer ones at middle;
median groove closed at anterior end; carinae impunctate; pronotal setae absent.
Elytra relatively long, narrow; stria subequal, intervals equally convex; Interval V not at all elevated at base; Stria 1
with four setae near apex; Stria III with three setae near apex; Stria IV with complete row of six or seven setae; medial
face of apical tubercle with one seta; apical striole with three setae; Stria VI with three setae near humerus; Stria VII with
six setae near apex, male with shallow median impression on metasternum and ventral tooth on anterior femur; middle
calcar small, acute; hind calcar bisinuate; female unknown.
The distinctive marks of this species are that the median groove is closed anteriorly and that
the inner carina is distinctly narrower than the outer one. The species most nearly sympatric is
O. crenatus. The latter species differs in having the median groove open anteriorly, the inner
carinae as broad as the outer ones, the orbital groove both wider and longer, and the occipital
angles more pointed.
Omoglymmius ( Orthoglymmius ) longiceps (Grouvelle)
Figs. 88, 96, 100
Rhysodes longiceps Grouvelle 1910: 324-325.
Omoglymmius (Orthoglymmius) longiceps (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE female, labelled: “BURMA,” without definite locality or
collector (MNHN). According to the original description, there was a type series of four
specimens, but we have been unable to locate the other three.
Quaest. Ent., 1982, 18 (1-4)
174
Bell and Bell
Description. — Length 6-7 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present; head
relatively elongate; median lobe tapered posteriorly, its tip obtuse; anteriomedial margin curved, emarginate; medial angles
acute, nearly contiguous; posteriomedial and posteriolateral margins oblique, slightly sinuate, margined by pilosity;
occipital angle distinct; orbital groove narrow, ended anterior to middle of eye; postorbital tubercle large, 0.67 as long and
0.67 as deep as eye, prominent in dorsal view; postorbital tubercle and postorbit pilose.
Pronotum moderately long, length/greatest width 1.21; widest near middle, margin in form of obtuse angle at widest
point; sides oblique and slightly curved both anterior and posterior to angle; apex strongly narrowed; base distinctly
narrowed; margin not sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of
pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra moderately long; striae subequal; intervals equally convex; base of Interval V not elevated; Stria I with five
setae near apex; Stria II with three setae near apex; Stria IV with five or six setae in complete row; medial face of apical
tubercle with one seta; apical striole with two setae; Stria VI with one or two setae near humerus; Stria VII with about five
setae near apex; both sexes with lateral pit in abdominal Sternum V; male with shallow concavity at middle of
metasternum; abdominal Sterna 1 1- V each with transverse row of punctures.
Male with ventral tooth on anterior femur; middle calcar acute; hind calcar with obtuse triangular point and
rectangular “shoulder” proximad to it, separated from point by shallow emargination (Fig. 100).
In addition to the holotype, we have studied one male specimen, labelled “Carin Ghecu
1300-1400 m, L. Fea, II-III 88, coll. Jul. Moser” (MNHB). This specimen bears the label “/?.
feae Grouv.” but resembles the holotype of R. longiceps in having the postorbital tubercles
pollinose, and in the shape of the pronotum. In our opinion, it represents the male of R.
longiceps. A female, labelled: “Carin-Ghecu 300-1400 m.; L. Fea, V-88” (GEN), and a male,
labelled: “Tenasserim, Meetan, Fea, April, 1887” (GEN) also appear to be O. longiceps. O.
coomani is closely related, but appears to differ consistently in the narrower pronotum without
distinct lateral angles, the shorter, more rounded occipital angles, and in the somewhat more
elongate median lobe.
Omoglymmius (Orthoglymmius) cavifrons (Grouvelle)
Figs. 89, 97, 101
Rhysodes cavifrons Grouvelle 1914: 33-34.
Omoglymmius ( Orthoglymmius ) cavifrons (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) sex not recorded, labelled: “Kosempo.
Formosa, coll. H. A. Sauter, 191 1” (MNHN). PARALECTOTYPES: The original description
mentions a type series of four specimens. We have seen one specimen from the MNHB
collection labelled “Kosempo, 11-08, Sauter, S.V.” which is a probable paralectotype. The
remaining two specimens, if still in existence, would also be paralectotypes.
Description. - Length 5-6 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present;
median lobe short, its tip obtuse, anterior to middle of eye; anteriomedial margin distinctly curved, sinuate; medial angles
contiguous or nearly so, acute; posteriomedial and posteriolateral margins oblique, margined with pilosity; occipital angle
nearly rectangular; orbital groove rather broad, ended posterior to middle of eye; postorbital tubercle rather large, 0.67 as
deep and 0.67 as long as eye; postorbital tubercle and postorbit pilose.
Pronotum rather elongate, length/greatest width 1.26; widest near middle, lateral margins curved and convergent
anteriorly; apex strongly narrowed; margins straight, almost parallel posteriorly, base only slightly narrowed; margin not
sinuate anterior to hind angle; paramedian grooves narrower than carinae at level of middle of pronotum; carinae broad,
impunctate, pronotal setae absent.
Elytra rather long, narrow; striae subequal; intervals equally convex; base of Interval V not at all elevated; Stria I with
three or four seta near apex; Stria II with two setae near apex; Stria IV with six setae in form of complete row; medial side
of apical tubercle with one seta; apical striole with two or three setae; Stria VI with three setae near humerus; Stria VII
with about five setae near apex.
Abdominal Sterna II-V each with transverse row of punctures; both sexes with lateral pits in Sternum V; male with
ventral tooth on anterior femur; middle calcar acute; hind calcar with two obtuse angles, one above the other, separated by
shallow sinuation (Fig. 101).
This species differs from O. coomani and other species from the mainland of Asia in having
the pronotum longer and narrower, with the lateral margins straighter and more nearly
Revision of Omoglymmius Ganglbauer
175
parallel. It differs sharply from O. microtis of the Ryukyu Islands in having the postorbital
tubercles much larger and more strongly pilose, and in having the medial angles of the temporal
lobes contiguous or nearly so.
Range. - O. cavifrons is restricted to Taiwan. In addition to the lectotype, we have seen the
following specimens: three males, two females, Chipen Spa, Taitung Hsien, 3-6, V-1972, M. Sakai leg. (SATO); one
male, one female, Chipon, Formosa, 1 V- 1 8-32, coll. L. Gressitt (CAS); one male, one female, Taiwan: Hori (Puli), July,
1954, native collector (BPBM). Miwa (1931) cites specimens from Kosempo, Sokutsu and Chip-Chip.
Omoglymmius (Orthoglymmius) crenatus (Grouvelle)
Figs. 90,98, 102
Rhysodes crenatus Grouvelle 1903: 119.
Omoglymmius crenatus (Grouvelle) Bell and Bell 1977
Omoglymmius ( Orthoglymmius ) crenatus (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE: According to the original description, female, Bhoutan, coll.
Maria Borsti, in collection of R. Oberthiir. We have not been able to locate the type specimen,
but have described and illustrated a female specimen corresponding well to the original
description, from Balu-Jhura, Bhutan (Bell 1977).
Description. — Length 5.5 mm. Antennal Segment XI as long as wide, its cone distinct; head relatively elongate;
clypeal setae present; median lobe tapered posteriorly, its tip acute, slightly anterior to widest point of head; anteriomedial
margin distinctly curved, emarginate; medial angles acute, nearly contiguous; posteriomedial margin shallowly
emarginate; posteriolateral margin oblique; both margined with pilosity; occipital angle distinct; orbital groove relatively
broad, ended opposite middle of eye; postorbital tubercle 0.5 as deep and 0.5 as long as eye; postorbital tubercle and
postorbit pilose.
Pronotum moderately elongate, length/greatest width 1.23; widest slightly anterior to middle, markedly and very
abruptly narrowed at apex; slightly narrowed to base; sides oblique, not sinuate anterior to hind angles; paramedian
grooves narrower than carinae at level of middle of pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra relatively long, narrow; Striae subequal; intervals equally convex except for more elevated base of Interval V,
latter with fine microsculpture opaque; Stria I with four setae near apex; Stria II with three setae near apex; Stria IV with
complete row of four or five setae; one seta on medial face of apical tubercle; apical striole with four setae; Stria VI with
two setae near humerus; Stria VII with four setae near apex; both sexes with shallow median impression on metasternum;
abdominal Sterna 1 1 - V each with transverse row of punctures; both sexes with deep lateral pit in Sternum V; male with
ventral tooth on anterior femur; middle calcar acute; hind calcar with two very obtuse lobes, one proximad to the other,
separated by shallow sinuation (Fig. 102).
In this species, the base of Interval V is more distinctly elevated than in any other member
of the subgenus. O. coomani is similar but has the base of Interval V less elevated and without
microsculpture. It also has the lobes of the hind calcar less rounded and the postorbital
tubercles smaller. O. cavifrons resembles this species in having the orbital groove dilated, but
differs in having the base of Interval V not at all elevated, and shape of the frontal space and
pronotum different. O. alticola is possibly sympatric with O. crenatus in the eastern Himalaya.
O. alticola differs in having the orbital grooves fine, base of Interval V not at all elevated,
median groove of the pronotum closed anteriorly, and inner pronotal carinae conspicuously
narrower than the outer ones.
Range. - Foothills of the Himalayas in India, Bhutan and also in Laos. We have seen the
following additional Specimens:BHUTAN; one female, Balu-Jhura, 200 m, 28-4-72, coll. Basel Natural History
Museum Exped. (BSL); INDIA: one male, Haldwani Dist., Kumaon, coll. H. Gower Champion, British Museum,
1953-156 (BMNH); LAOS: two males, ULR farm, 3200', Xieng Khouang Prov., 2 mi. NW, Long Tieng, 18-Jan-1968
and 3-Mar-1968, coll. G. L. Peters (GLP).
Quaest. Ent., 1982, 18 (1-4)
176
Bell and Bell
Omoglymmius ( Orthoglymmius ) coomani (Arrow)
Figs. 91, 99, 103
Rhysodes coomani Arrow 1942: 180.
Omoglymmius ( Orthoglymmius ) coomani (Arrow) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: “Tonkin: Hoabinh, coll.
A. de Cooman, BM 1940-13” (BMNH). PARALECTOTYPES (all same collection data as
lectotype), one male labelled “co-type”, BM 1929-299; one male, same data as lectotype; one
male, BM 1929-299; one male, BM-1925-251 (all BMNH).
Description. — Length 5. 8-6. 5 mm. Antennal Segment XI as long as wide, its cone distinct; clypeal setae present;
median lobe tapered posteriorly; its tip obtusely pointed; anteriomedial margin of temporal lobe distinctly curved,
emarginate; medial angles acute, contiguous, posteriomedial and posteriolateral margins oblique, margined by pilosity;
occipital angles more obtuse than in O. crenatus, orbital groove narrow, short, ended anterior to middle of eye; postorbital
tubercle rather small, 0.33 as long and 0.33 as deep as eye; postorbital tubercle and postorbit pilose.
Pronotum moderately long, length/greatest width 1.23; widest near middle, curved to both base and apex; base more
narrowed than in related species; side not sinuate anterior to hind angle; paramedian grooves narrower than carinae at
level of middle of pronotum; carinae broad, impunctate; pronotal setae absent.
Elytra rather narrow; striae subequal; intervals equally convex except that base of Interval V is slightly elevated
(varied in degree of distinctness); Interval V not microsculptured near base; Stria I with four setae near apex; Stria II with
two or three near apex; Stria IV with complete row of seven to nine setae; medial face of apical tubercle with one to three
setae; apical striole with three to five setae; Stria VI with two to four setae near humerus; Stria VII with six to eight setae
near apex.
Both sexes with lateral pit in Sternum V; male with ventral tooth on anterior femur; middle calcar acute; hind calcar
with two lobes separated by sinuation; these lobes less rounded than in O. crenatus (Fig. 103).
This rather nondescript species varies in the degree to which the base of Interval V is
elevated. Those specimens in which the elevation is rather distinct are most liable to be
mistaken for O. crenatus. The latter species has a large postorbital tubercle, occipital angle
more distinct and orbital groove longer and more dilated. Those specimens with the base of
Interval V only slightly elevated are liable to be mistaken for O. longiceps or O. feae. In the
former species, the median lobe is shorter and more obtuse, while the latter species has a
glabrous postorbital tubercle.
Range. - O. coomani is known from Vietnam, Thailand and Java. In addition to the type
series, we have seen the following specimens: java: one female, Mt. Djampangs (MNHN); Thailand:
two females, Khao-Yai Nat. Pk., 750 m, VI 1-26-62, coll. E. S. Ross, D. Q. Cavagnaro (CAS); two males, E. slope of Doi
Sutep, 875-950 m, 1 5- V 1 1- 1 962, coll. E. S. Ross and D. Q. Cavagnaro (CAS); one male, “SIAM”, Mouhot, colln. G.
Lewis, 1901-31 (BMNH). VIETNAM: one female, Hoabinh, Tonkin, labelled with the manuscript name “R. tonkinensis”
in Grouvelle's handwriting (MNHN); one female, 30 km NW of Pleiku, elevation 300 m, 10-V-1960, coll. L. W. Quate
(BPBM); one female, nr. Tam-Dao, 900 m (N. Vietnam), 30-X-1963. coll. Kabakov (LEN); one male, five females,
Tam-Dao, Son-Zuong, 200-300 m, 31-1. 1-2, 20-11, 24-11, 1962, coll. Kabakov (LEN); two males, one female, NE
Thai-Nguen, 1 9-X 1 1 - 1 962, coll. Kabakov (LEN); three males, three females, 50 km No. Thai-Nguen, 300-400 m, 2-1 1 1,
3-3, 5-2, 19-12, 1963, coll. Kabakov (LEN); one male, NE Thai-Nguen, 1 1-1-1964, coll. Kabakov (LEN); one female,
Tong-Hoa Phu (MNHN).
Subgenus Carinoglymmius new subgenus
Type species. - Rhysodes carinatus Grouvelle 1903
Description. — Antennal Segment XI without stylet; basal setae of antenna present or absent; clypeal setae absent;
frontal grooves deep, broad; margins of temporal lobe rounded posteriorly in one species, angulate posteriorly in other
species; temporal setae absent; eye large, deeper than long; cornea facetted; ommatidia distinct; marginal groove of
pronotum complete, dilated; outer carina of pronotum very narrow, linear; elytral setae relatively well-developed, with
complete or partial rows in Striae 11, IV, VI; metasternal punctures evenly scattered; spur of middle tibia straight or nearly
so; abdominal Sterna 1 1 - V each with single transverse row of very coarse punctures; female with lateral pits better
developed on Sternum V than on Sternum IV; hind coxa with a seta; Sternum VI with pair of setae in both sexes; males, as
far as known, with ventral tooth on anterior femur but without proximal tooth on anterior tibia; hind wings and genitalia
Revision of Omoglymmius Ganglbauer
177
not investigated.
The straight spur of the middle tibia and presence of deepest pits on the fifth, rather than
fourth abdominal sternum separate this subgenus from all subgenera except Orthoglymmius.
Absence of clypeal setae and very narrow outer carina separate Carinoglymmius from the
latter subgenus.
The Subgenus Carinoglymmius is restricted to a small part of the Oriental Region, where it
is known from Borneo, Sumatra, Mentawei, and the Andaman Islands.
Phylogeny. - O. nicobarensis is clearly less specialized and more isolated than the other two
species, having the inner pronotal carina broad, pronotal setae absent, and the elytral stria
equally deep. O. carinatus and O. hexagonus form a pair of highly specialized and closely
related species, with the inner carinae linear, the outer carinae each with a row of punctures,
pronotal setae present, and elytral Stria II and IV much deeper than the others. In these two
species, both antennae and legs are much thickened in a manner reminiscent of some species of
Rhyzodiastes.
KEY TO SPECIES
1 Inner carina of pronotum more than twice width of outer carinae; outer carina
impunctate; elytral striae of equal depth; intervals not carinate; pronotal setae
absent O. nicobarensis (Grouvelle), p. 177
V Inner carina very narrow, subequal to outer carina; outer carina punctate; Stria
II, IV much deeper than other striae; some elytral intervals carinate; pronotum
with row of setae in marginal groove 2
2 (1) Posterior margin of temporal lobe obtusely pointed; pronotum clearly hexagonal
O. hexagonus (Grouvelle), p. 178
2‘ ' Posterior margin of temporal lobe rounded; sides of pronotum more evenly
curved O. carinatus (Grouvelle), p. 179
Omoglymmius ( Carinoglymmius ) nicobarensis (Grouvelle) NEW COMBINATION
Figs. 104, 108
Rhysodes nicobarensis Grouvelle 1895b: 762.
Type material. - HOLOTYPE female, labelled: “I. ANDAMAN” (MNHN). This
specimen is labelled as the type, and the original description, despite the specific name, gives
the locality as the Andaman Islands. A second specimen identified as this species by Grouvelle
(MNHN), is described above as Omoglymmius (Laminoglymmius) inaequalis new species. It
was collected in the Nicobar islands, and is responsible for the listing of R. nicobarensis from
the latter islands by Hincks (1950). Our placement of R. nicobarensis in subgenus
Hemiglymmius (1978) was based on this specimen and not the true R. nicobarensis.
Description. — Length 5.5 mm. Antennal Segment XI longer than wide, with well-developed cone; basal setae
absent; medial lobe of head pointed posteriorly; anteriomedial margin of temporal lobe shallowly sinuate; medial angles
obtuse, distinctly separated; orbital groove abbreviated; postorbital lobe small.
Pronotum slightly longer than in related species, length/greatest width 1.29; widest near middle, sides curved, apex
strongly narrowed, base only slightly narrowed; lateral margin not sinuate anterior to hind angle;xparamedian grooves
dilated, as broad as inner carinae, almost three times as broad as outer carina; marginal groove dilated; inner carina broad;
outer carina linear, only 0.33 as broad as inner one, impunctate; pronotum; without setae.
Elytra moderately elongate; striae of equal depth; Interval V slightly elevated at base; Stria I with one seta in apex;
Stria II with four near apex; Stria IV with complete series of approximately seven setae; apical striole with two setae; Stria
Quaest. Ent., 1982, 18 (1-4)
178
Bell and Bell
Plate 8. Figures 104-112, Subgenus Carinoglymmius. Figs. 104-106, Head and pronotum, dorsal aspect; Fig. 104,
Omoglymmius ( Carinoglymmius ) nicobarensis (Grouvelle); Fig. 105, O. (C.) carinatus (Grouvelle); Fig. 106, O. (C.)
hexagonus (Grouvelle); Fig. 107, Left elytron, dorsal aspect, O. (C.) hexagonus (Grouvelle); Figs. 108-110, Head, lateral
aspect; Fig. 108, O. (C.) nicobarensis (Grouvelle); Fig. 109, O. (C.) carinatus (Grouvelle); Fig. 110, O. (C.) hexagonus
(Grouvelle); Fig. Ill, Hind tibia, male, apical portion, O. (C.) hexagonus (Grouvelle); Fig. 112, Antennal Segments
IX-XI, O. (C.) hexagonus (Grouvelle);
VI with complete series of about six; Stria VII with four near apex; female with rather deep lateral pit in abdominal
Sternum V; male unknown.
This species is easily recognized within the subgenus by the combination of broad inner and
linear outer pronotal carinae. This feature is duplicated in other subgenera from the Andaman
and Nicobar Islands: Omoglymmius (Laminoglymmius) inaequalis Bell and Bell, and
Omoglymmius (s. str.) solitarius (Arrow), a striking and unexplained example of convergent
evolution.
Omoglymmius (Carinoglymmius) hexagonus (Grouvelle) NEW COMBINATION
Figs. 106, 107, 110-112
Rhysodes hexagonus Grouvelle 1903: 120-121.
Omoglymmius ( Orthoglymmius ) hexagonus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: (SUMATRA)
“Palembang”, collector and date unspecified (MNHN). PARALECTOTYPES one male,
Revision of Omoglymmius Ganglbauer
179
three females, same data (MNHN).
Description. — Length 5. 0-6. 2 mm. Antennae very short, thick; Segments IV-XI transverse, cylindrical (Fig, 1 12);
Segment XI with cone very low, apex almost flat; Segments II-IV each with broad apical pollinose ring; Segments V-XI
each with narrow basal pollinose ring; Segments V-X each with short basal setae without pollinose ring; head elongate;
median lobe nearly parallel-sided, its apex broadly rounded; frontal grooves dilated; anteriomedial margin very deeply
emarginate; medial angles obtuse, well separated; temporal and antennal lobes separated by broad pollinose space;
occipital angle obtuse but distinct; postorbital tubercle distinct though small.
Pronotum with length/greatest width 1.22, its outline hexagonal, sides nearly parallel in middle 0.33; oblique,
moderately convergent in basal third; oblique, strongly convergent in apical third; paramedian groove dilated, almost three
times as broad as either carina; marginal groove dilated, with approximately 10 marginal setae; setae clavate; inner and
outer carinae subequal, both very narrow, sublinear; inner one straight, impunctate; outer one slightly curved, with row of
punctures.
Elytra elongate, cylindrical; Intervals III and V carinate, convex; Intervals I, II and IV very low and narrow; all striae
coarsely punctate (Fig. 107); Stria II with seven to ten setae; Stria IV with eight to eleven setae (most posterior of these on
medial face of carinate Interval V); apical stride with three or four setae; Stria VI with eight or nine setae; Stria VII with
five to eight setae near apex; setae clavate; male with prominent ventral tooth on anterior femur; calcar of middle tibia
small, acute; calcar of hind tibia rather long, its margins nearly parallel, its tip obtusely rounded (Fig. 111).
This species and O. carinatus are very similar, having the inner pronotal carinae very
narrow, the outer carinae punctate, the marginal groove setose, and Intervals III and V of the
elytra elevated and carinate. In O. hexagonus the occipital angles are obtuse and distinct, while
in the latter species they are completely rounded.
Range. - Sumatra and Borneo. In addition to the type material, we have seen one specimen,
labelled:“W.SARAWAK, Mt. Matang, coll. G. E. Bryant, Dec. 1913” (BMNH). Possibly also Mentawei. We
have examined two specimens from Mentawei, labelled: “Mentawei:Sipora, Sereinu, V-VI-94, Modigliani” (GEN). One
of these appears to us to be O. hexagonus and the other O. carinatus. Both are labelled “ Rhysodes carinatus Grouvelle.”
The O. hexagonus has a “typus” label, and the R. carinatus a “syntype” label. It is possible that O. carinatus is not
distinct from O. hexagonus , but more material is needed to decide the question.
Omoglymmius (Carinoglymmius) carinatus (Grouvelle) NEW COMBINATION
Figs. 105, 109
Rhysodes carinatus Grouvelle 1903: 121.
Omoglymmius (Orthoglymmius) carinatus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) female, labelled: “BORNEO: River
Sambey pres Ngabang, 1897, coll. J. B. Ledru” (MNHN). PARALECTOTYPE male,
labelled: “Mentawei.Sipora, Sereinu, V-VI-94, Modigliani” (GEN) As mentioned above a
second male with the same label is O. hexagonus , if the latter is specifically distinct.
Description. — Length 6 mm. Antennae very short, thick; Segments IV-XI transverse, cylindrical; Segment XI
with cone very low, so apex is almost flat; Segments II-IV each with broad apical pollinose ring; Segments V-XI each with
narrow basal pollinose ring; Segments V-X each with short basal setae within pollinose ring.
Head broader than in O. hexagonus ; median lobe nearly parallel-sided, its apex rounded; temporal lobe separated from
antennal lobe by broad pollinose space; anteriomedial margin of temporal lobe deeply emarginate; medial angles rounded,
well separated; temporal lobe evenly rounded posteriorly, occipital angle indistinct; postorbital tubercle distinct though
small.
Pronotum with length/greatest width 1.21; widest near middle, narrowed to both base and apex; sides more evenly
curved than in O. hexagonus ; outline not clearly hexagonal; paramedian groove dilated, almost three times as broad as
either carina; marginal groove dilated, with seven to nine setae; inner and outer carinae subequal, both very narrow; inner
ones straight, impunctate; outer one more curved than in O. hexagonus , with a row of punctures which extends almost to
anterior end.
Elytra elongate, cylindrical; Intervals III, V carinate, convex; Intervals 1, II and IV very low and narrow; all striae
coarsely punctate; Stria II with seven or eight setae; Stria IV with 10-13 setae (the most posterior of these are located on
the medial face of the carinate Interval V); apical stride with three or four setae; Stria VI with five or six setae; Stria VII
with 6 setae in apex; female with moderately deep lateral pit in abdominal Sternum V; male with legs as in O. hexagonus.
This species is close to O. hexagonus , but differs in having a wider head, temporal lobes
rounded posteriorly, and the sides of the pronotum not distinctly hexagonal.
Quaest. Ent., 1982, 18 (1-4)
180
Bell and Bell
Subgenus Omoglymmius sensu stricto Ganglbauer 1892
Type species. - Rhysodes germari Ganglbauer 1892
Description. — Antennal Segment XI without stylet; basal setae of antennae absent; clypeal setae absent; frontal
grooves deep, varied in width; margin of temporal lobe rounded lateral to occipital angle in all species except O. germari ;
temporal setae one to four or none; eye deeper than long, in most species fully developed, more than 0.5 of depth of head; in
few species, reduced in size, but not less than 0.33 of depth of head, and extended well below middle of side of head; cornea
facetted, transparent, colorless; marginal groove of pronotum complete; setae of pronotum and elytra varied in
development; spur of middle tibia curved anteriorly; punctures of abdominal sterna scattered in most species, forming
single transverse row in a few species; lateral pits on abdominal Sternum IV in female, and at least suggested in male;
ventral surface in most species without opalescent sheen; hind wings fully developed in most species (but probably reduced
in those species with reduced eyes).
This subgenus is most likely to be confused with Orthoglymmius and Nitiglymmius.
Members of the former subgenus have clypeal setae, and have the lateral pits of the abdomen
best developed on Sternum V. They also have the posteriolateral margin of the temporal lobe
oblique, and the punctures of the abdomen arranged in a single transverse row on each sternum.
Omoglymmius (s. str.) germari has the temporal lobes shaped somewhat like those of
Orthoglymmius but has the sternal punctures scattered. A few species of Omoglymmius s. str.
have the sternal punctures as in Orthoglymmius , but have the temporal lobes rounded laterally.
A few species of Omoglymmius s. str. approach Nitiglymmius in having the eyes reduced,
but the process has not gone as far, and the cornea is facetted and transparent, while the eye is
0.40 or more of the depth of the head and extends well below the middle in lateral view.
The Subgenus Omoglymmius s. str. ranges from the Solomon Islands and the Carolines in
the east to Europe in the west. There is only one species in Europe, two in India, and one in the
Ryukyu Islands. There are possibly two species in northern Australia. Rhysodes
ichthyocephalus Lea appears, on the basis of an excellent photograph of the type, to belong to
this subgenus, and B. P. Moore has another, undescribed Australian species. As we have not
studied these species, they have not been included in our keys.
Phylogeny. - This subgenus contains more than 70 species which are very uniform in
external appearance. The few characters of diagnostic value occur in many combinations, and
some species widely separated localities are superficially very nearly alike. A few species, such
as O. germari , are obviously somewhat isolated from the remainder. A few others, such as O.
duplex and O. quadruplex, show evidence of close relationship with one another. Beyond these
observations, attempts to construct a phylogeny would be highly speculative on the basis of
present data. In the future, an analysis based on additional characters, such as male genitalia,
would be more profitable.
The key to species is of necessity very long and cumbersome. Therefore, we have added
several regional keys.
KEY TO SPECIES (WORLD)
1 Posteriolateral margin of temporal lobe oblique, meeting posteriomedial margin
at prominent occipital angle; posteriomedial margin also oblique, posterior part
of temporal lobe nearly rectangular O. germari (Ganglbauer), p. 194
V Posteriolateral margin of temporal lobe rounded; occipital angle not prominent;
posteriomedial margin convex, oblique, or emarginate 2
2 (E) Antennal lobe, postantennal area densely microsculptured, not separated by
antennal groove O. continuus new species, p. 226
Revision of Omoglymmius Ganglbauer
181
2' Antennal lobe glabrous, raised above level of postantennal area, or else
separated from latter by distinct antennal groove
3 (2') Inner carina distinctly narrower than outer carina at middle; pronotal grooves
coarsely, densely punctate O. malabaricus (Arrow), p. 207
3' Inner carina equal to or wider than outer carina at middle; pronotal grooves
impunctate or sparsely punctate
4 (30 Temporal lobe nearly circular; frontal space very small, longer than wide;
median lobe short, rhomboid; carinae of pronotum only moderately narrowed at
base
4' Temporal lobe reniform or oblique; frontal space larger, in most species wider
than long; median lobe either not rhomboid, or, if rhomboid, elongate; inner
carina either pointed posteriorly or else constricted just anterior to base, with
extreme base widened
5 (4) Medial angles of temporal lobes acute, contiguous; Stria IV with three to five
setae along its length; total length 5. 1-5.2 O. sakuraii (Nakane), p. 207
5' Medial angles of temporal lobes obtuse, slightly separated; Stria IV with one or
two setae near apex; total length 6. 1-7.1 O. laticeps Bell, p. 206
6 (4') Postorbital, suborbital tubercles absent
6' Either postorbital or suborbital tubercle present
7 (6') Fourth interval in form of sharp raised carina
O. bicarinatus new species, p. 230
T Fourth interval flat or somewhat elevated, but not carinate
8 (70 Pronotum with two or three marginal setae, inserted in prominent punctures
O. bucculatus (Arrow), p. 229
8' Pronotum without marginal setae
9 (80 Outer carina at middle equal to or only slightly narrower than inner carina, 0.66
or more of width of inner carina
9' Outer carina at middle 0.5 or less of width of inner carina
10 (9) Elytron with short pollinose longitudinal scarp at base of Stria IV, or with
glabrous scarp ( O . summissus), or with pollinose spot but with scarp indistinct
( O . politus)
10' Elytron without longitudinal scarp or isolated pollinose spot at base of Stria IV
11 (10) Either or both pairs of pronotal carinae with six or more punctures (in most
species with many more)
IF Pronotal carinae without punctures, or with one or two punctures on inner
carina or with one to five on outer carina
12(11) Inner carina with many punctures {very fine in O. vadosus)
12' Inner carina with none to two punctures
13 (12) Abdominal punctures indistinct, limited to lateral bands of pollinosity on Sterna
III- V ; elytral punctures shallow, scarcely pollinose
O. vadosus new species, p. 227
13' Punctures of abdominal sterna distinct, separated or barely coalescent laterally;
elytral striae with distinct, pollinose punctures
14 (13') Median lobe of head narrow, its margins only slightly curved; outer carina of
pronotum broadest at anterior margin, slightly flattened anteriorly; outer carina
with lateral margin not sinuate anterior to hind angle
O. summissus new species, p. 216
3
4
5
6
7
63
9
10
60
11
45
12
43
13
30
14
Quaest. Ent., 1982, 18 (1-4)
182
Bell and Bell
14 ' Median lobe relatively broad, its margins more strongly curved; outer carina
widest near middle, in many specimens sinuate anterior to hind angle 15
15 (14") Orbital groove complete, continuously pollinose to base of temporal lobe 16
15' Pollinosity of orbital groove incomplete, ended posteriorly near posterior margin
of eye, or else interrupted there 18
16 (15) Head relatively broad posteriorly, its lateral margin abruptly rounded near base;
latter transverse; anteriomedial margin transverse; frontal space broadly
U-shaped O. fringillus new species, p. 240
16' Head not broad posteriorly, lateral margin evenly rounded from eye to base;
frontal space more narrow; anteriomedial margin oblique 17
17 (16') Outer carina markedly narrowed posteriorly, width at base less than 0.5 of
greatest width; median lobe narrow, its junction with clypeus distinctly
constricted; apex of median lobe obtuse; postantennal area convex, extensively
pollinose O. wittmeri new species, p. 227
17' Outer carina scarcely narrowed posteriorly; width at base 0.9 of greatest width;
median lobe relatively broad, its junction with clypeus scarcely constricted; apex
of median lobe rounded; postantennal area less extensively pollinose
O. gurney i new species, p. 235
18 (15') Tip of median lobe narrow, subacute; pronotum narrow, lateral margins
strongly curved; eye somewhat reduced, with posterioventral margin oblique;
antennal segments V-XI impunctate O. semperi new species, p. 217
18' Tip of median lobe broadly rounded or obtuse; pronotum quadrate or
subquadrate, in most specimens with lateral margins less rounded; eye round,
not reduced; antennal Segments VII-X, in some species with punctures
indistinct 19
19 (18") Outer antennal segments, including X, with numerous coarse punctures 20
19' Outer antennal segments with punctures sparse, in some species extended only
to Segment IX 22
20 (19) Postantennal region of temporal lobe in form of narrow, pollinose ridge;
pronotum subquadrate O. oroensis new species, p. 240
20' Postantennal region glabrous posterior to antennal groove; pronotum distinctly
narrowed anteriorly 21
21 (20') Medial angle of temporal lobe rounded; posteriomedial margin rounded; elytral
punctures round, coarse O. puncticornis new species, p. 241
21' Medial angle obtusely pointed; posteriomedial margin slightly sinuate; strial
punctures elliptical, fine, sparse O. viduus new species, p. 226
22 (19') Pronotum elongate, lateral margins only slightly curved; temporal lobe with
30-50 punctures; median lobe obtusely rounded; marginal groove deeper
posteriorly O. scopulinus new species, p. 237
22' Pronotum shorter; temporal lobe with 20 or fewer punctures; medial angle
produced; posteriomedial margin sinuate in most specimens; marginal groove
not deeper posteriorly 23
23 (22') Median lobe with approximately 10 fine punctures
O. vicinus (Grouvelle), p. 239
23' Median lobe impunctate 24
24 (23') Median lobe narrow, its tip acute 25
Revision of Omoglymmius Ganglbauer
183
24' Median lobe relatively broad, its tip obtuse 26
25 (24) Pronotum subquadrate, widest anterior to middle; outer carinae dilated,
divergent near base O. classicus new species, p. 238
25' Pronotum subcircular, markedly narrowed at both base and apex; outer carinae
narrow, parallel at bases O. princeps new species, p. 236
26 (24') Outer carinae narrowed to neither dilated nor divergent bases 27
26" Outer carinae scarcely narrowed to bases, latter either sinuate medially or else
distinctly dilated 28
27 (26) Pronotum subquadrate, lateral margins nearly parallel; base of pronotum
scarcely narrowed O. lindrothi new species, p. 232
21' Pronotum oval, lateral margins strongly curved; base of pronotum distinctly
narrowed O. rusticus new species, p. 234
28 (26') Medial margin of temporal lobe distinctly angulate opposite tip of median lobe;
frontal space more nearly U-shaped O. modicus new species, p. 233
28' Medial margin of temporal lobe scarcely angulate opposite tip of median lobe;
frontal space more nearly V-shaped 29
29 (28') Medial angles of temporal lobe produced, blunt; posteriomedial margin
emarginate; lateral margins of pronotum markedly curved; lateral pits of
abdominal Sternum IV shallow in female O. manni new species, p. 234
29' Medial angles not produced; posteriomedial margin not emarginate; lateral
margins of pronotum less curved, subparallel; lateral pits of Sternum IV deep,
round in female O. regius new species, p. 235
30 (12') Median lobe very narrow, elongate, margins nearly parallel, tip acute; base of
outer carina markedly narrowed O. crassicornis new species, p. 21 1
30' Median lobe moderate in width or broad, margins either parallel or not parallel;
base of outer carina less narrowed 31
31 (30') Median lobe with sides parallel, tip obtuse; greatest width of outer carina at
anterior 0.33; base of Interval V elevated laterad to longitudinal scarp of Stria
IV; Antennal Segments V-XI punctate 32
3L Median lobe lance-shaped, margins not parallel; outer carina either of uniform
width or else with greatest width at middle; base of Interval V not elevated;
outer antennal segments with punctures indistinct or absent 33
32 (31) Lateral margin of outer carina deeply sinuate anterior to hind angle
O. amplus new species, p. 21 1
32' Lateral margin scarcely sinuate anterior to hind angle
O. modiglianii new species, p. 212
33 (31') Lateral margin of temporal lobe almost straight; anteriomedial margin of
temporal lobe abruptly bent; frontal space very broad
O. morditus new species, p. 228
33' Lateral margin of temporal lobe rounded; anteriomedial margin abruptly bent
or rounded; frontal space narrower 34
34 (33') Head as broad as long or broader than long; frontal space broad, U-shaped 35
34' Head longer than broad; frontal space V- or U-shaped 38
35 (34) Outer carina slightly narrower than inner carina at middle
O. caelatus Bell and Bell, p. 230
35' Outer carina equal in width at middle to inner carina 36
Quaest. Ent.. 1982, 18 (1-4)
184
Bell and Bell
36 (35') Antennal groove very narrow; temporal lobes very flat; length 5. 0-6. 8 mm .
O. oceanicus Bell and Bell, p. 231
36' Antennal groove broader; temporal lobes convex; length 6. 5-7. 2 mm 37
37 (360 Pronotum nearly quadrate, scarcely narrowed posteriorly
O. batchianus (Arrow), p. 223
37' Pronotum distinctly narrowed posteriorly, markedly narrowed anteriorly
O. humeralis (Grouvelle), p. 224
38 (340 Antennal Segments V-VIII, and in some specimens also IX, X finely punctate 39
38' Antennal Segments V-XI impunctate 41
39 (38) Inner carina of pronotum constricted just anterior to base, latter broadened . .40
39' Inner carina not constricted just anterior to base; latter truncate
O. re nut us new species, p. 236
40 (39) Preorbital pit extensively pollinose; median lobe broad
O. trepidus new species, p. 242
40' Preorbital pit with pollinosity restricted; median lobe narrow
O. cavea new species, p. 243
41 (38') Temporal lobe punctate over most of its surface; median lobe narrow, tip acute;
sides of pronotum nearly parallel O. philippensis (Chevrolat), p. 209
41' Punctures of temporal lobe limited to lateral margin; median lobe slightly
broader, tip more obtuse; pronotum widest at middle, sides curved 42
42 (41') Eye large, round; basal scarp of Stria IV distinct; male with ventral tooth on
anterior femur O. imugani new species, p. 210
42' Eye slightly reduced; basal scarp of Stria IV very small, scarcely more than
lateral margin of pollinose spot; male without ventral tooth on anterior femur.
O. politus new species, p. 210
43 (11') Median lobe broad, margins rounded; abdominal Sterna IV, V with punctures
not coalescent O. opticus new species, p. 225
43' Median lobe narrow, elongate; Sterna IV, V with punctures coalescent near
lateral margins, forming shallow pits 44
44 (43') Anterior part of temporal lobe forming narrow, pollinose ridge; glabrous area of
temporal lobe separated from antennal lobe by nearly length of antennal lobe;
temporal setae one to three (in most specimens two); temporal lobe with up to
three punctures near lateral margin or none O. duplex new species, p. 220
44' Anterior part of temporal lobe glabrous nearly to antennal lobe, separated from
latter only by rather broad antennal groove; 1 temporal seta; temporal lobe with
many punctures O. bouchardi new species, p. 221
45 (10') Inner, outer, or both pairs of pronotal carinae punctate 46
45' Pronotal carinae entirely impunctate 56
46 (45) Inner carina with three or more (in most species many) punctures 47
46' Inner carina with one or two punctures or none 49
47 (46) Head twice as long as wide; median lobe elongate, margins almost parallel
O. nasalis new species, p. 228
47' Head slightly or not at all longer than wide; median lobe lance-shaped, margins
not parallel 48
48 (47') Median lobe constricted at junction with clypeus; tip of median lobe
subtruncate; Antennal Segments IV-IX impunctate; base of outer carina not
Revision of Omoglymmius Ganglbauer
185
48 (47') Median lobe constricted at junction with clypeus; tip of median lobe
subtruncate; Antennal Segments IV-IX impunctate; base of outer carina not
sloped into basal impression; eye slightly reduced, about 0.33 of length of
temporal lobe in profile view O. data new species, p. 217
48' Median lobe not constricted at junction with clypeus; tip of median lobe acute;
all antennal segments impunctate; eye not reduced, about 0.5 of length of
temporal lobe; base of outer carina sloped gradually into basal impression
O. ephemeris new species, p. 244
49 (46') Intervals II, IV distinctly elevated anteriorly; head elongate anterior to eye;
frontal grooves very narrow O. mycteroides new species, p. 237
49' Intervals II, IV not elevated; head not elongate; frontal grooves moderately
broad
50 (49') Prosternum with precoxal carinae; temporal seta absent
O. thoracicus new species, p. 216
50' Prosternum without precoxal carinae; one temporal seta
51 (50') Stria IV with five or six setae along its entire length
51' Stria IV with one or two setae near apex
52 (51) Outer carina scarcely narrowed at base O. coelebs new species, p. 215
52' Outer carina strongly narrowed to base; latter bluntly pointed
53 (52') Median lobe not constricted at junction with clypeus; tip of median lobe
rounded; pronotum elongate O. malaicus (Arrow), p. 214
53' Median lobe constricted at junction with clypeus; tip of median lobe acute;
pronotum more nearly quadrate
54 (53') Bases of outer carinae markedly divergent
O. fraudulentus new species, p. 214
54' Bases of outer carinae scarcely divergent O. nemoralis new species, p. 213
55 (51') Antennal Segments I-XI with faint, irregular punctures; lateral pits of
abdominal Sternum IV narrow, oblique, slit-like; temporal lobe, outer carina of
pronotum finely punctate O. sectatus new species, p. 243
55' Antennal Segments I-IV punctate; Segments V-X impunctate; lateral pits of
SternumlV shallow, round; temporal lobe, outer carina coarsely punctate
O. evasus new species, p. 212
56 (45') Temporal lobe punctate; temporal seta one
56' Temporal lobe impunctate; temporal setae two to four
57 (56) Median lobe narrow, elongate; anterior femur of male with ventral tooth; hind
calcar acute, angular
57' Median lobe broader, lance-shaped; anterior femur of male without ventral
tooth; hind calcar prominent, apex rounded. O. gracilicornis (Grouvelle), p. 244
58 (57) Abdominal Sterna III-V with coarse punctures, these coalescent near lateral
margin; Stria IV with one or two setae near apex. O. consors new species, p. 221
58' Abdominal Sterna III-V with finer, isolated punctures, not at all coalescent
laterally; Stria IV with five setae along its length. . O. hiekei new species, p. 218
59 (56') Precoxal carinae absent; outer carina as broad as inner one at middle
O. quadruplex new species, p. 219
59' Precoxal carina present; outer carina 0.66 as broad as inner carina at middle
O. pectoralis new species, p. 218
50
51
52
55
53
54
57
59
58
Quaesi. Ent., 1982, 18 (1-4)
186
Bell and Bell
60 (9X) Outer carina of two planes meeting at sharp edge, one vertical, other sloped
toward paramedian groove; pronotum hexagonal to subquadrate; Stria IV with
five to seven setae along its length O. tabulatus new species, p. 238
60' Outer carina not formed of two planes; pronotum not hexagonal; Stria IV with
one or two setae near apex 61
61 (600 Outer carina less than 0.4 as wide as inner carina at middle; Antennal Segments
V-X punctate O. solitarius (Arrow), p. 208
61 ' Outer carina about 0.5 as wide as inner one at middle; antennal Segments V-X
with punctures absent or indistinct 62
62 (6L) Pronotol carinae impunctate; temporal lobe with eight to 10 punctures near
lateral margin O. impletus Bell and Bell, p. 231
62' Outer carina punctate; temporal lobe with 20-30 punctures
O. patens new species, p. 242
63 (60 Suborbital tubercle present 64
63' Postorbital tubercle present 68
64 (63) Outer carina more than 0.6 as wide as inner carina at middle; marginal groove
not dilated 65
64' Outer carina about 0.4 as wide as inner carina at middle; marginal groove
dilated O. pulvinatus (Grouvelle), p. 252
65 (64) Outer carina broadest just anterior to base; base of inner carina strongly
narrowed; inner carina impunctate O. sedlaceki new species, p. 252
65' Outer carina narrowed posteriorly; base of inner carina less narrowed; inner
carina punctate 66
66 (650 Median lobe broad, tip subtruncate; pronotum nearly quadrate; metasternal
punctures limited to midline, margins O. biroi new species, p. 251
66" Median lobe narrow, tip acute; pronotum not quadrate; metasternum entirely
punctate 67
67 (660 Median lobe punctate; outer carina equal to or slightly narrower than inner
carina at middle; one temporal seta O. cheesmanae (Arrow), p. 250
67' Median lobe impunctate; outer carina distinctly narrower than inner carina at
middle; temporal seta absent O. asetatus new species, p. 250
68 (630 Postorbital tubercle visible only in lateral view 69
68' Postorbital tubercle visible in dorsal view 72
69 (68) Lateral margins of pronotum nearly parallel; median lobe broad, rounded;
pollinosity of orbital groove extended to posterior margin of eye
O. quadraticollis (Arrow), p. 223
69' Lateral margins more curved; pronotum not subquadrate; median lobe broad to
narrow, obtuse to rounded; pollinosity of orbital groove less extensive 70
70 (690 Inner carina truncate at base; outer carina narrowed to base
O. repetitus new species, p. 222
70' Inner carina pointed at base; outer carina dilated at base 71
71 (700 Median lobe broad, tip rounded; medial margin of base of outer carina sinuate;
pollinose area of orbital groove broad; temporal lobe with scattered, very minute
punctures; both pronotal carinae with very minute punctures
O.follis new species, p. 245
7L Median lobe angulate at tip; medial margin of base of outer carina not sinuate;
Revision of Omoglymmius Ganglbauer
187
pollinosity of orbital groove limited; temporal lobe with few minute punctures
near lateral margin; pronotal carinae impunctate
O. iridescens new species, p. 245
72 (680 Outer carina slightly narrower, more convex than inner one; carinae impunctate;
anterior end of outer carina elevated, obliquely truncate
O. planiceps new species, p. 248
72' Outer carina as wide and no more convex than inner one; anterior end of outer
carina neither elevated nor obliquely truncate; carinae punctate or impunctate .
73 (720 Metasternum entirely punctate
73' Metasternum with punctures limited to midline, margins
74 (73) Inner, outer carinae with numerous fine punctures; outer carina narrowed to
base; strial punctures round, pilose, coarse O. sus new species, p. 248
74' Outer carina punctate; inner carina impunctate; outer carina dilated at base;
strial punctures fine, especially in Striae I-III
75 (740 Postorbital tubercles very large, divergent; outer carina with many fine
punctures; medial angle of temporal lobe rounded O. capito (Grouvelle), p. 249
15' Postorbital tubercles smaller, scarcely divergent; outer carina with one or two
fine punctures; medial angle obtuse O. lentus new species, p. 249
76 (730 Postorbital tubercle very large; apex of pronotum less narrowed; strial punctures
elliptical, very fine O. auratus new species, p. 247
76' Postorbital tubercle small; pronotum more narrowed anteriorly; strial punctures
round, moderately fine
77 (760 Median lobe broad, tip rounded; width of pronotum at middle subequal to width
at base O. massa new species, p. 246
11' Median lobe narrower, tip obtusely rounded; width of pronotum at middle
clearly less than width at base O. denticulatus new species, p. 247
73
74
76
75
77
KEY TO EURASIAN SPECIES (INCLUDES RYUKYU AND ANDAMAN ISLANDS)
1 Outer carina equal to or wider than inner carina at middle 2
1 ' Outer carina 0.25 as wide as inner carina at middle (Andaman Is.)
O. solitarius (Arrow), p. 208
2 (1) Inner carina with anterior half distinctly narrower than posterior half; greatest
width of inner carina distinctly less than that of outer carina (South India). . .
O. malabaricus (Arrow), p. 207
2' Inner carina not distinctly narrower in anterior half; inner and outer carinae
equal in width 3
3 (2') Median lobe constricted at base of clypeus; temporal seta absent 4
3' Median lobe not constricted at junction with clypeus; one temporal seta (Malay
Peninsula) O. malaicus (Arrow), p. 214
4 (3) Orbital groove absent; prosternum with precoxal carinae; posteriolateral margin
of temporal lobe rounded 5
4' Orbital groove represented by strip of pollinosity; prosternum without precoxal
carinae; posteriolateral margin of temporal lobe sinuate (Europe, western Asia).
O. germari (Ganglbauer), p. 194
Quaest. Ent., 1982, 18 (1-4)
188
Bell and Bell
5 (4) Medial angles of temporal lobes acute, contiguous; Stria IV with two to four
setae along its length; total length 5. 1-5.2 mm (Ryukyu Is.)
O. sakuraii (Nakane), p. 207
5' Medial angles of temporal lobe obtuse, slightly separated; Stria IV with 1-2
setae near apex; total length 6. 1-7.1 mm (Bhutan) O. laticeps Bell, p. 206
KEY TO SPECIES OF GREATER SUNDA AND PHILIPPINE ISLANDS
1 At least one pair of pronotal carinae punctate
V Pronotal carinae entirely impunctate
2 (1) Inner carina with several to many punctures (only three or four in O.
summissus)
2' Inner carina impunctate or with only one or two punctures on each carina
3 (2) Pronotum quadrate; outer carina widest near anterior margin, where flattened,
angulate; outer carina only slightly narrower at base than at middle; eye large
(Sumatra) O. summissus new species, p. 216
3' Pronotum not quadrate, its lateral margins markedly curved; outer carina
distinctly narrowed to base, apex, widest at middle; eye reduced
4 (30 Base of Stria IV with small pollinose longitudinal scarp; pollinosity of orbital
groove extended at least to middle of eye; depth of eye about 0.6 of depth of
head; posterior margin of eye oblique, nearly straight (Philippines, island
unknown) O. semperi new species, p. 217
4' Base of Stria IV without longitudinal scarp; orbital groove absent; depth of eye
less than 0.5 of depth of head, its posterior margin curved (Luzon)
O. data new species, p. 217
5 (2') Base of Stria IV with longitudinal pollinose scarp, or without scarp but with
pollinose spot {O. politus )
5' Base of Stria IV without longitudinal pollinose scarp and without pollinose spot
6 (50 Pronotum widest at middle or else with lateral margins almost parallel; median
lobe of head narrow, its tip acute; base of Interval IV not elevated
6 ' Pronotum widest well anterior to middle; median lobe wide, margins almost
parallel, tip rounded; base of Interval IV distinctly elevated
7 (6) Base of outer carina very markedly narrowed; median lobe very narrow, margins
nearly parallel; punctures of abdominal Sterna IV and V each form transverse
row (Negros) O. crassicornis new species, p. 21 1
7' Base of outer carina only moderately narrowed; median lobe broader; punctures
of Sterna IV, V numerous, scattered
8 (7') Temporal lobe puncture over most of surface; median lobe narrow, tip acute;
sides of pronotum nearly parallel (Luzon, Mindoro, Palawan, Siargao)
O. philippensis (Chevrolat), p. 209
8" Temporal lobe punctate only near lateral margin; median lobe broader, tip more
obtuse; pronotum widest at middle, sides curved
9 (8') Eye large, round; basal scarp of Stria IV distinct; male with ventral tooth on
anterior femur (Luzon) O. imugani new species, p. 210
2
15
3
5
4
6
11
7
10
8
9
Revision of Omoglymmius Ganglbauer
189
9' Eye slightly reduced; basal scarp of Stria IV very small, scarcely more than
slight elevation of lateral margin of pollinose spot; male without ventral tooth on
anterior femur (Luzon) O. politus new species, p. 210
10 (6') Lateral margin of outer carina deeply sinuate anterior to hind angle (Sumatra)
O. amplus new species, p. 21 1
10' Lateral margin scarcely sinuate anterior to hind angle (Mentawei)
O. modiglianii new species, p. 212
1 1 (5") Prosternum with precoxal carinae (Java) O. thoracicus new species, p. 216
1 V Prosternum without precoxal carinae 12
12 (1 L) Outer carina not narrowed at base (Palawan). O. coelebs new species, p. 215
1 7! Outer carina narrowed at base 13
13 (120 Stria IV with one seta near apex, none or one at base (Mindanao)
O. evasus new species, p. 212
1 3' Outer carina with four or five setae, forming continuous row 14
14 (130 Bases of outer carinae strongly divergent (Sumatra)
O. fraudulentus new species, p. 214
14' Bases of outer carinae not divergent (Sarawak)
O. nemoralis new species, p. 213
15 (10 Basal transverse scarp of elytron with continuous pollinose area medial to base
of Stria IV 1 6
15' Basal transverse scarp without continuous pollinose area medial to base of Stria
IV 17
16(15) One temporal seta; median lobe with sides parallel, tip truncate (Sumatra,
Borneo) O. consors new species, p. 221
16' Two to four temporal setae; median lobe lance-shaped, tip acute (Mindanao) .
O. quadruplex new species, p. 219
17 (15') Base of Stria IV with longitudinal pollinose scarp 18
17' Base of Stria IV without scarp 19
18 (17) Temporal lobe with many punctures; one temporal seta; temporal lobe glabrous
anteriorly, separated from antennal lobe only by pollinose antennal groove
(Sumatra) O. bouchardi new species, p. 221
18' Temporal lobe with one or two punctures near lateral margin; in most
specimens, two temporal setae (in a few, one or three); temporal lobe connected
to antennal lobe by pollinose longitudinal ridge (Luzon, Palawan, Sibuyan,
Mindanao) O. duplex new species, p. 220
19 (17') Prosternum with precoxal carinae (Java) O. pectoralis new species, p. 218
19' Prosternum without precoxal carinae (Luzon) O. hiekei new species, p. 218
KEY TO SPECIES OF “WALLACEA” (CELEBES, MOLUCCAS, LESSER SUNDAS,
KEI AND SCHOUTEN ISLANDS)
1 Fifth interval elevated, carinate (Yapan). O. bicarinatus new species, p. 230
V Fifth interval not elevated nor carinate 2
2 (L) Outer carina of pronotum much narrower than inner carina; outer carina with
one to three setae inserted in conspicuous punctures (Sumbawa)
Quaest. EnU, 1982, 18 (1-4)
190
Bell and Bell
O. bucculatus (Arrow), p. 229
2 ' Pronotal carinae of nearly equal width at middle; outer carina without setae 3
3 (20 Head about twice as long as wide; median lobe elongate; its margins parallel
(Buru) * O. nasalis new species, p.228
3' Head at most slightly longer than wide 4
4 (3') Postorbital tubercle present 5
4' Postorbital tubercle absent 6
5 (4) Pronotum quadrate, its margins nearly straight and parallel; median lobe broad,
its apex obtuse; orbital groove extensively pollinose (Tanimbar)
O. quadraticollis (Arrow), p. 223
5' Pronotum not quadrate, its margins rounded; median lobe narrower, its apex
acute; orbital groove with pollinosity restricted to anterior end (Celebes)
O. repetitus new species, p. 222
6 (40 Lateral margin of temporal lobe almost straight; anteriomedial margin of
temporal lobe abruptly bent; frontal space very broad (Morotai)
O. morditus new species, p. 228
6' Lateral margin of temporal lobe rounded; anteriomedial margin bent or
rounded; frontal space narrower 7
7 (60 Abdominal sterna with extensive lateral pollinose area, but without distinct
punctures (Amboina) O. vadosus new species, p. 227
1' Abdominal sterna with distinct punctures, in some specimens coalescent
laterally 8
8 (70 Anterior portion of temporal lobe extensively pollinose, glabrous portion of
temporal lobe broadly separated from antennal lobe 9
8' Anterior part of temporal lobe not pollinose; glabrous portion of temporal lobe
separated from antennal lobe only by narrow antennal groove 10
9 (8) Pronotum subquadrate; one to three temporal setae; medial angle of temporal
lobe rounded; Segment I of antenna with apex inflated laterally (Mangole)
O. continuus new species, p. 226
9 ' Pronotum not subquadrate, its base distinctly narrowed; one temporal seta;
medial angle of temporal lobe obtuse; Segment I of antenna with apex not
inflated (Mangole) O. wittmeri new species, p. 227
10 (8') Pronotum with sides strongly curved; temporal lobe with 25-30 rather coarse
punctures; outer carina with 35-40 punctures (Kei Is.)
O. viduus new species, p. 226
10' Pronotum subquadrate, its margins only slightly curved; temporal lobe with
10-15 punctures; outer carina with 10 or fewer punctures 11
11 (100 Frontal space 0.33 of width of temporal lobe; eye relatively convex (Dammar)
O. opticus new species, p. 225
1 Y Frontal space wider, about 0.43 of width of temporal lobe; eye less convex 12
12 (1 10 Pronotum nearly quadrate, scarcely narrowed posteriorly (Batjan)
O. batchianus (Arrow), p. 223
12' Pronotum distinctly narrowed posteriorly, and rather strongly so anteriorly
(Ternate, Halmahera) O. humeralis (Grouvelle), p. 224
Revision of Omoglymmius Ganglbauer
191
KEY TO SPECIES FROM SOLOMON ISLANDS
1 Outer carina much narrower than inner one, its dorsal surface flat, meeting
lateral surface at sharp angle; Stria IV with four or five setae (Bougainville,
New Georgia) O. tabulatus new species, p. 238
Y Outer carina subequal to inner carina at middle; outer carina more or less
convex, its dorsal surface not in form of lateral margin with lateral face 2
2 (L) Head elongate, about twice as long as wide; median lobe elongate, narrow, its
sides nearly parallel; elytron without longitudinal scarp at base of Stria IV
(New Georgia, Guadalcanal) O. mycteroides new species, p. 237
2' Head not elongate, at most slightly longer than wide; median lobe broader, its
sides not parallel; elytron with short longitudinal scarp at base of Stria IV 3
3 (2') Pronotum with sides strongly curved, conspicuously wider at middle than at
base or apex; lateral margin sinuate anterior to hind angle 4
3' Pronotum with sides only slightly curved, nearly parallel (pronotum
subquadrate); lateral margin not sinuate anterior to angle 5
4 (3) Inner carina punctate; pronotum more expanded at middle, more constricted to
base and apex (Bougainville) O. princeps new species, p. 236
4' Inner carina impunctate; pronotum less expanded at middle (New Georgia) .
O. renutus new species, p. 236
5 (3') Pollinosity of orbital groove complete to hind margin of eye, conspicuous; elytral
striae with continuous pollinosity (Bougainville, Choiseul, Faro)
O. gurney i new species, p. 235
5' Pollinosity of orbital groove incomplete; elytral striae with pollinosity confined
to punctures 6
6 (50 Pronotum elongate, its sides nearly parallel; temporal lobe with 30-35
punctures; medial angle of temporal lobe broadly rounded; inner carina with
15-18 punctures; total length larger, about 7.3 mm (Santa Cruz)
O. scopulinus new species, p. 237
6' Pronotum not elongate; temporal lobe with less than 30 punctures; medial angle
of temporal lobe produced, oblique; inner carina with two to eight punctures 7
7 (6') Outer carina as broad as inner carina at middle; base of outer carina only
slightly narrower than its greatest width, latter slightly greater than width of
inner carina at base; medial margin of outer carina sinuate near base 8
7" Outer carina slightly narrower than inner carina at middle; base of outer carina
more narrowed, its width equal to or slightly less than width of inner carina at
base; medial margin of outer carina not sinuate near base 10
8 (7) Anteriomedial margin of temporal lobe curved gradually; frontal space more
V-shaped 9
8' Medial margin of temporal lobe bent abruptly near middle, frontal space more
U-shaped (Savo,Ngela) O. modicus new species, p. 233
9 (8) Posteriomedial margin of temporal lobe distinctly emarginate between occipital
and medial angles; latter produced, closer together; margins of pronotum
strongly curved; lateral pits of Sternum IV in female relatively shallow and
indefinite (Malaita) O. manni new species, p. 234
9' Posteriomedial margin scarcely emarginate; medial angles not produced, more
Quaest. Ent., 1982, 18 (1-4)
192
Bell and Bell
separated from one another; margins of pronotum less curved: lateral pits of
Sternum IV in female deeper, rounder (Isabella) . . O. regius new species, p. 235
10 (7') Pronotum subquadrate, lateral margins nearly parallel; base of pronotum
scarcely narrowed (Guadalcanal) O. lindrothi new species, p. 232
10' Pronotum with lateral margins strongly curved; base of pronotum strongly
narrowed (Russell Is., San Cristobal) O. rusticus new species, p. 234
KEY TO SPECIES FROM NEW GUINEA AND ADMIRALTY ISLANDS
1 Neither postorbital nor suborbital tubercles present 2
V Either postorbital or suborbital tubercle present 12
2 (1) Frontal space elongate, narrow, V-shaped or slightly U-shaped; antennal groove
narrow 3
2' Frontal space broad, strongly U-shaped; antennal groove broad 8
3 (2) Base of Stria IV with longitudinal pollinose scarp 4
3' Base of Stria IV without longitudinal pollinose scarp 6
4 (3) Median lobe narrow, tip acute; pollinosity very limited at lateral end of antennal
groove 5
4' Median lobe broader, tip obtuse; pollinosity expanded at lateral end of antennal
groove O. trepidus new species, p. 242
5 (4) Outer carina 0.5 as wide as inner carina at middle; lateral pit of Sternum IV of
female rounded O. patens new species, p. 242
5' Outer carina as wide as inner carina at middle; lateral pit of Sternum IV of
female triangular, broad O. cavea new species, p. 243
6 (3') Pronotal carinae impunctate; Stria IV with two to four setae; male without
ventral tooth on anterior femur; hind calcar of male large, obtuse; total length
4.5 mm O. gracilicornis (Grouvelle), p. 244
6' Pronotal carinae punctate; Stria IV with one seta near apex; male with ventral
tooth on anterior femur; hind calcar acute; most specimens longer than 4.5 mm .7
7 (6') Apex of median lobe acute; median lobe with seven to 10 punctures; lateral pits
of abdominal Sternum IV rounded; middle calcar of male elongate
O. ephemeris new species, p. 244
T Apex of median lobe obtuse; median lobe with one puncture or none; lateral pits
of Sternum IV narrow, oblique; middle calcar of male very small
O. sectatus new species, p. 243
8 (27) Pronotum subquadrate, margins only slightly curved 9
8' Pronotum not subquadrate, margin strongly curved
O. puncticornis new species, p. 241
9 (8) Outer carina narrowed to base; margin not sinuate anterior to hind angle; length
4. 6-6. 2 mm 10
9' Outer carina narrowed to subbasal constriction, posterior to which it is
broadened; margin sinuate anterior to hind angle; length 6.0-7. 2 mm 11
10 (9) Median lobe with a few scattered punctures (New Guinea)
O. vicinus (Grouvelle), p. 239
10' Median lobe impunctate (Admiralty Is.) O. classicus new species, p. 238
Revision of Omoglymmius Ganglbauer
193
1 1 (90 Frontal space broad-U-shaped; posterior margin of temporal lobe nearly
transverse O. fr ingill us new species, p. 240
11' Frontal space more elongate, heart-shaped; posterior margin of temporal lobe
more evenly rounded O. oroensis new species, p. 240
12 (F) Suborbital tubercle present 13
12' Postorbital tubercle present 17
13 (12) Outer carina more than 0.6 as wide as inner carina at middle; marginal groove
not dilated 14
13' Outer carina about 0.4 as wide as inner carina at middle; marginal groove
dilated O. pulvinatus (Grouvelle), p. 252
14(13) Outer carina broadest just anterior to base; base of inner carina strongly
narrowed; inner carina impunctate O. sedlaceki new species, p.252
14' Outer carina narrowed posteriorly; base of inner carina less narrowed; inner
carina punctate 1 5
15 (14x) Median lobe broad, tip subtruncate; pronotum subquadrate, lateral margins
only slightly curved; posterior margin of temporal lobe nearly transverse;
punctures of metasternum restricted to midline, lateral margins.
O. biroi new species, p. 25 1
15' Median lobe narrow, tip acute to subacute; pronotum not subquadrate, lateral
margins strongly curved; entire disc of metasternum punctate 16
16 (15') Median lobe punctate; outer carina equal to or slightly narrower than inner
carina at middle; one temporal seta O. cheesmanae (Arrow), p. 250
16' Median lobe impunctate; outer carina distinctly narrower than inner carina at
middle; temporal seta absent O. asetatus new species, p. 250
17 (12') Postorbital tubercle visible only in lateral view, hidden in dorsal view 18
17' Postorbital tubercle visible in dorsal as well as lateral view 19
18 (17) Median lobe broad, tip obtuse; medial margin of base of outer carina sinuate;
antennal groove extended laterally into broad pollinose area; temporal lobe,
pronotal carinae with scattered fine punctures . . O.follis new species, p. 245
18' Median lobe narrow, lance-shaped, tip acute; medial margin of outer carina not
sinuate near base; antennal groove extended laterally into short, narrow
pollinose area; temporal lobe with very few punctures; pronotal carinae
impunctate O. iridescens new species, p. 245
19 (17') Metasternum with punctures limited to midline, margins 20
19' Metasternum with punctures scattered over entire disc 22
20 (19) Postorbital tubercle larger; anterior margin of outer carina abruptly truncate;
punctures of elytral striae very fine, elliptical over most of disc, obsolete near
base O. auratus new species, p. 247
20' Postorbital tubercle smaller, anterior margin of outer carina evenly sloped; strial
punctures coarse, round on most of disc, becoming fine, elliptical near base 21
21 (20') Median lobe broad, tip rounded; postorbital tubercle of moderate size; medial
angles of temporal lobes sharper, more produced
O. massa new species, p. 246
21' Median lobe narrow, tip obtuse; postorbital tubercle very small; median angles
obtuse, scarcely produced O. denticulatus new species, p. 247
22 (19') Outer carina more convex than, slightly narrower than inner one at middle
Quaest. Ent., 1982, 18 (1-4)
194
Bell and Bell
O. planiceps new species, p. 248
22' Outer carina equal in convexity, width to inner one at middle 23
23 (22') Both pairs of carinae punctate; strial punctures round, coarse, pollinose
O. sus new species, p. 248
23' Inner carina impunctate; outer carina with few scattered punctures; strial
punctures shallow, those of Striae I-III finer than those of outer striae 24
24 (230 Postorbital tubercles relatively small, not divergent posteriorly; medial angle of
temporal lobe slightly produced; posteriomedial margin slightly emarginate
between medial, occipital angles O. lentus new species, p.249
24' Postorbital tubercles large, divergent posteriorly; me angle of temporal lobe
rounded; posteriomedial margin not emarginate O. capito (Grouvelle), p. 249
KEY TO SPECIES FROM THE CAROLINE ISLANDS
1 Outer carina of pronotum punctate; inner carina narrowest anterior to base,
broadened from there to base 2
T Outer carina impunctate; inner carina tapered to point at base (Yap)
O. impletus Bel! and Bell, p. 231
2 (1) Temporal lobe with margin sinuate between medial angle and occipital angle
(Palau) O. caelatus Bell and Bell, p. 230
2' Temporal lobe with margin evenly convex between medial and occipital angles
(Kusiae, Ponape, Palau) O. oceanicus Bell and Bell, p. 231
Omoglymmius (sens u stricto ) germari (Ganglbauer)
Figs. 113, 121
Rhysodes exaratus Erichson 1847: 301 (nee Dalman 1823; nee Serville 1825)
Rhysodes aratus Chevrolat 1873a: 209 (nee Newman 1838)
Rhysodes americanus Reitter 1882: 140 (nee Castelnau 1836)
Rhysodes germari Ganglbauer 1892: 534
Omoglymmius (sensu stricto ) german (Ganglbauer) Bell and Bell 1978
Type material. - None designated. NEOTYPE (here designated) female, labelled:
“Majevica, Bosna, CNHM 1955, Karl Branczik colln., ex Eduard Knirsch” (in
Yugoslavia)(CNHM). PARANEOTYPES three males, four females, labelled: “Majevica,
Bosna, VI. Zoufal” (MNHB); one male, labelled: ’’Majevica, Bosna (Bosna = Bosnia), VI
Zoufal, coll. G. Falkenstrom’’ (LUN).
Description. — Length 6. 8-8.0 mm. Antennal segments impunctate; median lobe impunctate, rhomboid, its apex
subacute; frontal space U-shaped, anteriomedial margin of temporal lobe abruptly curved; medial angles obtuse, slightly
separated; posteriomedial margin oblique; occipital angle obtuse, distinct; posteriolateral margin oblique, parallel to
posterior half of anteriomedial margin; posteriolateral margin slightly sinuate just posterior to eye; orbital groove pollinose,
extended to level of hind margin of eye; temporal lobe with five to 12 coarse punctures, restricted to lateral portion;
temporal seta absent; small postorbital tubercle present, or this tubercle merely suggested or entirely absent.
Pronotum moderately elongate, length/greatest width 1.29; widest near middle, apex markedly narrowed; base nearly
equal to greatest width; lateral margin with slight sinuation anterior to hind angle; outer carina slightly narrower than
inner one at middle; medial margin of outer carina slightly sinuate anterior to base; base of outer carina narrowed; inner
carinae narrowed at base; prpnotal carinae finely punctate; punctures more numerous on outer carina; inner carina with
two to 10 punctures; pronotum without setae; prosternum without precoxal carinae.
Revision of Omoglymmius Ganglbauer
195
Plate 9. Figures 1 13-129, Subgenus Omoglymmius sensu stricto. Figs. 1 13-120, Head and pronotum, dorsal
aspect; Fig. 1 13, Omoglymmius (s. str.) germari (Ganglbauer); Fig. 114, O. (s. str.) laticeps Bell ; Fig. 1 15,
O. (s. str.} sakuraii (Nakane); Fig. 116, O. (s. str.) malabaricus (Arrow); Fig. 117, O. (s. str.) solitarius
(Arrow); Fig. 1 18, O. (s. str.) philippensis (Chevrolat); Fig. 1 19, O. (s. str.) imugani new species; Fig. 120, O.
(s. str.) politus new species; Figs. 121-128, Head, lateral aspect; Fig. 121, O. (s. str.) germari (Ganglbauer);
Fig. 122, O. (s. str.) laticeps Bell; Fig. 123, O. (s. str.) sakuraii (Nakane); Fig. 124, O. (s. str.) malabaricus
(Arrow); Fig. 125, O. (s. str.) solitarius (Arrow); Fig. 126, O. (s. str.) philippensis (Chevrolat); Fig. 127, O.
(s. str.) imugani new species; Fig. 128, O. (s. str.) politus new species; Fig. 129, Prothorax, ventrolateral
aspect, O. (s. str.) sakuraii (Nakane).
Quaest. Ent., 1982, 18 (1-4)
196
Bell and Bell
Plate 10. Figures 130-139, Subgenus Omoglymmius sensu stricto. Figs. 130-138, Head and pronotum, dorsal aspect; Fig.
130, Omoglymmius (s. str.) crassicornis new species; Fig. 131, O. (s. str.) amplus new species; Fig. 132, O. (s. str.)
modiglianii new species; Fig. 133, O. (s. str.) malaicus (Arrow); Fig. 134, O. (s. str.) evasus new species; Fig. 135, O. (s.
str.) nemoralis new species; Fig. 136, O. (s. str.) fraudulentus new species; Fig. 137, O. (s. str.) coelebs new species; Fig.
138, O. (s. str.) thoracicus new species; Fig. 139, Prothorax, ventral aspect, O. (s. str.) thoracicus new species.
Revision of Omoglymmius Ganglbauer
197
Plate 1 1. Figures 140-149, Subgenus Omoglymmius sensu stricto. Figs. 140-146, Head and pronotum, dorsal aspect; Fig.
140, Omoglymmius (s. str.) summissus new species; Fig. 141, O. (s. str.) semperi new species; Fig. 142, O. (s. str.) data
new species; Fig. 143, O. (s. str.) hiekei new species; Fig. 144, O. (s. str.) pectoralis new species; Fig. 145, O. (s. str.)
quadruplex new species; Fig. 146, O. (s. str.) duplex new species; Figs. 147-148, Head, lateral aspect; Fig, 147, O. (s. str.)
semperi new species; Fig. 148, O. (s. str.) data new species; Fig. 149, Prothorax, ventral aspect, O. (s. str.) pectoralis new
species;
Quaest. Ent., 1982, 18 (1-4)
198
Bell and Bell
Plate 12. Figures 150-158, Subgenus Omoglymmius sensu stricto. Figs. 150-151, Head, pronotum and anterior portion of;
elytra, dorsal aspect; Fig. 150, Omoglymmius (s. str.) bouchardi new species; Fig. 151, O. ( s . str.) consors new species;
Figs. 152-155, Head and pronotum, dorsal aspect; Fig. 152, O. (s. str.) quadraticollis (Arrow); Fig. 153, O. fs. str.)
batchianus (Arrow); Fig. 154, O. (s. str.) humeralis (Grouvelle); Fig. 155, O. fs. str.) repetitus new species; Figs. 156-158,
Head, lateral aspect; Fig. 156, O. (s. str.) repetitus new species; Fig. 157, O. f s . str.) quadraticollis (Arrow); Fig. 158, O.
fs. str.) batchianus (Arrow).
Revision of Omoglymmius Ganglbauer
199
Plate 13. Figures 159-171, Subgenus Omoglymmius sensu stricto. Figs. 159-165, Head and pronotum, dorsal aspect; Fig.
159, Omoglymmius (s. str.) opticus new species; Fig. 160, O. (s. str.) viduus new species; Fig. 161, O. (s. str.) continuus
new species; Fig. 162, O. (s. str.) wittmeri new species; Fig. 163, O. (s. str.) vadosus new species; Fig. 164, O. (s. str.)
nasalis new species; Fig. 165, O. (s. str.) morditus new species; Figs. 166-167, Head, lateral aspect; Fig. 166, O. (s. str.)
continuus new species; Fig. 167, O. (s. str.) wittmeri new species; Figs. 168-169, Antennal Segments I, II; Fig. 168, O. (s.
str.) wittmeri new species; Fig. 169, O. (s. str.) continuus new species; Figs. 170-171, Hind tibia, male, apical portion; Fig.
170, O. (s. str.) continuus new species; Fig. 171, O. (s. str.) wittmeri new species;
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Plate 14. Figures 172-181, Subgenus Omoglymmius sensu stricto. Figs. 172-179, Head and pronotum, dorsal aspect; Fig.
172, Omoglymmius (s. str.) bucculatus (Arrow); Fig. 173, O. (s. str.) caelatus Bell & Bell; Fig. 174, O. (s. str.) impletus
Bell & Bell; Fig. 175, O. (s. str.) oceanicus Bell & Bell; Fig. 176, O. (5. str.) lindrothi new species; Fig. 177, O. (s. str.)
modicus new species; Fig. 178, O. (s. str.) rusticus new species; Fig. 179, O. (s. str.) manni new species; Fig. 180, Hind
tibia, male, apical portion, O. (s. str.) manni new species; Fig. 181, Head, pronotum and anterior portion of elytra, O. (s.
str.) bicarinatus new species.
Revision of Omoglymmius Ganglbauer
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Plate 15. Figures 182-191, Subgenus Omoglymmius sensu stricto. Figs. 182-188, Head and pronotum, dorsal aspect; Fig.
182, Omoglymmius (s. str.) regius new species; Fig. 183, O. (s. str.) gurneyi new species; Fig. 184, O. (s. str.) princeps new
species; Fig. 185, O. (s. str.) renutus new species; Fig. 186, O. (s. str.) scopulinus new species; Fig. 187, O. (s. str.)
mycteroides new species; Fig. 188, O. (s. str.) tabulatus new species; Fig. 189, Head, lateral aspect, O. (s. str.) gurneyi ;
new species; Figs. 190-191, Hind tibia, male, apical portion; Fig. 190, O. (s. str.) regius new species; Fig. 191, O. (s. str.)
tabulatus new species;
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Bell and Bell
Plate 16. Figures 192-201, Subgenus Omoglymmius sensu stricto. Figs. 192-200, Flead and pronotum, dorsal aspect; Fig.
192, Omoglymmius (s. str.) classicus new species; Fig. 193, O. (s. str.) vicinus (Grouvelle); Fig. 194, O. (s. str.) oroensis
new species; Fig. 195, O. (s. str.) fringillus new species; Fig. 196, O. (s. str.) puncticornis new species; Fig. 197, O. (s. str.)
trepidus new species; Fig. 198, O. (s. str.) patens new species; Fig. 199, O. (s. str.) cavea new species; Fig. 200, O. (s. str.)
sectatus new species; Fig. 201, Middle tibia, male, apical portion, O. (s. str.) sectatus new species.
Revision of Omoglymmius Ganglbauer
203
Plate 17. Figures 202-215, Subgenus Omoglymmius sensu stricto. Figs. 202-208, Head and pronotum, dorsal aspect; Fig.
202, Omoglymmius (s. sir.) ephemeris new species; Fig. 203, O. (s. str.) gracilicornis (Grouvelle); Fig. 204, O. (s. str.)
follis new species; Fig. 205, O. (s. str.) iridescens new species; Fig. 206, O. (s. str.) massa new species; Fig. 207, O. (s. str.)
denticulatus new species; Fig. 208, O. (s. str.) auratus new species; Fig. 209, Middle tibia, male, apical portion; O. (s. str.)
ephemeris new species; Fig. 210, Hind tibia, male, apical portion, O. (s. str.) gracilicornis (Grouvelle); Figs. 211-215,
Head, lateral aspect; Fig. 211, O. (s. str.) iridescens new species; Fig. 212, O. (s. str.) follis new species; Fig. 213, O. (s.
str.) massa new species; Fig. 214, O. (s. str.) denticulatus new species ; Fig. 215, O. (s. str.) auratus new species;
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Bell and Bell
Plate 18. Figures 216-227, Subgenus Omoglymmius sensu stricto. Figs. 216-221, Head and pronotum, dorsal aspect; Fig.
216, Omoglymmius (s. str.) sus new species; Fig. 217, O. (s. str.) planiceps new species; Fig. 218, O. (s. str.) lentus new
species; Fig. 219, O. (s. str.) capito (Grouvelle); Fig. 220, O. (s. str.) cheesmanae (Arrow); Fig. 221, O. (s. str.) asetatus
new species; Figs. 222-227, Head, lateral aspect; Fig 222, O. (s. str.) sus new species; Fig. 223, O. (s. str.) planiceps new
species; Fig. 224, O. (s. str.) lentus new species; Fig. 225, O. (s. str.) capito (Grouvelle); Fig. 226, O. (5. str.) cheesmanae
(Arrow); Fig. 227 , O. (s. str.) asetatus new species;
Revision of Omoglymmius Ganglbauer
205
Plate 19. Figures 228-237, Subgenus Omoglymmius sensu stricto. Figs. 228-233, Head and pronotum, dorsal aspect; Fig.
228, Omoglymmius (s. str.) biroi new species; Fig. 229, O. (s. str.) pulvinatus (Grouvelle); Figs. 230-233, O. (s. str.)
sedlaceki new species; Fig. 230, Mt. Kaindi form A; Fig. 231, Kamira form C; Fig. 232, Kassem form B; Fig. 233,
Sepalakambang form E; Figs. 234-236, Head, lateral aspect; Fig. 234, O. (s. str.) biroi new species; Fig. 235, O. (s. str.)
pulvinatus (Grouvelle); Fig. 236, O. (s. str.) sedlaceki new species; Fig. 237, Head, dorsal aspect, O. (s. str.) sedlaeeki\ new
species, Mt. Otto form D.
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Bell and Bell
Elytron rather long, narrow for subgenus; elytral striae impressed, coarsely punctate; base of Stria IV with very
small glabrous longitudinal scarp; Stria IV with one seta near apex; Stria VII with several near apex; metasternum with
disc entirely coarsely punctate; abdominal sterna with punctures coarse, scattered; lateral pit of female deep, rounded;
anterior femur of male with ventral tooth; middle calcar of male very small; hind calcar of male large, obtuse, its apex
slightly lobate.
The strongly oblique posteriolateral margin of the temporal lobe separates this species from
all other members of the subgenus. The temporal lobes of O. (Boreoglymmius) lewisi (Nakane)
are similarly oblique, but the latter species differs in presence of the basal setae on the
antennae, a much larger postorbital tubercle, and in having the base of the pronotum distinctly
narrowed. Early authors repeatedly confused O. germari with the North American O.
(Boreoglymmius) americanus (Castelnau). The latter species differs in the presence of basal
setae, in having the temporal lobes rounded, and in having the hind calcar of the male acute.
Range. - Widely distributed in Europe, in and south of the Alps, and perhaps isolated, in the
Caucasus and in the Kopet Dagh Mountains of northeastern Iran. Like Rhysodes sulcatus, an
“Urwaldrelikte”, now restricted to small areas of primeval forest. Dajoz (1975) lists localities
and gives a bibliography of published records. In addition to these records, we have examined
specimens from the following localities: FRANCE: one male, Monguillon, Basses Alpes (SATO); IRAN: one
female, Astrabad, Hauser coll. (MNHB); two males, one female, Kopet Dagh, coll. Hauser (CNHM); ITALY: two males,
Lazio, Bassano di Sutri, coll. A. Fiori, P. Luigioni (MNHB); U.S.S.R.: (AZERBAIDZHAN S.S.R.) one male, Lenkoran,
Tangyarud, coll. Kirichenko (LEN); one female, Hamarat, Caspian Sea area, coll. Leder (UVM); (GRUZIAN S.S.R.)
one female, Batum, coll. G. Lewis (BMNH), two females, same locality (MNHN); one male, Lagodekhi Nature Reserve,
coll. Kirekchuk (UVM), one male, one female, same locality (LEN); seven males, four females, Talyche (MNHB);
CAUCASUS (unlocated); two females, Kadi-Keri (MNHB). The specimens listed for UVM were obtained by exchange
from the Academy of Sciences, Leningrad.
Variation. - This species is unusually varied in the development of the postorbital tubercles.
This variation is in part geographical. Almost all specimens from west of the Black Sea have
the tubercle relatively markedly developed. The single exception is the specimen from
Monguillon, France. Most specimens from the Caucasus and eastern Iran have the tubercle
very slightly developed or absent, though a few have it as well developed as in European
specimens. It is possible that more than one species is involved, but further study is necessary.
Bionomics. - Dajoz (1975) summarizes available information as follows (quoted, in
translation): “found in Slovakia in the rotten wood of a beech ( Fagus silvatica L.) invaded by
ants of the genus Camponotus, but this species is not a myrmecophile. In Bulgaria, in the red
bark (?) of an old Oak ( Quercus sp.), larvae and soft imagos in September. In the Gargano
Mountains, found in dead beech wood with its larva. In Greece, at Mount Ossa, at about 1400
meters, in July, we (Dajoz) have found them under the bark of dead beeches in the company of
larvae of Cerambycidae, Buprestidae, and diverse Clavicornia such as Dechomus sulcicollis
Germar and Uleiota planata Fab.”
The specimen from Lagodekhi, U.S.S.R. is labelled as being from beech forest, in old dead
trunk of beech.
Omoglymmius (sensu stricto) laticeps (Bell)
Figs, 114, 122
Omoglymmius laticeps Bell 1977: 157-158.
Omoglymmius (sensu stricto) laticeps (Bell) Bell and Bell 1978.
Type material. - HOLOTYPE female, labelled: “BHUTAN: Nobding, 41 km. E of
Wangdi Phodrang, 2800 m. Basel Natl. Hist. Exped., 1972” (BSL). PARATYPES two
females, labelled: “BHUTAN: Dorju La, Wangdo Phodrang, 1300 m.” (BSL); one female,
Revision of Omoglymmius Ganglbauer
207
labelled: “BHUTAN: Tango, 22 km. N of Thimphu” (BSL).
Description. — Length 6.1 -7.1 mm. Antennal segments impunctate; head relatively short, broad; median lobe short,
broad, obtusely pointed at tip; frontal space short, narrow, nearly V-shaped, its margins only shallowly sinuate; medial
angles rounded, nearly contiguous; posteriomedial margin rounded evenly into posteriolateral margin; orbital groove
replaced by row of punctures, extended nearly to posterior margin; temporal lobe otherwise without punctures; temporal
setae absent; postorbital tubercle minute; in profile view appearing as a slight convexity.
Pronotum with length/greatest width 1.26; widest near middle, apex and base markedly narrowed, lateral margins
markedly curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal at middle; medial margin of
outer carina shallowly sinuate anterior to base; base of outer carina markedly narrowed; inner carina slightly narrowed to
base; inner carina with two or three fine punctures near base; carinae otherwise impunctate except for extremely minute
punctures; median groove coarsely, sparsely punctate; paramedian groove narrow, with two or three coarse punctures;
pronotum without setae; prosternum with fine precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without scarp; Stria IV with one
or two setae near apex; subapical stride with one seta; Stria VII with five to seven setae near apex; metasternum with very
coarse punctures concentrated in midline and near lateral margins; female with lateral pit of abdominal sternum IV deep,
round; female with minute ventral tooth on anterior femur; male unknown.
This species, the only Omoglymmius s. str. known from the Himalaya, is very similar to O.
sakuraii Nakane of the Ryukyu Islands. The two species are similar by combination of a
precoxal carina, a reduced frontal space, very reduced punctation of the pronotal carinae, and a
group of punctures replacing the orbital groove. O. sakuraii differs in having acute medial
angles and in having more setae on Stria IV.
Omoglymmius (sensu stricto) sakuraii (Nakane)
Figs. 115, 123, 129
Rhysodes (Omoglymmius) sakuraii Nakane 1973: 5 (description in Japanese).
Omoglymmius sakuraii Nakane 1978: 130-131 (redescription in English).
Omoglymmius ( sensu stricto) sakuraii (Nakane) Bell and Bell 1978.
Type material. - HOLOTYPE female, Hatsuno, Amami-Oshima, Japan, 30-III- 1 965, S.
Sakurai lgt. (KU). We have not seen the type, but have studied two males from the type
locality (SATO).
Description. — Length 5. 1-5.2 mm. Antennal segments impunctate; head relatively short, broad; median lobe short,
broad, its tip obtuse to subtruncate; frontal space short, narrow, nearly V-shaped, its margin only shallowly sinuate; medial
angles acute, contiguous; posteriomedial angle rounded, in form of very obtuse angle with posteriolateral margin; orbital
groove absent; one to four coarse punctures medial to eye; temporal setae absent, postorbital tubercle minute.
Pronotum longer and narrower than in O. laticeps, length/greatest width 1.43 widest near middle; sides subparallel
except near narrowed base and apex; lateral margin not sinuate anterior to hind angle; carinae subequal at middle; medial
margin of outer carina sinuate anterior to base; base of outer carina narrowed; inner carina narrowed slightly to base; both
pairs of carinae impunctate; median and paramedian grooves narrow, punctured as in O. laticeps ; pronotum without setae;
prosternum with fine precoxal carinae (Fig. 129).
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without trace of scarp; Stria IV
with two to four setae, most specimens with one seta anterior to middle; apical striole without seta; Stria VII with four
setae near apex; metasternum with very coarse punctures concentrated in midline and near lateral margin; abdominal
sterna with punctures very coarse, scattered; female with deep round lateral pit in Sternum IV; male with shallow pit in
this position; male with ventral tooth on anterior femur (condition of anterior femur of female not mentioned in original
description); middle calcar of male minute; hind calcar of male small, triangular, obtuse.
This species is very similar to O. laticeps , but differs in having the pronotum narrower and
more parallel-sided, and the medial angles acute and contiguous.
Omoglymmius (sensu stricto) malabaricus (Arrow)
Figs. 116, 124
Rhysodes malabaricus Arrow 1901: 86-87.
Quaest. Ent., 1982, 18 (1-4)
208
Bell and Bell
Omoglymmius (sensu stricto) malabaricus (Arrow) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) male, labelled: “S. INDIA, Malabar,
Pascoe collection” (BMNH). PARALECTOTYPE one specimen, sex not recorded, same data
as lectotype (BMNH).
Description. — Length 6. 5-7.0 mm. Antennal segments impunctate; head relatively short, broad; median lobe short,
broad, obtusely pointed at apex; frontal space short, broad U-shaped, its lateral margin bent abruptly; temporal lobe
continued anteriorly as oblique, glabrous carina; medial angles rounded, nearly contiguous; posteriomedial margin evenly
rounded into posteriolateral margin; orbital groove fine, obsolescent; temporal lobe with eight to ten coarse punctures near
posterior margin; temporal setae absent; postorbital tubercle minute.
Pronotum with length/greatest width 1.38; widest near middle; apex and base strongly narrowed; lateral margins
curved; lateral margin distinctly sinuate anterior to hind angle; outer carina broader than inner carina; latter distinctly
broader behind than in front of middle; base of outer carina narrowed, narrowly truncate; inner carina narrowed near base;
outer carina with three or four punctures near lateral margin; inner carina impunctate; both median and paramedian
grooves coarsely punctate; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without any trace of scarp;
Stria IV with three to six setae; apical stride without setae; Stria VII with several setae near apex; metepisternum with
coarse punctures near lateral margins, but impunctate in midline; punctures of abdominal Sterna III-V coarse, in form of
irregular transverse rows; lateral pits of Sternum IV of female deep, round; male anterior femur with ventral tooth.
The punctate pronotal grooves and short, broad head make this isolated species superficially
similar to the two preceding ones. However, O. malabaricus differs from them in lacking
precoxal carinae; in having the inner carinae strongly narrowed anteriorly; and in having the
anterior end of the temporal lobe in the form of an oblique ridge. It is the only Omoglymmius s.
str. . in southern India.
Range. - In addition to the type material, we have seen specimens from the following
localities: four females, Anamalai Hills, Cinchona, 1050 m., VI- 1957 coll. P. S. Nathan (BPBM).
Omoglymmius (sensu stricto ) solitarius (Arrow)
Figs. 117, 125
Rhysodes solitarius Arrow 1942: 179.
Omoglymmius (sensu stricto ) solitarius (Arrow) Bell and Bell 1978.
Type material. - HOLOTYPE male, labelled: “ANDAMAN, 76.14” (BMNH).
Description. — Length 6 mm. Antennal Segments V-X punctate; head slightly longer than broad; median lobe
short, narrow, its sides nearly parallel, its apex broadly rounded; frontal space large, V-shaped; its lateral margins only
slightly curved; medial angles produced, their apices obtuse, narrowly separated; posteriomedial margin emarginate;
posteriolateral margin rounded; orbital groove short, ended opposite middle of eye; temporal lobe with nine or 10 shallow
punctures, restricted to lateral third; one temporal seta; postorbital tubercle absent.
Pronotum rather short, length/greatest width 1.24; broadest near middle, margins markedly, evenly curved; base and
apex markedly narrowed; lateral margin not sinuate anterior to hind angle; outer carina only 0.3 as wide as inner carina at
middle; outer carina of nearly even width, markedly curved; its medial margin not sinuate near base; inner carina widest at
middle, tapered anteriorly; narrowed just anterior to base, broadened at extreme base; pronotal carinae entirely
impunctate; pronotal setae absent; precoxal carina absent.
Elytron relatively long, narrow; striae impressed, coarsely punctate; elytral Intervals III, V slightly elevated and
dilated anteriorly; Stria II with three setae near apex; subapical stride with one seta; Stria VII with five or six setae near
apex; metasternum coarsely punctate near lateral and anterior margins and in midline; abdominal Sterna III-V with
scattered coarse punctures; male with shallow lateral pit on Sternum IV (female unknown); male with ventral tooth on
anterior femur; calcar of middle tibia prominent, acute; hind calcar shorter than middle one, triangular, slightly obtuse;
This species can be distinguished from all others in the subgenus by the very narrow, evenly
curved outer carinae. It is the only member of the subgenus from the Andaman Islands, where
there are members of several other subgenera which also have the outer carinae markedly
narrowed.
Revision of Omoglymmius Ganglbauer
209
Omoglymmius ( sensu stricto) philippensis (Chevrolat)
Figs. 118, 126
Rhyzodes philippensis Chevrolat, 1875: 183 (note variant spelling of generic name).
Rhysodes philippensis (Chevrolat) Grouvelle 1903.
Rhysodes philippinensis (Chevrolat) Hincks 1950.
Omoglymmius (sensu stricto) philippensis (Chevrolat) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) male, labelled: “Ins. Philippine,
Chevrolat, Chevl. Or. Typ., philippensis ” (NMW). According to the original description, the
type was received from Henri Deyrolle. PARALECTOTYPES: It is uncertain whether
additional specimens exist from the original type series.
Description. — Length 5. 0-6. 7 mm. Antennal Segments I- IV coarsely punctate; V either punctate or impunctate;
VI-X1 impunctate; head 1.5 longer than broad; median lobe short, narrow, lance-shaped, its apex pointed; frontal space
large, V-shaped, its lateral margin curved; medial angles obtuse, slightly separated; posteriomedial margin oblique,
shallowly emarginate; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, becoming obsolete
near middle of length of eye; temporal lobe with approximately 10 coarse punctures in posteriolateral portion; one
temporal seta; postorbital and suborbital tubercles absent; eye large, round.
Pronotum moderately elongate, length/greatest width 1.27; widest near middle, base and apex narrowed; lateral
margins moderately curved; lateral margin not sinuate anterior to hind angle; carinae subequal at middle; medial margin
of outer carina distinctly sinuate just anterior to base; outer carina widest near middle, distinctly narrowed to base; inner
carina narrowed near base, but extreme base broadened, truncate; outer carina with 10-28 (most specimens 12-15) coarse
punctures; inner carina impunctate or with only one or two punctures; pronotum without setae; prosternum without
precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small longitudinal
pollinose scarp; Stria IV with one to five setae or none (in most specimens with one at base and one or two near apex), in a
few with unbroken row of four or five setae; apical striole with one seta or none; apex of Stria VII with several setae;
metasternum coarsely punctate; punctures of Sterna I1I-V scattered, numerous; female with lateral pit of Sternum IV
deep, round; male with ventral tooth on anterior femur; middle calcar acute, rather small about 0.5 of length of spur; hind
calcar triangular, its apex slightly lobate, obtuse.
This species is the most widespread of a large group of very similar forms, and is the
standard for comparison for them. It is a species with a rather long head, impunctate inner
carina and punctate outer carina, with a very small scarp at the base of Stria IV. Among
species sympatric with it in Luzon, O. imugani is the most similar, but differs in shape of
pronotum and in reduction of punctuation on outer carinae and temporal lobes. O. duplex
differs in having more than one temporal seta and in shape of the head. O. hiekei differs in
having the outer carinae impunctate and in lacking the longitudinal scarp on Stria IV. O.
semperi and O. data have the inner carinae punctate and the eyes reduced. In Palawan, O.
philippensis is sympatric with O. coelebs. The latter species has the outer carinae broad to the
base, and the basal scarp of Stria IV entirely absent. Other Philippine species will be compared
with O. philippensis when they are described below.
Range. - Luzon, Mindoro, Palawan, and Siargao. We have seen the following specimens:
LUZON: one female, Los Banos, coll. Baker (MCZ), one male, same locality, 27-Juli, 1914, leg. Boettcher (MNHB); four
males, Mt. Makiling, Laguna, V-l-31, F. C. Hadden Colin. (CAS); four males, three females, Mt. Makiling, Baker
(NMNH); one male, Pangil, Laguna, V-21-31, F. C. Hadden Colin. (CAS); one male, Quezon Park, Tayabas, alt. 1000
ft., VI-5-32, F. C. Hadden Colin. (BPBM); MINDORO; one male (MNHN); PALAWAN: one female, Binaluan, leg.
Boettcher (MNHB); SIARGAO, two males, Dapa, leg. Boettcher (MNHB).
Variation. - There appears to be considerable variation in development of setae of Stria IV;
however, these are minute and it is difficult to tell whether or not they have been broken off in
some specimens. The specimen from Palawan might be a distinct species; it has five setae in
Stria IV (maximum of four in other O. philippensis ).
Quaest. Ent., 1982, 18 (1-4)
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Bell and Bell
Omoglymmius { sensu stricto) imugani new species
Figs. 119, 127
Type material. - HOLOTYPE female, labelled: “Imugin, N. Viscaya, Baker” (NMNH).
PARATYPES one female, labelled: “Philippinen, Imugan, 6-6-1916, leg. Boettcher”
(MNHB), one male, same data, 16-6-1917 (MNHB), one female, same data, 30-6-1917
(MNHB).
Description. — Length 4. 8-6. 7 mm. Antennal Segments I- 1 1 1 coarsely punctate, Segment IV with few punctures or
impunctate, Segments V-Xl impunctate; head about 1.5 longer than wide; median lobe short, lance-shaped, its apex
obtusely pointed in some specimens, rounded in others; frontal space large, V-shaped; its lateral margin weakly curved;
medial angles obtuse, slightly separated; posteriomedial margin oblique, weakly emarginate; posteriolateral margin evenly
curved, occipital angle distinct; orbital groove fine, tending to be broken into series of punctures; temporal lobe impunctate
except for those in orbital groove; one temporal seta; postorbital and suborbital tubercles absent; eye large, round.
Pronotum moderately long, length/greatest width 1.26; widest near middle; apex more narrowed than base; lateral
margins curved; lateral margin not sinuate anterior to hind angle or else slightly sinuate there; carinae subequal at middle;
medial margin of outer carina slightly sinuate anterior to base; outer carina widest at middle, slightly narrowed to base;
inner carina narrowed near base, at most slightly broadened at base; outer carina with six to nine coarse punctures; inner
carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose scarp;
Stria IV with one seta or none at base and one or two at apex or none; subapical striole with one seta or none; apex of Stria
VII with several setae; metasternum coarsely punctate along margins and in posterior half of midline; anterior half of
midline punctate in some specimens, impunctate in others; disc of metasternum impunctate; punctures of abdominal
Sterna III-V sparse, in form of transverse row on each sternum; female with lateral pit of Sternum IV deep, round; male
with ventral tooth on anterior femur; middle calcar acute; hind calcar triangular, its apex lobate, obtuse.
This species differs from O. philippensis in having the pronotum more narrowed anteriorly,
and in having the punctation reduced, especially on the temporal lobes, outer pronotal carina,
metasternum and abdomen, and in having a wider median lobe. In most respects, O. imugani is
nearly identical to O. politus , but the two differ sharply in male secondary sexual characters;
the former species has the ventral tooth of the anterior femur present and the middle calcar
large, while in the latter species, the femoral tooth is absent and the middle calcar is minute. In
O. imugani , the eye is large, while in O. politus , it is reduced.
Omoglymmius (sensu stricto ) politus new species
Figs. 120, 128
Type material. - HOLOTYPE male, labelled: “Philippinen, Luzon Prov., Lepanto, Mt.
Polis, 2-1917, leg. Boettcher” (MNHB). PARATYPE female, same data as holotype
(MNHB).
Description. — Length 5. 7-6. 6 mm. Antennal Segments I-III coarsely punctate; Segments IV-XI impunctate; head
1 .5 longer than wide; median lobe short, lance-shaped, its apex obtusely pointed, its width greater than in O. imugani ;
frontal space large, V-shaped, its lateral margins feebly curved; medial angles obtuse, slightly separated; posteriomedial
margin oblique, slightly emarginate; posteriolateral margin evenly curved, occipital angle distinct; orbital groove in form of
row of fine punctures; temporal lobe otherwise impunctate; one temporal seta; postorbital, suborbital tubercles absent; eye
slightly reduced.
Pronotum moderately long; length/greatest width 1.29; widest near middle; apex more narrowed than base; lateral
margins curved; shallowly sinuate anterior to hind angle; carinae subequal at middle; medial margin of outer carina
slightly sinuate anterior to base; outer carina widest near middle, narrowed to base; inner carina narrowed near base, but
dilated at extreme base; outer carina with six to 10 coarse punctures; inner carina impunctate; pronotum without setae;
prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with pollinose scarp minute,
indistinct; Stria IV with one seta at apex, none at base; subapical striole with one seta; apex of Stria VII with several setae;
metasternum coarsely punctate near margins and in midline, its disc otherwise impunctate; punctures of abdominal Sterna
III-V sparse, in form of one transverse row on each sternum; female with lateral pit on Sternum IV deep, round; male
without ventral tooth on anterior femur; middle calcar minute; hind calcar as in O. imugani.
Revision of Omoglymmius Ganglbauer
211
This species is very similar to O. imugani except in the secondary sexual characters of the
male. The series of both species at hand are so small that it is impossible to be sure of the
reliability of nonsexual characters. The slightly reduced eye, broader median lobe, more
elongate pronotum with more distinct sinuation of the lateral margin, and broader base of the
inner carina are all possible differences from O. imugani.
Omoglymmius ( sensu stricto ) crassicornis new species
Fig. 130
Type material. - HOLOTYPE female (Negros I.). PHILIPPINES, labelled: “Dumaguete,
P.I., May 1925, John Chapman” (NMNH). PARATYPE female, labelled: “Dumaguete, P.I.,
Horns of Negros, J. W. Chapman Coll.” (MCZ).
Description. — Length 6. 0-6. 5 mm. Outer segments of antennae stouter than in related species; Antennal Segments
I-VI punctate; fine, sparse punctures on Segments VII, VIII in one specimen; IX-XI impunctate; head 1.5 longer than
broad; median lobe narrow, lance-shaped, longer than in O. philippensis, its apex obtusely pointed; frontal space large,
V-shaped, its lateral margin shallowly curved; medial angles obtuse, slightly separated; posteriomedial margin oblique,
posteriolateral margin evenly curved; occipital angle distinct; orbital groove distinct, extended to posterior margin of eye;
temporal lobe with four to nine coarse punctures in posteriolateral portion; one temporal seta present; postorbital and
suborbital tubercles absent; eye large, round.
Pronotum moderately elongate, length/greatest width 1.17 to 1.34; widest near middle, base, apex narrowed; lateral
margins curved, subangulate in middle in one specimen; lateral margin not sinuate anterior to hind angle; carinae subequal
at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest near middle, distinctly
narrowed to base; inner carina narrowed near base, but extreme base broadened, its width nearly equal to width of inner
carina at middle; outer carina with 12-17 coarse punctures; inner carina impunctate or with only one or two coarse
punctures; pronotum without setae; prosternum without procoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small longitudinal
pollinose scarp; Stria I with 0-1 seta near apex; Stria II with one or two setae near apex; Stria IV with one or two setae
near apex; subapical stride with one seta or none; Stria VII with several setae near apex; metasternum with coarse
punctures near margin and in midline, but with disc otherwise impunctate; punctures of Sterna III-V coarse, in part in
form of a single transverse row on each sternum; female with lateral pit on Sternum IV deep, round; male unknown.
This species is the only Omoglymmius s. str. known from the island of Negros. It is similar
to O. philippensis but differs in the following points: punctures of the abdominal sterna form
transverse rows, punctures of the metasternum are restricted to the margins and midline; the
median lobe of the head is longer; punctures of the antenna extend to more distal segments, and
the outer antennal segments are stouter.
Variation. - The two specimens differ markedly in shape of the pronotum. Nevertheless,
they are markedly similar otherwise, and are probably conspecific. Probably one of the two is
an extreme variant. More specimens will have to be collected to determine the limits of
variation.
Omoglymmius (sensu stricto ) amplus new species
Fig. 131
Type material. - HOLOTYPE female, labelled: “SUMATRA, Palembg., Grouvelle”
(LEI). The locality probably refers to the Province, rather than the city of Palembang.
PARATYPE female labelled: “Sumatra, dono Grouvelle, 1901”, (GEN).
Description. — Length 6.5 mm. All antennal segments coarsely punctate; head 1.5 longer than wide; median lobe
short, rather broad, its tip broadly rounded; frontal space large, V-shaped, its lateral margin curved; medial angles obtuse,
narrowly separated; posteriomedial margin oblique, nearly straight; posteriolateral margin evenly curved; occipital angle
distinct; orbital groove fine, extended to posterior margin of eye; temporal lobe with five to seven coarse punctures near
temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Quaest. Ent., 1982, 18 (1-4)
212
Bell and Bell
Pronotum moderately long, length/greatest width 1.20; widest slightly anterior to middle; apex rather suddenly
narrowed; margin markedly oblique near apex; base markedly narrowed; lateral margin strongly sinuate anterior to hind
angle; inner carina slightly narrower than outer one at middle; medial margin of outer carina shallowly sinuate just
anterior to base; outer carina widest at anterior third, constricted opposite lateral sinuation, broadened at extreme base;
base of outer carina directed obliquely laterad; inner carina narrowed just anterior to base; extreme base broadened;
outer carina with 21 punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carina.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose
scarp; Interval V with tubercle laterad to base of Stria IV; Stria II with one or two setae near apex; Stria IV with one
seta near apex; subapical stride without seta; apex of Stria VII with several setae; metasternum with coarse, irregularly
scattered punctures; punctures of abdominal Sterna III-V numerous, scattered; female with lateral pit of Sternum IV
deep, round; male unknown.
This species and the next resemble O. philippensis in most respects, but have the pronotum
broadest anterior to the middle. Of the other species from Sumatra, O. summissus has a
subquadrate pronotum, punctate inner carinae, and lacks a longitudinal basal scarp on Stria
IV; O. fraudulentus likewise lacks the scarp, and has the temporal lobes very coarsely punctate;
O. bouchardi has the pronotum nearly impunctate, and has the punctures of the abdominal
sterna largely confluent.
Omoglymmius (sensu stricto) modiglianii new species
Fig. 132
Type material. - HOLOTYPE female, labelled: “MENTAWEI, Si Oban, IV-VIII,
Modigliani 94” (GEN). PARATYPE female (teneral), same data as holotype (GEN).
Description. — Length 5. 7-6.0 mm. All antennal segments punctate; punctures coarse on proximal segments, finer
on distal segments; head 1.5 longer than wide; median lobe short, its tip broadly rounded; frontal space large, nearly
U-shaped, its lateral margin more markedly curved than in O. antplus\ medial angles nearly rectangular, narrowly
separated; posteriomedial margin oblique, slightly sinuate; posteriolateral margin evenly curved; occipital angle distinct;
orbital groove fine, extended to posterior margin of eye; temporal lobe with 13-15 very coarse punctures in its lateral half;
one temporal seta; postorbital, suborbital tubercles absent; eye large, rounded.
Pronotum moderately elongate; length/greatest width 1.26, widest anterior to middle, apex rather suddenly narrowed;
margin oblique near apex; base strongly narrowed; lateral margin sinuate anterior to hind angle; inner carina slightly
narrower than outer one at middle; medial margin of outer carina shallowly sinuate just anterior to base; outer carina
widest at anterior third, scarcely constricted opposite lateral sinuation; base of outer carina not turned obliquely laterally;
inner carina narrowed just anterior to base, but extreme base broadened; outer carina with about 20 coarse punctures;
inner carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with small longitudinal scarp;
Interval V laterad to base of Stria IV with low tubercle; Stria I with one seta near apex or none; Stria II with up to two
near apex or none; Stria IV with one seta near apex; subapical stride with one seta; apex of Stria VII with several setae;
metasternum with coarse, irregularly distributed punctures; punctures of abdominal Sterna III-V numerous, scattered;
female with lateral pit of Sternum IV deep, round; male unknown.
This species is the only member of the subgenus known from the Mentawei Islands. It is
close to O. ampins , but differs in having the pronotum more elongate, its lateral margin less
sinuate, the base of the outer carina not turned outwards, and the temporal lobe with the
punctures coarser and more numerous.
Omoglymmius ( sensu stricto) evasus new species
Fig. 134
Type material. - FIOLOTYPE male, labelled: “E. slope Mt. McKinley, Davao Prov.,
MINDANAO, elev. 3300 ft., IX-28-1946, F. G. Werner leg., lot #92, dead stump, CNHM
Phil. Zool. Exp. (1946-47)” (CNHM). PARATYPES one female, same data as holotype
(CNHM); one male, three females, same locality as holotype, IX- 1946, H. Hoogstrall leg.
(CNHM).
Revision of Omoglymmius Ganglbauer
213
Description. — Length 5. 0-7. 2 mm. Antennal Segments I - 1 1 1 coarsely punctate; Segment IV with a few, rather fine
punctures; Segments V-XI impunctate; head 1.3 longer than broad; median lobe rather short, its apex broadly rounded;
frontal space moderate-sized, V-shape, its lateral margins curved; medial angles obtuse, slightly separated; posteriomedial
margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, extended to hind margin
of eye; temporal lobe with three to nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital
tubercles absent; eye large, round.
Pronotum moderately elongate; length/greatest width 1.34, widest near middle, base and apex narrowed; apex much
less narrowed than in O. philippensis\ lateral margins moderately curved; lateral margin shallowly sinuate anterior to hind
angle; carinae subequal in middle; medial margin of outer carina sinuate just anterior to base; outer carina widest in
anterior third, transversely truncate at apex; inner carina narrowed near base, but extreme base broadened, truncate; outer
carina with 15-18 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal
carinae.
Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
II with one seta near apex; Stria IV with one seta near apex; most specimens with one seta at base of Stria V near
humerus; subapical stride without setae; Stria VII with several setae near apex; metasternum with coarse punctures near
margin and scattered near midline; disc otherwise impunctate; punctures of abdominal Sterna III-V coarse, sparse,
scattered; female with lateral pit of Sternum IV round, rather shallow; male with small, obtuse ventral tooth on anterior
femur; middle calcar prominent, acute; hind calcar triangular, its apex obtuse.
This species differs conspicuously from O. philippensis in the shape of the pronotum,
particularly of the outer carinae, which are truncate anteriorly. Absence of the longitudinal
scarp at the base of Stria IV, the smaller number of punctures on the temporal lobes, and
metasternum are among numerous additional differences. The truncate apex of the outer carina
will separate O. evasus from similar species in nearby regions, although O. nemoralis of
Sarawak approaches it. The latter species has the setae of the elytra much more numerous, and
the punctures of the outer carina fewer in number and limited to its posterior portion.
Range. - Mindanao, Philippine Islands. In addition to the type material, we have seen the
following specimens: two males, two females, Baracatan, 1500 m., June 27-29-1977, M. Sato leg. (SATO); two
females, Mainit Hot Spring, 28- V 1 1- 1 970, M. Sato leg. (SATO); one male, four females, Todaya, July 30, 1970, M. Sato
leg. (SATO).
Omoglymmius (sensu stricto) nemoralis new species
Fig. 135
Type material. - HOLOTYPE male, labelled: “Mt. Matang, W. Sarawak, 17-1-14, G.
Bryant Coll. 1919-147” (BMNH). PARATYPES one male, same data as holotype (BMNH);
one male, labelled: “Sarawak. Matang Riv., 3 1/2 mile, J.E.A. Lewis, 1910-116” (BMNH);
one female, Semengoh For. Res., 15 mi. s. Kuching, 22-IX-1966, J.F.G. Clarke, Thelma M.
Clarke (NMNH).
Description. — Length 5. 6-6.0 mm. Antennal Segments I- V III coarsely punctate; Segments IX-XI impunctate;
head 1.5 longer than broad, distinctly more elongate than in O. evasus\ median lobe rather narrow, its apex obtusely
pointed; frontal space moderate-sized, V-shaped, its lateral margins curved; medial angles obtuse, slightly separate;
posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, not
extended to hind margin of eye, interrupted in some specimens; temporal lobe with seven to nine coarse punctures in lateral
portion; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately elongate; length/greatest width 1.26; widest near middle; base, apex narrowed; lateral margins
moderately curved; lateral margin shallowly sinuate anterior to hind angle; carinae subequal in width at middle; medial
margin of outer carina scarcely sinuate anterior to base; outer carina widest in anterior third, transversely truncate at apex;
inner carina narrowed near base, but extreme base broadened, truncate; outer carina with nine to 16 coarse punctures,
anterior 0.25-0.5 of outer carina impunctate; inner carina impunctate; pronotum without setae; prosternum without
precoxal carinae.
Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
II with two setae near apex; Stria IV with six or seven setae forming uninterruped row from base to apex; base of Stria V
without seta; subapical striole with one seta or none; Stria VII with several setae near apex; metasternum coarsely
punctate, with most punctures near midline or margins; punctures of Sterna III-V coarse, sparse, scattered; female with
lateral pit of Sternum IV round, moderately deep; male with small, obtuse ventral tooth on anterior femur; middle calcar
Quaest. Ent., 1982, 18 (1-4)
214
Bell and Bell
of male acute, smaller than in O. evasus\ hind calcar triangular, apex obtuse.
This species is allopatric with its closest ally, O. evasus of Mindanao. From the latter species
it can be separated by the more numerous elytral setae and the absence of punctures from the
anterior part of the outer carina. O. fraudulentus of Borneo and Sumatra and O. malaicus of
Malay Peninsula are also very similar. In the former species, the outer carinae are markedly
divergent at their bases, while in the latter, the pronotum is narrower, with its margins more
nearly parallel, with apex of the median lobe rounded. The only other member of the subgenus
known to occur on Borneo is O. consors. The latter species differs strikingly in completely
impunctate outer carinae and the presence of a transverse band of pollinosity on the basal scarp
of the elytron.
This species is possibly O. borneensis (Grouvelle) 1903, p. 119-120). We have been unable
to locate type material for O. borneensis (Type locality R. Sambey, coll. Ledru), indicated as
being in the Oberthiir Collection. There are several points of disagreement between the original
description and O. nemoralis, and it is quite possible that Grouvelle’s name applies to O.
consors , O. fraudulentus, or to some other member of the subgenus, not yet found in Borneo.
Omoglymmius (sensu stricto) fraudulentus new species
Fig. 136
Type material. - HOLOTYPE male, labelled: “SUMATRA, Palembg” (MNHN);
PARATYPES three males, eight females, labels identical to holotype (MNHN).
Description. — Length 5. 5-6. 4 mm. Antennal Segments I-1V coarsely punctate; a few punctures on V, VI;
Segments VII-XI impunctate; head 1.5 longer than broad; median lobe rather narrow, its apex obtusely pointed; frontal
space moderate-size, V-shaped, its lateral margins curved; medial angles obtuse, slightly separated; posteriomedial margin
less distinctly oblique than in the two preceding species, posteriolateral margin evenly curved; occipital angle indistinct;
orbital groove fine, interrupted; temporal lobe with six to 10 coarse punctures in lateral portion; one temporal seta;
postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately elongate; length/greatest width 1.29; widest near middle; apex narrowed; base slightly narrower
than greatest width; lateral margins moderately curved; lateral margin markedly sinuate anterior to hind angle; carinae
subequal in width at middle; outer carinae abruptly bent near base, strongly divergent there; outer carina widest anterior
to middle, narrowed, obtusely rounded at apex; outer carina with three or four (Borneo specimen) or 14-16 (Sumatra
specimens) coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal carina.
Elytron relatively narrow, long, striae impressed, coarsely punctate; base of Stria IV wihout longitudinal scarp; Stria II
with two setae near apex; Stria IV with complete series of five setae; subapical stride with one seta; Stria VII with several
setae near apex; metasternum coarsely punctate; punctures of abdominal Sterna II1-V coarse, rather dense; female with
lateral pit on Sternum IV round, rather shallow; male with small, obtuse ventral tooth on anterior femur; calcars as in O.
nemoralis.
This species is distinguished from its closest relatives by the sharply divergent bases of the
outer carinae. Among the other members of the subgenus on Sumatra, O. consors comes
closest, but differs in having a transverse band of pollinosity on the basal scarp of the elytron,
and in having the outer carina completely impunctate. In Borneo, the most similar species is O.
nemoralis , which does not have the bases of the outer carinae divergent.
Range. - Sumatra and Borneo. In addition to the type material, we have seen one female
from Sumatra, labelled: “Tebbing Tinggi, VI-37 , Brues” (MCZ), and one male from Borneo, labelled: “Miri Riv.
VIII, Mjoberg Colin., W. 1S/I. Funge Bequest-’ (CAS). The latter specimen differs from those of Sumatra having only three
or four punctures on the outer carina. It does not seem to differ otherwise, and is provisionally assigned to this species.
Omoglymmius (sensu stricto ) malaicus (Arrow)
Fig. 133
Rhysodes malaicus Arrow 1901: 85-86.
Revision of Omoglymmius Ganglbauer
215
Omoglymmius ( sensu stricto) malaicus (Arrow) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) male, labelled: “Penang, Lamb, Pascoe
Coll.” (BMNH). PARALECTOTYPE female, same label as lectotype (BMNH). Penang is a
small offshore island near the Malay Peninsula.
Description. — Length 6.0 mm. Antennal Segments I-V punctate, punctures of Segment V very small; Segments
VI-XI impunctate; head 1.5 longer than broad; median lobe rather narrow, its sides parallel, its apex rounded; frontal
space moderate-size, U-shaped, its lateral margins curved; medial angles obtuse, very narrowly separated; posteriomedial
margin oblique; posteriolateral margin evenly curved; occipital angle distinct; orbital groove fine, interrupted; temporal
lobe with seven to nine coarse punctures in lateral portion; one temporal seta; postorbital, suborbital tubercles absent; eye
large, round.
Pronotum elongate, length/greatest width 1.43; widest slightly anterior to middle, narrowed at apex, scarcely
narrowed to base, lateral margins nearly parallel; lateral margin not sinuate anterior to hind angle; carinae subequal in
width at middle; outer carinae not divergent at base; inner carina constricted anterior to base, latter broadened; outer
Carina with 10-14 coarse punctures; inner carina impunctate; pronotum without setae; prosternum without precoxal
carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
1 1 with two setae near apex; Stria IV with complete series of four to seven setae; subapical striole with one seta; Stria VII
with several setae near apex; metasternum coarsely punctate; punctures of abdominal Sterna 1 1 I-V coarse, rather dense;
female with lateral pit round, rather deep; male with ventral tooth on anterior femur more prominent than in preceding
species; middle calcar acute, prominent; hind calcar triangular, its apex obtuse.
This is the only member of the subgenus known from the Malay Peninsula. It is closest to O.
fraudulentus of Sumatra, but differs in shape of the pronotum, and lacks the divergent bases of
the outer carinae which are so prominent in the latter species.
Range. - Malay Peninsula and offshore islands. In addition to type material, we have seen
the following specimens: one female, labelled: “Poulo Penang, Raffray, compared with type, G.J.A., correctly
named, G.J.A.” (MNHN); one specimen, sex not recorded, labelled “Penang-Bryant” (BMNH); one specimen, sex not
recorded, labelled “Perak, Doherty” (BMNH).
Omoglymmius (sensu stricto) coelebs new species
Fig. 137
Type material. - HOLOTYPE male, labelled: “N. Palawan, P. I. Binaluan,
Nov.-Dez.-1913, leg. G. Boettcher, from H. P. Loding, 1935” (NMNH).
Description. — Length 6.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-VII with few punctures;
Segments VIII-XI impunctate; head 1.5 longer than broad; median lobe with margins nearly parallel, apex rounded;
frontal space moderate in size, U-shaped, its margins curved; medial angles obtuse, seperated; posteriomedial margin
oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove fine, not attaining posterior margin
of eye, interrupted; temporal lobe with eight or nine coarse punctures in lateral portion; one temporal seta; postorbital,
suborbital tubercles absent; eye large, round.
Pronotum moderately elongate, length/greatest with 1.26; base, apex narrowed; lateral margins moderately curved;
lateral margin not sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina
scarcely sinuate anterior to base; outer carina of almost even width, transversely truncate at both apex and base; inner
carina constricted just anterior to base; latter broad, truncate; outer carina with 15-19 coarse punctures; anterior 0.25 of
outer carina impunctate; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively narrow, long; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria I
with one seta near apex; Stria II with one to three setae near apex; Stria IV with complete series of four or five setae;
apical striole with one seta; Stria VII with several setae near apex; metasternum with coarse punctures near anterior and
lateral margins, otherwise nearly impunctate; punctures of Sterna 1 1 I-V coarse, rather dense, forming broad transverse
band on each sternum; male with small, obtuse ventral tooth on anterior femur; middle calcar acute, small; hind calcar
triangular, its apex obtuse, slightly lobate; female unknown.
This species is closest to O. evasus and O. nemoralis, from which it differs conspicuously in
having the outer carinae not narrowed posteriorly. On Palawan it is sympatric with O.
philippensis. The latter species is longer and narrower, and has the pronotum narrowed both at
the base and apex. In addition, it has a small longitudinal scarp at the base of Stria IV.
Quaest. Ent., 1982, 18 (1-4)
216
Bell and Bell
Omoglymmius (sensu stricto) thoracicus new species
Figs. 138, 139
Type material. - HOLOTYPE male (JAVA), labelled: “Tangk.-Prahoe, Juni- 1933, V.
Doesburg” (LEI).
Description. — Length 5.9 mm. All antennal segments coarsely punctate; head 1.5 longer than wide; median lobe
oval, its apex broadly rounded; frontal space broad, U-shaped, its lateral margins abruptly bent; medial angles obtuse,
slightly separated; posteriomedial margin oblique; posteriolateral margin evenly rounded; occipital angle distinct; orbital
groove absent; temporal lobe with nine to 12 rather fine punctures in posteriolateral half; temporal seta absent; postorbital
and suborbital tubercles absent; eye large, round.
Pronotum moderately elongate; length/greatest width 1.24; widest near middle; base and apex narrowed; lateral
margins strongly curved; lateral margin slightly sinuate anterior to hind angle; carinae subequal in width at middle; medial
margin of outer carina distinctly sinuate just anterior to base; outer carina widest anterior to middle, narrowed to base and
apex; inner carina narrowed just anterior to base, latter broad, truncate; outer carina with 13-16 rather fine punctures;
inner carina impunctate; pronotum without setae; prosternum with sharp precoxal carina on each side, extended 0.75 of
distance from coxa to anterior margin (Fig. 1 39).
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Striae
I, II without setae; Stria IV with one seta near apex; subapical stride with one seta; apex of Stria VII with several setae;
metasternum coarsely punctate; punctures of abdominal Sterna 1II-V scattered, numerous; male with small, obtuse,
ventral tooth on anterior femur; middle calcar acute, subequal to spur; hind calcar small, triangular, obtuse, margin
emarginate between calcar and tibial spur; female unknown.
In dorsal aspect this species looks like O. philippensis and its relatives, except that the
punctures of the temporal lobes and outer carinae are finer and the temporal seta is absent. In
ventral view it is easily distinguished by well-developed precoxal carinae. It shares this
character with another Javanese species, O. pectoralis. The latter species, however, entirely
lacks punctures on the outer carinae, and has four temporal setae.
Omoglymmius ( sensu stricto ) summissus new species
Fig. 140
Type material. - HOLOTYPE female, labelled: “SUMATRA, Palembg.” (MNHN). (The
type was part of a group of specimens of several species mounted on a single pin; consequently,
the label on the holotype was copied by us.)
Description. — Length 6.1 mm. Antennal Segments I-VI1 coarsely punctate; Segments VIII-XI impunctate;
median lobe short, lance-shaped, its apex obtuse; frontal space large, V-shaped, its lateral margins only slightly curved;
medial angles very obtuse, separated; posteriomedial margin slightly oblique; posteriolateral margin evenly rounded;
occipital angle indistinct; trace of orbital groove present near anterior margin of eye, at junction with distinct preorbital
pit; temporal lobe with 10-14 coarse punctures in posteriolateral half; one temporal seta; postorbital, suborbital tubercles
absent; eye large, round.
Pronotum moderately elongate; length/greatest width 1.31; widest near middle; base slightly narrowed; apex narrowed
abruptly; lateral margins nearly straight, almost parallel; lateral margin not sinuate anterior to hind angle; inner carinae
slightly wider than outer ones at middle; medial margin of outer carina not sinuate anterior to base; outer carina widest
anterior to middle; its apex obliquely truncate; inner carina narrowed near base, broadened, truncate at base; outer carina
with 12-18 coarse punctures; inner carina with four or five punctures in posterior half; pronotum without setae; prosternum
without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with slight suggestion of
longitudinal scarp, but latter entirely glabrous; Stria II with two setae near apex; Stria IV with three setae in anterior
third, one near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum entirely coarsely
punctate; punctures of abdominal Sterna III-V rather fine, scattered, not at all confluent; female with deep, round lateral
pit on Sternum IV; male unknown.
This species differs from O. philippensis and similar species by punctures on the inner
carina. Shape of the pronotum is closest to that of O. malaicus. The latter species has a longer,
narrower pronotum with outer carinae more narrowed anteriorly, and without punctures on the
inner carina. Among Sumatran species, O. consors and O. bouchardi have both inner and outer
Revision of Omoglymmius Ganglbauer
217
carinae impunctate, while O. fraudulentus has only the outer carinae punctate. The latter
species also differs from O. summissus in having the bases of the outer carinae divergent.
Omoglymmius (sensu stricto ) semperi new species
Fig. 141. 147
Type material. - HOLOTYPE male, labelled: “Phillip. Islands, Semper” (MNHN)
PARATYPES two females, with labels identical to that of holotype and the additional label
“Fry Colin. 1905-100” (BMNH).
Description. — Length 5. 4-6. 4 mm. Antennal Segments Ml coarsely punctate; Segment III with a few fine
punctures; Segments 1V-IX impunctate; head 1.5 longer than wide; median lobe short, lance-shaped, its apex acute; frontal
space V-shaped, its margins scarcely curved; medial angles obtuse, well separated; posteriomedial margin oblique, slightly
emarginate; posteriolateral margin evenly curved; occipital angle distinct; orbital groove narrow but distinct, extended
posteriorly at least to middle of eye; temporal lobe with nine to 12 coarse punctures in posteriolateral portion; one temporal
seta; postorbital, suborbital tubercles absent; eye reduced, its depth about 0.6 of depth of head, its posterior margin
oblique, nearly straight.
Pronotum elongate; length/greatest width 1.44; widest near middle, base, apex narrowed; lateral margin curved,
scarcely sinuate anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate
anterior to base; outer carina widest near middle, its apex markedly narrowed; inner carina narrowed just anterior to base;
latter broadly truncate; outer carina with 25-30 coarse punctures; inner carina with 10-14 coarse punctures concentrated
in anterior and posterior thirds of carinae, middle third impunctate or nearly so; pronotum without setae; prosternum
without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with small, pollinose longitudinal
scarp; Stria II without setae; Stria IV with one seta at apex; subapical stride with one seta; Stria VII with several setae
near apex; metasternum entirely coarsely punctate; punctures of abdominal Sterna 1II-V coarse, rather dense, those of 111
in form of irregular transverse row; those of IV, V scattered; female with deep round lateral pit on Sternum IV; male with
small obtuse ventral tooth on anterior femur; middle calcar acute, very small, less than 0.3 as long as spur; hind calcar
triangular, its apex obtuse, slightly lobate, its upper margin angulate.
This species and the next one differ from O. philippensis by punctures on the inner carina
and by reduction of the eyes. In this species, the eyes are only slightly reduced in depth, but are
sharply reduced in length, with the posterior margin forming an oblique, nearly straight line.
This species is dedicated to the collector, Professor Carl Semper (1832-1893) of the
University of Wurzburg, whose distinguished career included several years of pioneer collecting
in the Philippines. It is not known on what island he collected this species. Its similarity to O.
data suggests that it might be a localized montane form from northern Luzon.
Omoglymmius ( sensu stricto ) data new species
Figs. 142, 148
Type material. - HOLOTYPE male, labelled: “P.I., Mt. Data, Mt. Province, 5000 ft.,
Ill- 1 6- 193 1 , F.C. Hadden Colin.” (CAS). PARATYPE female, labelled: “Philippines, Mt.
Prov., Abatan, Buguias, 60 km. S. of Bontoc, 1800-2000 m., 9-VI-1964, coll. H.M. Torrevillas”
(BPBM).
Description. — Length 6. 0-6. 6 mm. Antennal Segments 1, II coarsely punctate; Segments 1I1-XI impunctate; head
1.5 longer than wide; median lobe lance-shaped, notched at junction of clypeus; apex obtuse; frontal space V-shaped, its
margins sharply curved; medial angles obtuse, contiguous; posteriomedial margin slightly curved; posteriolateral margin
evenly curved; occipital angle very indistinct; orbital groove absent; temporal lobe with 25-30 rather fine, scattered
punctures; one temporal seta; postorbital, suborbital tubercles absent; eye strongly reduced, its depth less than 0.5 of depth
of head, its posterior margin curved.
Pronotum moderately elongate; length/greatest width 1.21; widest near middle; base moderately narrowed; apex
strongly narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; carinae subequal in width at
middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, its apex
narrowed; inner carina only slightly constricted just anterior to base, latter broadly truncate; outer carina with 25-30
Quaest. Ent., 1982, 18 (1-4)
218
Bell and Bell
rather fine punctures; inner carina with eight to 10 punctures; pronotum without setae; prosternum without precoxal
carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal carina;
Stria II without setae; Stria IV with one apical seta; subapical striole without seta; apex of Stria VII with several setae;
metasternum entirely coarsely punctate; punctures of abdominal Sterna as in O. semperi\ female with round lateral pit
on Sternum IV; male with small obtuse ventral tooth on anterior femur; middle calcar minute; hind calcar triangular,
its apex obtuse, its upper margin not angulate.
This species is similar to O. semperi in most characters, bur differs in form of the eye, which
is rounder and more reduced, in the shorter, broader pronotum, and in shape of the hind calcar
of the male.
Omoglymmius (sensu stricto ) hiekei new species
Fig. 143
Type material. - HOLOTYPE male, labelled: “Philippinen, Luzon, Dalbalan (underside
label), S. Boettcher, 3-1-17” (MNHB). PARATYPES one female, same data as holotype
(MNHB; one male, same locality and collector as holotype (underside label), dated 1-1917
(MNHB); one female, labelled “Mt. Makiling, Luzon, Baker” (MCZ).
Description. — Length 6. 0-6. 5 mm. Antennal Segments 1, II punctate; Segments III-XI impunctate; head about
1.5 longer than wide; median lobe narrow, lance-shaped, its apex obtusely pointed; frontal space large, U-shaped, its
lateral margin curved; medial angles obtuse, slightly separated; posteriomedial margin oblique, slightly emarginate;
posteriolateral margin evenly curved; occipital angles distinct though slight; temporal lobe with orbital groove extended as
irregular row of punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum long; length/greatest width 1.40; widest near middle; both ends narrowed; apex slightly more so than base;
lateral margins curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal in width at middle; medial
margin of outer carina slightly sinuate anterior to base; outer carina widest at middle, narrowed to base and apex; inner
carina narrowed just anterior to base; latter broadly truncate; outer carina impunctate or with one or two minute punctures
near base; inner carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
II with two setae near apex; Stria IV with five setae in continous series; subapical Stria with one seta; apex of stria VII
with several setae; metasternum with coarse punctures near margins and midline, and few scattered on disc; punctures of
abdominal Sterna III-V sparse, in form of irregular transverse row on each sternum; female with lateral pits of Sternum
IV deep, round; male with very small, obtuse ventral tooth on anterior femur; middle calcar acute, prominent, nearly as
long as spur; hind calcar triangular, its tip obtuse, slightly lobate.
This species is characterized by the absence or near absence of punctures from the pronotal
carinae. It is most similar to O. imugani and O. politus , but the latter two species have six to 10
conspicuous punctures on the outer carina. In addition, O. imugani has very few setae on the
elytra, while O. politus has slightly reduced eyes and the male lacks the ventral tooth on the
anterior femur.
We dedicate this species to Dr. Fritz Hieke, of the Museum fur Naturkunde der
Humboldt-Universitat, in token of his help in this work, in arranging for us to borrow the
historically important rhysodid collection of his museum.
Omoglymmius ( sensu stricto) pect oralis new species
Figs. 144, 149
Type material. - HOLOTYPE female, labelled: “JAVA, Mt. Smetou” (label is
handwritten, and spelling of place name may be inaccurate)(MNHN).
Description. — Length 7.0 mm. Antennal Segments I- 1 1 1 coarsely punctate; Segments IV-XI impunctate; head
1 .25 longer than broad; median lobe short, oval, its apex obtusely pointed; frontal space rather narrow, its sides curved;
medial angles nearly rectangular, contiguous; posteriomedial margin slightly curved; posteriolateral margin evenly curved;
occipital angle indistinct; orbital groove absent; temporal lobe impunctate; four temporal setae; antennal lobe elevated
above anterior end of temporal lobe: postantennal groove represented by broad pollinose scarp; postorbital, suborbital
Revision of Omoglymmius Ganglbauer
219
tubercles absent; eye large, round.
Pronotum rather short, length/greatest width 1.19; subquadrate, widest near middle; base, apex scarcely narrowed;
lateral margins only slightly curved; lateral margin not sinuate anterior to hind angle; outer carina about 0.6 as wide as
inner one at middle; inner carina only slightly narrowed anterior to base; latter broadly truncate; both pairs of carinae
impunctate; marginal groove dilated; pronotum without setae; prosternum with sharp precoxal carina on each side,
extending 0.75 of distance from coxa to anterior margin (Fig. 149).
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp;
Stria IV with two setae near base and two near apex; subapical stride with 1 seta; Stria VII with several setae near
apex; metasternum impunctate except for row along each lateral margin; abdominal Sterna 1II-V each with coarse
punctures tending to form transverse row; female with deep, reniform lateral pit on Sternum IV; male unknown.
This species resembles O. thoracicus, also of Java, in having precoxal carinae, but is
otherwise very different. In dorsal aspect it resembles O. quadruplex of Mindanao, but the
pronotum is more quadrate, and the anterior ridge of the temporal lobe less extensively
pollinose.
Omoglymmius (sensu stricto ) quadruplex new species
Fig. 145
Type material. - HOLOTYPE male, labelled: “P.I., Kabasalan, Zamboanga, 20 ft.,
III-9-32, coll. H.C. Muzzell, F.C. Hadden Colin.” (BPBM). PARATYPES seven males, one
female, same data as holotype (BPBM); two males, one female, same data as holotype, no
collector cited (BPBM); one male, three females, same locality as holotype, V-4-1932, no
elevation given (CAS); one male, same locality as holotype, F. C. Hadden Colin., no collector
cited, VI- 1-32 (CAS).
Description. — Length 5.2-7. 1 mm. Antennal Segments I-V sparsely, coarsely punctate; Segments VI-XI
impunctate; head slightly longer than broad; median lobe short, oval, its tip obtusely pointed; frontal space rather narrow,
its sides curved; medial angles rectangular, slightly separated; posteriomedial margin slightly emarginate; posteriolateral
margin evenly curved; occipital angle indistinct; antennal ridge elevated above level of temporal lobe; latter forming
narrow, pollinose ridge anteriorly; temporal lobe impunctate; orbital groove broad, extended beyond middle of eye;
temporal setae four in most specimens (three present in a few specimens); two pairs of gular setae present; postorbital,
suborbital tubercles absent; eye large, round.
Pronotum rather broad, length/greatest width 1.23; widest near middle; base moderately narrowed, apex more
narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; outer carina slightly broader than
inner one at middle; medial margin of outer carina sinuate anterior to base; inner carina constricted near base, latter
broad, truncate; both pairs of carinae impunctate; marginal grooves not dilated; pronotum without setae; prosternum
without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; basal margin of elytron completely pollinose
medial to base of Stria IV; base of latter without longitudinal scarp; Stria I with one to five setae near apex; Stria II with
one seta near apex, Stria IV with one seta near base, one or two setae near apex; subapical stride with one seta; apex of
Stria VII with several setae; metasternum coarsely, sparsely punctured near lateral margins and midline; disc otherwise
impunctate; abdominal Sterna II I-V each with punctures in form of transverse row; lateral portions of Sterna 1 1 I-V I with
punctures confluent, forming irregular transverse impressions; lateral pit of Sternum IV of female scarcely distinct;
anterior femur of male with prominent, acute, ventral tooth; middle calcar acute, nearly as long as spur; hind calcar
broadly rounded, raised well above spur.
This species, O. pectoralis and O. duplex form a well-marked group with more than one
temporal seta, the head broad, and the glabrous part of the temporal lobe separated from the
antennal ridge by an elongate, pollinose ridge. From the first species, O. quadruplex is
separated by the absence of precoxal carinae; from the second, by the greater number of
temporal setae, the less extensive orbital groove, the extragular setae, several setae on the apex
of Stria I, pollinosity on the basal margin medial to Stria IV.
Range. - Mindanao in the Philippines. In addition to the type material we have seen the
following specimens from eastern Mindanao: one male, Lawa, Davao Prov., IV-24-30. coll. C. F. Clagg, F.
Psota Colin, “at light”(CNHM); two males, Madaum, Tagum, Davao Prov., sea level, F. G. Werner leg., CNHM Philip.
Zool. Exp. 1946-47 (CNHM).
Quaest. Ent., 1982, 18 (1-4)
220
Bell and Bell
Omoglymmius (sensu stricto) duplex new species
Fig. 146
Type material. - HOLOTYPE male, labelled: “Mt, Makiling, Laguna, P.I., 1-1-1931, F. C.
Hadden Colin.” (CAS). PARATYPES three males, six females, same data as holotype (CAS);
the following paratypes all have the same locality data as the holotype but with these dates: one
female, IV- 1 - 193 1 ; one female, IV-24-1931; one male, VI-1-31; one female, VI-19-31; one
male, one female, VI-29-31; one male, 3-3-32, 4000 ft., one male, II 1-5-32; two males, four
females, IV- 1-32; two males, three females, IV- 1 -32, 400 ft.; one male, one female, V-5-32, 800
ft.; one male, VII- 1-32, 400 ft. (all CAS); one male, one female, same locality data as holotype,
IV-6-32, 500 ft. (BPBM); one female, same locality data as holotype, VI-1-32, 500 ft.
(BPBM); two females, Mt. Makiling, Luzon, coll. C. F. Baker (NMNH); one male, two
females, same data as previous specimens, July 1923 (NMNH); one male, one female, Mt,
Makiling, Luzon, P.I., coll. H. C. Muzzall, F. C. Hadden Colin. ,B. Malkin Colin., XII-6- 1931
(CNHM).
Description. — Length 5. 7-7. 8 mm. Antennal Segments I- IV sparsely, coarsely punctate; Segments V-Xl
impunctate; head slightly longer than broad; median lobe short, oval, its tip obtusely pointed; frontal space rather broad,
U-shaped, its lateral margins strongly curved; median angles obtuse, contiguous; posteriomedial margin oblique;
posteriolateral margin evenly curved; occipital angle relatively distinct; antennal ridge elevated above level of temporal
lobe; anterior end of latter forming narrow, pollinose ridge; temporal lobe impunctate or with few minute punctures;
orbital groove represented by broad, rather irregular band of pollinosity which reaches posterior margin of temporal lobe;
temporal setae two in most specimens three or one in a few specimens; one pair gular setae present; postorbital, suborbital
tubercles absent; eye large, round.
Pronotum moderately elongate, length/greatest width 1.31; widest near middle; base moderately narrowed; apex more
narrowed; lateral margins curved; lateral margin not sinuate anterior to hind angle; medial margin of outer carina sinuate
anterior to base; carinae subequal at middle; inner carina constricted at base, latter broad, truncate; outer carinae
impunctate in most specimens (four to six punctures in the specimen from Mindanao); marginal grooves not or but slightly
dilated; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp of elytron not pollinose; base of Stria
IV with small longitudinal pollinose scarp; setae absent from apex of Stria I; Stria 1 1 with one seta near apex; Stria IV with
one to three near apex, but none near base; subapical stride with one seta; apex of Stria VII with several setae;
metasternum coarsely, sparsely punctate; abdominal Sterna III-V with coarse, sparse punctures; lateral portions of Sterna
I1I-VI with punctures confluent, in form of irregular impressions; lateral pit of Sternum IV of female scarcely distinct;
anterior femur of male with prominent, acute, ventral tooth; middle calcar acute, about 0.5 as long as spur; hind calcar
triangular, very obtuse.
This is the commonest and most widely distributed species in the Philippines. The broad
head, impunctate temporal lobes, the pollinose ridge connecting the antennal rim with the
temporal lobe, and, in most specimens, two temporal setae, will separate it from O.
philippensis , the other widespread species in the islands. The most similar is O. quadruplex.
The latter has more temporal setae, extragular setae, several setae on the apex of Stria I, a
broader pronotum, and a reduced orbital groove.
Range. - Philippines (Luzon, northern Palawan, Sibuyan Island and one specimen from
northern Mindanao). This latter specimen is different from all others in that it has four to six
punctures on the outer carina. In all other respects it seems to be O. duplex , though it is
possibly not conspecific. In addition to the type locality of Mt. Makiling, we have seen
Specimens from the following localities: Luzon: one female, Alabang, Rizal Prov., XI I- 1945, B. Malkin Colin.
(NMNH); two females, S.E. Bataan, July-Aug. 1945, coll. Darlington (MCZ); one male, one female, Mt. Bulasan, Okt.,
1917, leg Bottcher (MNHB); one male, two females, Dingalan Bay, June-Aug. 1945, coll. Darlington (MCZ); five
females, Imugin, N. Viscaya, Baker (NMNH); ten females, Imugan, May, 1916, May & June 1917, leg. Bottcher
(MNHB); two males, two females, Los banos, March, July, 1914, leg. Bottcher (MNHB); one female, Los Banos, 8-9,
2-914, Bottcher (MNNH); one male, Los Banos, Colin. Baker (NMNH); one male, Los Banos, Colin. Baker (MCZ); one
female, Los Banos, May-June, 1947, 2nd growth, 100-1500 ft. F. G. Werner leg. (CNHM); two males, one female,
Malinao, Tayabas, Colin. Baker, 5989 (NMNH); one male, Manila (MNHN); one female, Massisiat, Abra Prov.,
Revision of Omoglymmius Ganglbauer
221
V-1946, 3500 ft., CNHM Zool. Exp. 1946-47 (CNHM); one female, Montalvan, E. Simon (MNHN); one female,
Ripang, 11-1918, Bottcher, Colin, Jul. Moser (MNHB); one male, three females, Zambales Prov., X 1 1- 1 9 1 7, Bottcher
(MNHB); MINDANO: one female, Surigao, 4-2-1915, leg. Bottcher (MNHB); PALAWAN: one male, two females,
Binaluan, 13-1-1914, leg. Bottcher (MNHB); SIBUYAN I.: one male, Colin. Baker (NMNH).
Variation. - The specimens from Mindanao and Sibuyan are possibly not conspecific with
those of Luzon. The Mindanao specimen has four to six punctures on the outer carina; that
from Sibuyan differs from all others seen by us in having a ventral tooth on the hind femur of
the male (in addition to the usual one on the front femur).
Omoglymmius (sensu stricto) bouchardi new species
Fig. 150
Type material. - HOLOTYPE female, labelled: “SUMATRA Palembang, ty-Grouv”
(MNHN). PARATYPES two females, same locality data (but not cited as “ty”) (MNHN);
one female, same locality data as holotype, labelled also “ty-Grouv” (LEI); one female, same
locality data as holotype, labelled “Grouvelle, Fry Colin. 1905-100” (BMNH); one male;
labelled “Sumatra” (MNHN). The “ty” labels probably were intended to designate type
specimens. Grouvelle did not describe this species.
Description. — Length 6. 0-6. 8 mm. Antennal Segments 1-IV sparsely, coarsely punctate; Segments V-XI
impunctate; head 1.25 longer than broad, median lobe oval, its tip broadly rounded, its sides slightly parallel; frontal space
rather broad, U-shaped, its lateral margins markedly curved; median angles obtuse, narrowly separated; posteriomedial
margin curved; posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe separated from temporal lobe
by moderately wide antennal groove; temporal lobe with 15-20 rather fine punctures in posteriolateral portion; orbital
groove broad, extending to posterior margin of eye; one temporal seta; postorbital, suborbital tubercles absent; eye large,
round.
Pronotum moderately broad, length/greatest width 1.21; widest near middle, base moderately narrowed; apex more
strongly narrowed; lateral margin sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base;
inner carinae slightly broader than outer carinae at middle; inner carina constricted near base; latter broadened, truncate;
outer carina impunctate or with one or two punctures; marginal grooves slightly dilated; pronotum without setae;
prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp of elytron glabrous except for
longitudinal scarp at base of Stria IV and pollinose spot at base of Stria 11; Stria II with one seta near apex; Stria IV with
one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate;
abdominal Sterna II1-V with pollinose transverse sulci separated at midline; sternal punctures incorporated by sulci;
lateral pit of Sternum IV in female shallow; front femur of male with prominent, acute ventral tooth; middle calcar acute,
about 0.5 as long as spur; hind calcar triangular, its apex rounded, slightly lobate, its dorsal margin obtusely angulate.
This species and O. consors are Sumatran species with pronotal carinae impunctate or
nearly so, median lobe of head narrow and truncate, and abdominal sterna with the punctures
confluent, in form of transverse sulci. O. consors differs in having the basal margin of the
elytron entirely pollinose, and antennal lobes more markedly elevated above the temporal lobes.
The remaining species from Sumatra have the outer carinae punctate.
Grouvelle had intended to name this species in honor of M. Bouchard, who brought him the
specimens, and whose observations on Rhysodidae in the dead limbs of freshly felled trees in
the Sumatran forest were cited by Grouvelle (1903). We are pleased to carry out his intentions.
Range. - Sumatra. In addition to the type material we have seen two males, labelled “Z.O.
Serdang, Sumatra’s O.K. ± 1000', Dec. 89, 1.Z. Kannegieter” (MNHN).
Omoglymmius (sensu stricto ) consors new species
Fig. 151
Type material. - HOLOTYPE male, labelled: “SUMATRA, Palembang” (MNHN).
PARATYPES three males, two females, same data as holotype (MNHN).
Quaest. Ent., 1982, 18 (1-4)
222
Bell and Bell
Description. — Length 5. 3-6. 5 mm. Antennal Segments I-Ill faintly punctate; Segments 1V-X1 impunctate; head
1 .5 longer than broad; median lobe parallel-sided, its tip broadly rounded; frontal space broad, U-shaped, its lateral
margins markedly curved; medial angles obtuse to rounded, narrowly separated; posteriomedial margin curved;
posteriolateral margin evenly curved; occipital angle indistinct; antennal lobe raised above level of temporal lobe,
connected to latter by pollinose ridge; temporal lobe with three to six fine punctures; orbital groove complete though
narrow; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum rather elongate, narrow; length/greatest width 1.36; widest near middle, base slightly narrowed, apex
strongly narrowed; lateral margins nearly parallel; margin not sinuate anterior to hind angle; medial margin of outer
carina markedly sinuate anterior to base; inner carina slightly broader than outer one at middle; inner carina constricted
near base, latter truncate but only slightly broadened; both carinae impunctate; marginal grooves not dilated; pronotum
without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; basal margin of elytron entirely pollinose, base of
Stria IV concealed, thus longitudinal scarp not evident; Stria II with one or two setae near apex; Stria IV with one or two
setae near apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum coarsely punctate;
punctures of abdominal Sterna III-V in transverse sulci contiguous across midline or only narrowly interrupted there;
lateral pit of Sternum IV in female round, deeper than in O. bouchardi\ front femur of male with prominent acute ventral
tooth; middle calcar acute, about 0.5 as long as spur; hind calcar triangular, its apex obtuse, its dorsal margin not angulate.
This species differs from all others in presence of a complete band of pollinosity on the basal
transverse scarp of the elytron. Otherwise, it is rather similar to O. bouchardi, from which it
also differs in having the pronotum more elongate and parallel-sided, and transverse sulci of the
abdomen at most narrowly interrupted. In Borneo, completely impunctate outer carinae and
the transverse sulci of the abdominal sterna separate it from O. fraudulentus and O. nemoralis.
The name was chosen by Grouvelle, who died without publishing a description.
Range. - Sumatra, Borneo. In addition to the type material we have seen one male, labelled:
“Borneo Occ. Setipas pres Singawang, J. B. Ledru 1897” (MNHN).
Omoglymmius { sensu stricto) repetitus new species
Figs. 155, 156
Type material. - HOLOTYPE female, labelled: “Nord-Celebes, Toli-toli, Nov.-Dez. 1895,
H. Fruhstorfer” (MNHN).
Description. — Length 6.9 mm. Antennal Segments I-IV coarsely punctate; outer segments with punctures
successively finer; those of X scarcely visible; Segment XI impunctate; head slightly longer than wide; median lobe broad,
lance-shaped, its apex very obtuse; frontal space broad, U-shaped, its lateral margins markedly curved; medial angles
rounded, well separated; posteriomedial margin curved into posteriolateral margin; latter curved most markedly opposite
temporal seta; occipital angle absent; antennal lobes raised above level of temporal lobes, these and slope connected to
temporal lobe pollinose; posterior margin of temporal lobe narrowly fringed by pollinosity; orbital groove absent; temporal
lobe with about 12 very fine punctures near lateral margin; one temporal seta; small postorbital tubercle present; eye large,
round.
Pronotum rather short, length/greatest width 1.15; widest near middle; base slightly narrowed; apex more markedly
narrowed; lateral margins slightly curved, markedly so near apex; margin shallowly sinuate anterior to hind angle; inner
carina slightly broader than outer one at middle; medial margin of outer carina shallowly sinuate anterior to base; outer
carina widest anterior to middle; narrowed to both base and apex; inner carina narrowed posteriorly, its base obliquely
truncate; outer carina with seven or eight fine punctures; inner carina with about three fine punctures; pronotum without
setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with very small, pollinose,
longitudinal scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with several setae in apex;
metasternum coarsely punctate; abdominal Sternum III with transverse row of punctures, IV, V with transverse bands of
coarse punctures, confluent laterally on IV, V, VI; female with deep round lateral pit on Sternum IV; female with small,
very obtuse ventral tubercle on anterior femur; male unknown.
Pollinosity of antennal lobes is characteristic of this species. Presence of a small postorbital
tubercle separates it from all species of Borneo and the Philippines, and from all other species
from Wallacea except O. quadraticollis. The latter species has glabrous antennal lobes and a
differently-shaped pronotum. Several species from New Guinea are superficially similar in
having very small postorbital tubercles, but differ in having the antennal lobes glabrous, and
Revision of Omoglymmius Ganglbauer
223
striae not impressed.
Omoglymmius (sensu stricto) quadraticollis (Arrow)
Figs. 152, 157
Rhysodes quadraticollis Arrow 1901: 87.
Omoglymmius ( sensu stricto ) quadraticollis (Arrow) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) male, labelled: “Tenimbar I., Doherty,
1900-150” (BMNH). PARALECTOTYPES two females, same data as lectotype (BMNH).
The locality, usually spelled “Tanimbar”, is a group of islands between Timor and New
Guinea.
Description. — Length 5. 2-7.0 mm. Antennal Segments I-X punctate; Segment XI impunctate; head slightly longer
than wide; median lobe broad, nearly circular, its apex broadly rounded; frontal space broad, U-shaped, its margins
markedly curved; medial angles broadly rounded, nearly contiguous, posteriomedial margin curved into posteriolateral
margin; latter more evenly curved than in O. repetitus\ occipital angle absent; antenal lobes glabrous, raised above level of
temporal lobes, slope between antennal and temporal lobes broadly pollinose; posterior margin of temporal lobe narrowly
fringed by pollinosity; orbital groove represented by broad pollinose band tapered to point near posterior margin of eye;
temporal lobe with four to seven fine punctures mostly anterior to temporal setae; one temporal seta; small postorbital
tubercle present, visible only in profile view; eye large, round.
Pronotum rather short, length/greatest width 1.15; widest distinctly behind middle; base not narrowed; apex slightly
narrowed; pronotum nearly rectangular, lateral margins nearly straight, slightly convergent anteriorly; margin not sinuate
anterior to hind angle; carinae subequal in width at middle; medial margin of outer carina scarcely sinuate anterior to
base; outer carina of nearly even width except at rounded apex; inner carina narrowed posteriorly, its base narrowly
truncate; outer carina with 10-17 fine punctures; inner carina with about three fine punctures; pronotum without setae;
prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp of elytron pollinose;
longitudinal basal scarp of Stria IV at most merely suggested; Stria IV with one seta near apex; subapical stride with one
seta; Stria VIII with several setae near apex; metasternum finely, nearly uniformly punctate; abdominal Sterna II1-V with
coarse punctures in form of irregular transverse row on each sternum; both sexes with lateral pit in Sternum IV, deeper in
female; male with obtuse ventral tooth on anterior femur; middle calcar acute, very small, less than 0.25 of length of spur;
hind calcar nearly triangular, its apex acute, only slightly above level of spur, its dorsal margin indistinctly angulate.
This species is characterized by its almost rectangular pronotum, which is widest near the
base. A small postorbital tubercle separates it from all other species of Wallacea except for O.
repetitus of Celebes. The latter does not have a rectangular pronotum, and also has the median
lobe pointed posteriorly.
Omoglymmius (sens u stricto) batchianus (Arrow)
Figs. 153, 158
Rhysodes batchianus Arrow 1901: 86.
Omoglymmius ( sensu stricto) batchianus (Arrow) Bell and Bell 1978.
Type material. - HOLOTYPE female, labelled: “Batchian, Pascoe Coll. 93-60” (BMNH).
Description. — Length 7. 0-7.2 mm. Antennal Segments I-X punctate. Segment I coarsely so, punctures finer on
more distal segments, those of Segment X extremely fine; Segment XI impunctate; head slightly longer than broad;
median lobe broad, its tip obtusely pointed; frontal space broad, U-shaped, its margins markedly curved; medial angles
rounded, well separated; posteriomedial margin curved into posteriolateral margin; latter evenly curved; occipital angle
absent; antennal lobe raised above level of temporal lobe, slope between them broadly pollinose; orbital groove nearly
absent, represented by narrow strip of pollinosity medial to anterior margin of eye; temporal lobe with three to five fine
punctures between vestige of orbital groove and temporal seta; one temporal seta; postorbital, suborbital tubercles absent;
eye large, round.
Pronotum rather short, broad; length/greatest width 1.10; widest slightly behind middle; base very slightly narrowed;
apex distinctly narrowed; lateral margins nearly parallel in basal half, more curved, convergent in apical half; margin not
sinuate anterior to hind angle; inner carinae slightly wider than outer ones at middle; medial margin of outer carina not
sinuate anterior to base; outer carina of nearly even width except at slightly narrowed extreme apex; inner carina widest at
Quaest. Ent., 1982, 18 (1-4)
224
Bell and Bell
basal third, narrowed anteriorly, posteriorly, base obtuse, no trace of constriction anterior to base; outer carina with
20-25 very fine punctures; inner carina with four to six fine punctures; pronotum without setae; prosternum without
precoxal carinae.
Elytron rather short, striae shallowly impressed, finely punctate; base of Stria IV with pollinose longitudinal scarp;
Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum
coarsely punctate; abdominal Sterna III-V each with transverse band of coarse punctures; Sterna 111-V1 with punctures
confluent laterally in form of irregular transverse sulci; Sternum IV in both sexes in form of deep, round lateral pit;
male with obtuse ventral tooth on anterior femur; female with vestigial tooth or angle on anterior femur; middle calcar
obtuse, very small, less than 0.25 as long as spur; hind calcar nearly triangular, its apex obtuse, only slightly above level
of spur, its dorsal margin sinuate.
This is a species with a broad, almost quadrate pronotum and shallow striae with fine
punctures. It is close to O. humeralis of Ternate, but the latter species has the lateral margins
of the pronotum more strongly curved. Shallow elytral striae, fine punctures, and tapered bases
of inner carinae are points of resemblance to several species from New Guinea. Of these, the
closest are O. fringillus and O. oroensis, both of which lack postorbital and suborbital
tubercles. In these species, the glabrous part of the temporal lobe is much more broadly
separated from the antennal lobe.
Range. - Probably restricted to Batjan (also spelled “Batchian” and “Bacan”, a group of
islands in the North Moluccas. Neither Arrow nor Grouvelle adequately compared this species
with O. humeralis (Grouvelle) of Ternate and Halmahera. Differences in shape between
pronota of the two appear to be equal to specific differences in other groups of Omoglymmius s.
str ., but confirmation of their distinctness will have to await the collection of more material. In
addition to the holotype, we have studied one male, labelled “Batjan, Bernstein” (LEI).
Omoglymmius (sensu stricto) humeralis (Grouvelle)
Fig. 154
Rhysodes humeralis Grouvelle 1895a: 157.
Omoglymmius ( sensu stricto) humeralis (Grouvelle) Bell and Bell 1978.
Type material. - LECTOTYPE (here designated) female, labelled: “MOLUQUES,
Ternate, Raffray et Maindron 78” (MNHN). PARALECTOTYPE female, labelled:
“MOLUQUES, Gilolo, Raffray & Maindron 78” (MNHN). The original description gave the
locality as “Celebes”, but Grouvelle later (1903), listed it as “Ternate”. For this reason, we
chose the Ternate specimen as lectotype. “Celebes” perhaps referred to a political unit, and not
the island. “Gilolo” is an old name for Halmahera, the largest island in the North Moluccas.
The specimens from the two islands are not identical, and are quite possibly specifically
distinct, although this is difficult to judge on the basis of one specimen from each island. The
description below is based entirely on the lectotype. Points of difference of the paralectotype are
listed below, under the heading of “variation.”
Description. — Length 6.5-7. 1 mm. Antennal Segments I- IX punctate; Segment I coarsely so; punctures finer
distally, those of Segment IX very fine; Segments X, XI impunctate; head slightly longer than broad; median lobe broad,
rather long, its tip obtusely pointed; frontal space very broad, more than twice as broad as long; U-shaped, its sides
markedly curved; median angles obtuse, well separated; posteriomedial margin curved into posteriolateral margin; latter
evenly curved; occipital angle absent; antennal lobe raised above level of temporal lobe, slope between them broadly
pollinose; orbital groove nearly absent, represented by broad but very short strip of pollinosity opposite anterior margin of
eye; temporal lobe with 12-14 very fine punctures in lateral half; one temporal seta; postorbital, suborbital tubercles
absent; eye large, round.
Pronotum moderately long, length/greatest width 1.37; widest near middle, base distinctly narrowed; apex more
strongly so; lateral margins nearly evenly curved; margin not sinuate anterior to hind angle; inner carinae slightly wider at
middle than outer carinae; medial margin of outer carina shallowly sinuate anterior to base; outer carina widest just
anterior to middle; inner carina widest near middle, narrowed, obtuse anteriorly, narrowed to point posteriorly; no
constriction near base of inner carina; outer carinae with 1 8-23 very fine punctures; inner carina with six or seven very fine
Revision of Omoglymmius Ganglbauer
225
punctures; pronotum without setae; prosternum without precoxal carinae.
Elytra rather short; stria shallowly impressed, finely punctate; base of Stria IV with pollinose longitudinal scarp;
Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum
coarsely punctate; abdominal Sterna III-V each with sparse coarse punctures, in form of one irregular transverse row on
each sternum; punctures of Sterna 1V-VI confluent laterally, in form of rudimentary transverse sulci (these less distinct
than in O. batchianus)-, female with deep, round lateral pit on Sternum IV; female with minute, obtuse ventral tooth on
anterior femur; male unknown.
The most similar species to this one are O. batchianus, O. viduus and O. opticus. The first,
from Batjan, has the pronotum with sides almost parallel; the second one, from the Kei Islands,
has the pronotal margins more strongly curved, and the temporal lobe with more and coarser
punctures, while the third, from Dammar, has the frontal space much narrower.
Range. - In addition to the lectotype, we have studied the following specimens: two females,
labelled: “Ins. Ternate, Doherty VII" (MCZ); two females, labelled “Ternate, Kannegieter (Museum Natura Artis
Magistra)” (AMS).
Variation. - As stated above, the paralectotype, from Halmahera, is rather different from
the lectotype, and might represent another, unnamed species, possibly restricted to Halmahera.
We do not wish to name it until more specimens from Halmahera are available. The length is
6.5mm. The principal points of difference from the specimens from Ternate are as follows:
median lobe shorter, more broadly rounded posteriorly; frontal space smaller; medial angles more obtuse, more narrowly
separated; orbital groove represented by a much narrower strip of pollinosity; temporal lobe with only about six minute
punctures, these forming irregular row anterior to temporal seta; pronotum shorter, length/greatest width 1.18; inner
carinae subtruncate, much less narrowed at base; fewer pronotal punctures (about 10 on outer carina; one to three on inner
carina); basal scarp of Stria IV exceptionally long, its length almost twice distance from scarp to elytral humerus (in
Ternate specimens it is shorter than distance to humerus).
Omoglymmius (sensu stricto) opticus new species
Fig. 159
Type material. - HOLOTYPE female, labelled: “Dammer Jnsel” (MNHB). PARATYPE
female, same data as holotype. The label probably refers to the Damar Islands, which lie in the
Banda Sea, northeast of Timor and northwest of Tanimbar.
Description. — Length 6. 2-6. 7 mm. Antennal Segments I-X punctate, those of distal segments very fine; Segment
XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex very obtuse, nearly rounded;
frontal space rather narrow, U-shaped, its lateral margins distinctly curved; medial angles very obtuse, slightly separated;
posteriomedial margin evenly curved into posteriolateral margin; occipital angle absent; antennal lobe glabrous, separated
from temporal lobe by broad band of pollinosity continued as very short orbital groove; latter ended opposite anterior 0.25
of eye; temporal lobe with 10-12 fine, scattered punctures; one temporal seta; postorbital, suborbital tubercles absent; eye
large, round, more convex than in other Omoglymmius s. str., clearly visible in dorsal view.
Pronotum rather short; length/greatest width 1.18; widest near middle; base, apex scarcely narrowed; pronotum nearly
rectangular; lateral margins very feebly curved; margin not sinuate anterior to hind angle; inner carina slightly broader
than outer carina at middle; medial margin of outer carina very shallowly sinuate anterior to base; outer carina widest
anterior to middle; apex subtruncate; inner carina narrowed at base, slightly constricted anterior to base; outer carina with
two to four fine punctures; inner carina with one puncture, or none; pronotum without setae; prosternum without precoxal
carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; basal scarp pollinose; base of Stria IV with small
pollinose longitudinal scarp; Stria IV with one seta at apex; subapical striole with one seta; Stria VII with several setae
near apex; metasternum finely punctate; abdominal Sterna III-V with coarse, rather sparse punctures in form of broad
band on each sternum; female with deep, round lateral pit on Sternum IV; male unknown.
This species is closest in apperance to O. quadraticollis of the Tanimbar Islands. It
resembles the latter in having the pronotum nearly rectangular but differs in lacking the
postorbital tubercle, in having the pronotal carinae nearly impuctate: and in having the eyes
very convex. Shape of the pronotum and very convex eyes separate it from O. batchianus and
its relatives.
Quaest. Ent., 1982, 18 (1-4)
226
Bell and Bell
Omoglymmius ( sensu stricto) viduus new species
Fig. 160
Type material - HOLOTYPE male, labelled: “Key Ins.” (MNHB). PARATYPES one
male, one female, same data as holotype (MNHB); one female, labelled:“Key Inseln, coll.
Plason, Coll. Reithoffer” (BSL).
Description. — Length 6. 2-7. 2 mm. Antennal Segments I-X distinctly punctate; Segment XI impunctate; head
slightly longer than wide; median lobe broad, lance-shaped, its apex obtuse; frontal space broad, U-shaped; its lateral
margins markedly curved; medial angles acute, nearly contiguous; posteriomedial margin slightly sinuate; posteriolateral
margin nearly evenly curved; occipital angle very obtuse; postantennal groove narrow; orbital groove abbreviated, scarcely
extended to anterior margin of eye; temporal lobe with 20-25 rather fine punctures, scattered over its entire surface; one
temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Prothorax rather long, length/greatest width 1.32; widest near middle; base slightly narrowed; apex more markedly
narrowed; lateral margins curved; margin not or but slightly sinuate anterior to hind angle; inner carina slightly broader
than outer one at middle; medial margin of outer carina not distinctly sinuate anterior to base; outer carina widest near
middle, markedly narrowed to apex; inner carina with basal part narrowed parallel-sided; base truncate; outer carina with
21-27 fine punctures; inner carina with five to seven fine punctures; pronotum without setae; prosternum without precoxal
carinae.
Elytra relatively long, narrow; striae impressed, coarsely punctate; base of Stria 1 with longitudinal pollinose scarp;
Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with
coarse but very shallow punctures; abdominal Sterna 1I1-V with coarse, irregularly distributed punctures; these coalescent
laterally to form indistinct transverse sulci; both sexes with deep, round lateral pit on Sternum IV; male with ventral tooth
on anterior femur; middle calcar very small, indistinct; hind calcar triangular, its apex obtusely pointed; its dorsal margin
slightly sinuate.
This species differs from O. quadraticollis of the Tanimbar Islands in having the pronotum
much more elongate and less quadrate. Numerous punctures of the temporal lobes and outer
carina separate it from O. opticus of the Kei Islands. Deeper elytral striae and numerous
punctures of the outer carinae separate it from O. batchianus and similar forms.
Omoglymmius ( sensu stricto ) continuus new species
Figs. 161, 166, 169, 170
Type material. - HOLOTYPE male, labelled: “Mangole (Sula Inseln), VII-XII, 1977,
Wegener, DOY-K” (BSL). This is one of the islands east of Celebes, and lies southwest of
Batjan and north of Buru.
Description. — Length 6.9 mm. First segment of antenna with marked swelling around base of principal tactile
setae (Fig. 169); Segments I-X punctate; punctures of distal segments very fine; Segment XI impunctate; head slightly
longer than wide; median lobe oval, rather broad; its apex broadly rounded; frontal space broad, U-shaped; medial angles
broadly rounded, nearly contiguous; posteriomedial margin evenly curved; posteriomedial margin broadly curved, margin
more markedly curved at eye; occipital angle absent; antennal lobes pollinose, not sharply elevated above anterior end of
temporal lobe; anterior end of latter extensively pollinose, boundary of pollinosity extending obliquely posteriomedially;
pollinosity of orbital groove extending 0.75 of length of eye; temporal lobe with four to six very fine punctures; two or three
very coarse punctures, probably each bear temporal seta (but only one present on either side in holotype, the other perhaps
being broken off); postorbital, suborbital tubercles absent; eye large, round.
Pronotum subquadrate; rather short, length/greatest width 1.17; widest near base; sides nearly parallel; apex slightly
narrowed; lateral margin not at all sinuate anterior to hind angle; inner carina slightly wider than outer one; medial margin
of outer carina not sinuate anterior to base; outer carina of nearly even width; inner carina constricted anterior to base;
latter broad, truncate, outer carina with 11-12 very fine punctures; inner carina with nine to 1 1 very fine punctures; lateral
margin of inner carina sloped gradually into paramedian groove; pronotum without seta; prosternum without precoxal
carinae.
Elytron moderately narrow; inner ( I- 1 1 1 ) striae finely punctate, very shallowly impressed; outer striae deeper, more
coarsely punctate; base of Stria IV with very small longitudinal scarp; elytral setae absent except for several near apex of
Stria VII; metasternum densly punctate; abdominal Sterna III-V densely punctate; male with deep, round lateral pit on
Sternum IV; male with prominent ventral tooth on anterior femur; middle calcar acute, very small; hind calcar
subtriangular, its apex truncate, raised above level of spur, its ventral margin emarginate; its dorsal margin angulate (Fig.
1 70); female unknown.
Revision of Omoglymmius Ganglbauer
227
This species differs from other members of the subgenus in having the lateral margin of the
inner carina sloped gradually into the paramedian groove, as in most species of Rhyzodiastes.
The indistinct boundary between the antennal and temporal lobes is also unique. It differs
sharply from O. wittmeri, the other species known from Mangole in these characters, and also
in the subquadrate pronotum, in having the base of Antennal Segment I markedly swollen, in
having temporal lobes scarcely punctate, and, apparently, in having more than one temporal
seta.
Omoglymmius ( sensu stricto ) wittmeri new species
Figs. 162, 167, 168, 171
Type material. - HOLOTYPE male, labelled: “Mangole (Sula Inseln) VII-XII,1977,
Wegener, DRP-K” (BSL). PARATYPES two females, same data as holotype but collected by
V. & G. Wegener (BSL).
Description. — Length 5. 3-6. 7 mm. First segment of antenna scarcely swollen at base of principal tactile seta (Fig.
168); Antennal Segments I-X punctate; punctures of distal segments very fine; Segments XI impunctate; head slightly
longer than wide: median lobe lance-shaped, its apex very obtuse; frontal space broad, U-shaped; medial angles obtuse,
narrowly separated; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct;
antennal lobe glabrous, raised above level of temporal lobe; anterior end of temporal lobe extensively pollinose, boundary of
pollinosity extended obliquely posteriomedially; pollinosity of orbital groove extended posterior to eye, merged with narrow
fringe of pollinosity along posterior margin of temporal lobe; temporal lobe with 12-18 rather fine punctures; one temporal
seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum rather short; length/greatest width 1.14; widest slightly anterior to middle; base moderately narrowed; apex
more markedly so; lateral margins curved; lateral margin scarcely sinuate anterior to hind angle; carinae subequal at
middle of length; medial margin of outer carina shallowly sinuate anterior to base; outer carina widest at anterior third;
narrowed both at base and apex; inner carina constricted just anterior to base; latter truncate, scarcely broadened; outer
carina with 18-25 punctures; inner carina with four or five fine punctures; medial margin of outer carina separated from
paramedian groove by sharp, distinct scarp; pronotum without setae; prosternum without precoxal carinae.
Elytron narrow, elongate; elytral striae impressed, coarsely punctate; base of Stria IV with longitudinal scarp; Stria IV
with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with dense,
coarse but shallow punctures; abdominal Sterna III-V densely punctate; punctures near lateral margin of III, V, VI
coalescent; both sexes with deep round lateral pit on Sternum IV; male with prominent ventral tooth on anterior femur;
female with vestige of anterior femoral tooth; middle calcar small, triangular, about 0.3 as long as spur; hind calcar
triangular, its apex slightly acute, at level of spur, its distal margin straight (Fig. 171).
The glabrous, sharply defined antennal lobe, rounded pronotal margins, sharp outer
boundary of the inner carina, lack of a swelling on Antennal Segment I, and different shape of
the hind calcar differentiate this species from the sympatric O. wittmeri. The temporal lobe has
a continous pollinose margin extended from the postantennal groove to the posterior margin of
the temporal lobe. This feature is not found in other Moluccan species, but is duplicated in a
few species from other regions, such as O. fringillus of New Guinea and O. gurneyi of the
Solomon Islands. The former differs in the shape of the head, while the latter has a nearly
quadrate pronotum.
We dedicate this species to Dr. Wittmer of the Basel Natural History Museum in gratitude
for his help in this study.
Omoglymmius ( sensu stricto) vadosus new species
Fig. 163
Type material. - HOLOTYPE female, labelled: “Amboina,X-XI-07, coll. E. Muir”
(BPBM). Amboina, now Ambon, is in the South Molluccas, south of Ceram.
Quaest. Ent., 1982, 18 (1-4)
228
Bell and Bell
Description. — Length 7.0 mm. Antennal Segments I-X punctate; punctures of distal segments very fine; Segment
XI impunctate; head slightly longer than wide; median lobe broad, lance-shaped, its apex very obtuse; frontal space broad,
transverse, U-shaped; margin abruptly curved, medial angles obtuse, very narrowly separated; posteriomedial margin
oblique, posteriolateral margin nearly evenly curved; occipital angle distinct; antennal lobe raised above level of temporal
lobe, separated from latter by rather broad pollinose strip; latter continuous with short but broad orbital groove, ended
opposite middle of eye; temporal lobe with six to eight very minute punctures; one temporal seta (holotype has two coarse
punctures on right temporal lobe, and perhaps a second seta was broken off); postorbital, suborbital tubercles absent; eye
large, round.
Pronotum moderately long, length/greatest width 1.26; widest near middle; base, apex narrowed; lateral margins
markedly curved; margin scarcely sinuate anterior to hind angle; inner carina slightly broader than outer carina at middle;
medial margin of outer carina not sinuate near base; outer carina widest anterior to middle; inner carina tapered to base;
outer carina with 12-14 extremely minute punctures, visible only under high magnification; inner carina with two or three
minute punctures; pronotum without setae; prosternum without precoxal carinae.
Elytron long, rather narrow; striae shallowly impressed; punctures of inner striae rather fine, those of outer striae
coarser; base of Stria IV with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical stride without seta;
Stria VII with several setae near apex; metasternum sparsely, shallowly punctate; abdominal sterna with shallow,
pollinose, lateral transverse sulci, with one or two isolated punctures between the sulci on each sternum; female with deep
lateral pit on Sternum IV; male unknown.
O. vadosus is characterized by the extremely fine punctures on the head, pronotum, elytra
and metasternum. The propleuron is minutely pollinose as are the transverse sulci of the
abdomen.
Omoglymmius (sensu stricto) morditus new species
Fig. 165
Type material. - HOLOTYPE male, labelled: “Morotai Is., Dutch E. I., Sept. 1944,
Darlington” (MCZ). This island is in the North Moluccas, northeast of Halmahera.
Description. — Length 5.8 mm. Antennal Segments I, II coarsely punctate; III, IV impunctate, V-IX very minutely
punctate (appearing impunctate except under high magnification); Segments X, XI impunctate; head slightly longer than
broad; median lobe short, broad, its tip broadly rounded; frontal space very broad, U-shaped; its margins bent abruptly;
medial angles obtuse, well separated; posteriomedial margin oblique; occipital angle indistinct; posterior margin of head
nearly transverse, in form of obtuse angle with nearly straight lateral margin of temporal lobe; antennal lobe raised above
level of temporal lobe, slope between them broadly pollinose; orbital groove absent; posterior margin of temporal lobe
narrowly fringed by pollinosity; temporal lobe with about 12 fine punctures in posteriolateral half; one temporal seta;
postorbital, suborbital tubercles absent; eye large, round.
Pronotum rather short, length/greatest width 1.17; subquadrate; widest near middle, slightly narrowed to base and
apex; lateral margins very slightly curved; margins not sinuate anterior to hind angle; inner carina subequal to outer carina
at middle; medial margin of outer carina not sinuate anterior to base; outer carina of nearly even width except at extreme
base, apex; inner carina widest anterior to middle; tapered to point at base; outer carina with 13-15 punctures; inner carina
with three or four very fine punctures; pronotum without setae; prosternum without precoxal carinae.
Elytron rather long, narrow; striae deeply impressed, coarsely punctate; base of Stria IV with very small longitudinal
scarp; transverse basal scarp of elytron pollinose; Stria IV with one seta near apex; subapical stride with one seta; Stria
VII with several setae near apex; metasternum coarsely, shallowly punctate; abdominal Sterna III-V very coarsely,
sparsely punctate; punctures of each sternum tending to form transverse row; male with ventral tooth on anterior femur;
middle calcar very small, obtuse; hind calcar triangular, apex obtuse; upper margin slightly sinuate; female unknown.
Among Moluccan species, this one is most similar to O. batchianus in the subquadrate form
of the pronotum, but O. morditus differs from all other Moluccan species in the subquadrate
form of the head, with a distinct angle between the lateral and posterior margins of the
temporal lobe.
Omoglymmius ( sensu stricto ) nasalis new species
Fig. 164
Type material. - HOLOTYPE female, labelled: “Kajeli, Boeroe” (MNHN). The island,
now spelled “Buru”. is in the South Moluccas, south of Mangole, and west of Ceram and
Revision of Omoglymmius Ganglbauer
229
Amboina.
Description. — Length 7.7 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head nearly
twice as long as broad; preocular portion produced; median lobe elongate, its margins nearly parallel; its apex slightly
narrowed, rounded; medial angles obtuse, slightly separated; frontal space very broad, U-shaped, its margins markedly
curved; medial angles rounded; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle
indistinct; antennal lobe far from eye; pollinose oblique ridge between antennal lobe, frontal space; deep oblique groove
posteriolateral to ridge; surface of temporal lobe between oblique groove and eye pollinose, representing short but broad
orbital groove; temporal lobe with four to six punctures near temporal seta; one temporal seta; postorbital, suborbital
tubercles absent; eye large, round.
Pronotum rather short, length/greatest width 1.18; widest near middle; base slightly narrowed; apex more markedly
narrowed; lateral margins curved; margin distinctly sinuate anterior to hind angle; inner carinae slightly broader than
outer carinae at middle; medial margin of outer carina angulate but not sinuate near base; outer carina of nearly even
width except at extreme base, apex; inner carina widest near middle; broadly truncate anteriorly, constricted just anterior
to broadened, truncate base; outer carina with 25-28 coarse punctures; inner carina with 12-13 coarse punctures; pronotum
without setae; prosternum without precoxal carinae.
Elytron rather elongate, narrow; striae impressed; coarsely punctate, becoming finely so near apex; base of Stria IV
without longitudinal scarp; Stria IV without setae; subapical striole with one seta; Stria VII with several setae near apex;
metasternum densely, rather finely punctate; abdominal Sterna III-VI finely, densely punctate; female with large, deep,
round lateral pit in Sternum IV; male unknown.
The elongate snout separates this species from all others from Wallacea except for O.
bucculatus. The latter has pronotal setae, several temporal setae, and has the outer carinae
more narrowed, and lacks the diagonal groove anterior to the eye.
Omoglymmius (sensu stricto) bucculatus (Arrow)
Fig. 172
Rhysodes bucculatus Arrow 1901: 88-89.
Omoglymmius ( sensu stricto ) bucculatus (Arrow) Bell and Bell 1978.
Type material. - LECTOTYPE (Here designated) female, labelled: “SUMBAWA Id.
(Doherty) 1900-150” (BMNH). PARALECTOTYPES three females, one male, same data as
type (BMNH).
Description. — Length 5. 9-7. 4 mm. Antennal Segments I-X punctate; Segment I coarsely punctate; punctures
becoming finer on distal segments, nearly obsolete on Segment X; Segment XI impunctata; head nearly twice as long as
wide, preocular portion produced; median lobe elongate, gradually tapered posteriorly; apex obtuse; frontal space narrow,
V-shaped, its margins oblique or slightly curved; medial angles obtuse, slightly to distinctly separated; posteriomedial
margin slightly emarginate; posteriolateral margin evenly curved; occipital angle distinct; antennal lobe separated from
anterior end of temporal lobe by narrow postantennal groove; small pit present anterior to eye; orbital groove absent;
temporal lobe with nine to 15 moderately coarse punctures; one to three temporal seta; postorbital, suborbital tubercles
absent; eye large, round.
Pronotum various in length, length/greatest width 1.23 in short specimens, up to 1.33 in longer ones; widest near
middle; base, apex markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina 0.35
to 0.5 as wide as inner one at middle; lateral margins markedly curved; margin not sinuate anterior to hind angle; marginal
groove dilated; outer carina with medial margin sinuate near base; inner carina widest near middle, constricted just
anterior to base, latter broadened, truncate; outer carina with one to eight coarse punctures or none in addition to two large
setiferous punctures; inner carina with one to 1 3 coarse punctures or none; two or three prominent setae on each outer
carina; precoxal carinae absent.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
II with one to three setae near apex or none; Stria IV one seta near base and one to two near apex; subapical striole without
seta; Stria VII with several setae near apex; metasternum with coarse punctures near margins and in midline, otherwise
impunctate; punctures of abdominal Sterna III-V coarse, in form of transverse row on each sternum; both sexes with deep
lateral pit in Sternum IV, this larger in female; male with small acute ventral tooth on anterior femur; middle calcar acute,
as long as spur; hind calcar subtriangular, apex obtuse or truncate, proximal margin angulate.
Pronotal setae are unique in the subgenus. The species is also characterized by the elongate
head, and, in most specimens, by more than one temporal seta.
Range. - Restricted to the island of Sumbawa, in the Lesser Sundas. In addition to type
material, we have Studied the following specimens: one male, one female, labelled: “B. Aroe Hassa,
Quaest. Ent., 1982, 18 (1-4)
230
Bell and Bell
Sumbawa 2-5000’ Doherty IX, X” (year not indicated)(MCZ); two males, two females, labelled: “Sumbawa, W.
Doherty” (MNHN); two females, labelled: “Sumbawa”, collector not indicated (MNHN).
Variation. - We are provisionally treating this as one variable species, but it is possible that
two or more species are involved. Most of the specimens can be assigned to one or other of two
morphs. The first, and most common, is characterized by having the mentum densely punctate,
the pronotum more elongate, and the hind calcar narrowly truncate at apex. The lectotype
belongs to this form. The second form has the mentum impunctate except for a few punctures
near its base, the pronotum less elongate and the hind calcar with apex obtuse. One specimen
(BMNH), however, agrees with the first form in the shape of the pronotum, but agrees with the
second one in the absence of punctures on the mentum and in the shape of the hind calcar.
Omoglymmius ( sensu stricto) bicarinatus new species
Fig. 181
Type material. - HOLOTYPE male, labelled: “Ins. Jobi, N. Guinea (Doherty)” (MNHN).
This island, now called “Yapen”, formerly spelled “Japen”, is one of the Schouten Islands,
north of Geelfink Bay on the northwest side of New Guinea.
Description. — Length 4.8 mm. Antenna with Segments I- 1 1 1 coarsely punctate; Segments IV-X very minutely
punctate; Segment XI impunctate; head about 1.5 longer than wide; median lobe lance-shaped, its apex obtusely pointed;
frontal space rather narrow, its margins weakly curved; medial angles obtuse, widely separated; posteriomedial margin
emarginate; posteriolateral margin oblique, in form of obtuse angle with posterior margin, which is transverse; occipital
angle distinct; prominent preorbital pit; orbital groove absent; temporal lobe with about 16 fine punctures; one temporal
seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum relatively short, narrow; length/greatest width 1.14; shape irregular and probably somewhat deformed in
holotype; widest anterior to middle; base, apex narrowed; margins curved, margin not sinuate anterior to hind angle; outer
carina slightly narrower than inner carina at middle; medial margin shallowly sinuate anterior to base; outer carina widest
anterior to middle, narrowed to both base, apex; inner carina constricted just anterior to base, latter slightly broadened;
outer carina with 10-11 fine punctures; inner carina with six or seven fine punctures; pronotum without setae; prosternum
without precoxal carinae.
Elytron relatively short, narrow; striae impressed, coarsely punctate; Stria IV with longitudinal pollinose scarp at base;
Interval V forming narrow, elevated carina extended from humerus to subapical tubercle; Interval V bounded on both sides
by strip of pollinosity; Stria IV with one seta at base and one at apex; subapical stride with one seta; Stria VII with several
setae near apex; metasternum coarsely, densely punctate; abdominal Sterna 1II-V densely punctate; male with deep, round
lateral pit on Sternum IV; male without ventral tooth on anterior femur; middle calcar acute, longer than spur; hind calcar
triangular, small, shorter than spur, its apex acute.
The elevated cariniform fifth interval is characteristic of this species. Absence of a ventral
tooth on the anterior femur of the male is highly unusual; it is shared only with O. gracilicornis
of New Guinea, O. politus of Luzon, and two species from the Caroline Islands.
Omoglymmius ( sensu stricto ) caelatus Bell and Bell NEW COMBINATION
Fig. 173
Omoglymmius caelatus Bell and Bell 1981: 57-58.
Type material. - HOLOTYPE male, labelled: “PELELIU I., Palau Islands, West coast, 2
Feb. 1948, H. S. Dybas” (BPBM). PARATYPES one male, two females, labelled: “PELELIU
I. Palau Islands, East coast, 27 Jan. 1948, H. S. Dybas” (BPBM); one female, labelled: “Palau
Islands, Koror I., 17 Jan. 1948. H. S. Dybas” (BPBM). See Bell and Bell, 1981 for complete
data.
Description. — Length 5.8-7. 1 mm. Outer antennal segments faintly punctate; head slightly longer than wide;
frontal grooves very narrow; one temporal seta; posteriomedial margin emarginate; postorbital, suborbital tubercles absent.
Pronotum moderately long; length/greatest width about 1.28; outer carina with about 25-30 coarse punctures; inner
carina impunctate or with one or two minute punctures; inner carina constricted just anterior to base; latter broadened;
Revision of Omoglymmius Ganglbauer
231
marginal groove dilated.
Elytron with longitudinal scarp at base of Stria IV; striae shallow, scarcely impressed; punctures fine; male with
minute vestige of ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar with apex obtuse, dorsal
margin obtusely angulate.
For more complete description, see Bell and Bell (1981).
This species differs from O. impletus of Yap in having the outer carinae densely punctate
and from O. oceanicus of Kusiae, Ponape, and Palau in having the posteriomedial margin of the
temporal lobe emarginate, in having more punctures on the outer carina and in a greater
constriction anterior to the base of the inner carina. It is a rather nondescript species,
superficially similar to O. philippensis and its relatives in having a marked contrast between
the coarsely, densely punctate outer carina and an impunctate inner one. O. philippensis differs
in having the striae markedly impressed and coarsely punctate, and in having the ventral tooth
of the anterior femur prominent. O. patens of New Guinea is similar, but lacks the basal scarp
on Stria IV. Members of the O. lindrothi complex of the Solomon Islands are also similar, but
have the inner carina punctate.
Range. - Known only from the Palau Islands in the western Carolines.
Omoglymmius (sensu stricto) impletus Bell and Bell NEW COMBINATION
Fig. 174
Omoglymmius impletus Bell and Bell 1981: 58-60.
Type material. - HOLOTYPE male, labelled: “Yap group, Tomil Dist. Jul-Au 50, R. J.
Goss” (BPBM). PARATYPES two males, three females, same data as holotype (BPBM). See
Bell and Bell (1981) for complete data.
Description. — Length 5.2-6. 1 mm. Outer antennal segments impunctate; head slightly longer than wide; frontal
grooves very narrow; frontal space longer than broad, its margins slightly curved; posteriomedial margin emarginate,
narrowly fringed with pollinosity; one temporal seta; postorbital, suborbital tubercles absent.
Pronotum relatively short, length/greatest width 1.18; marginal groove dilated; outer, inner carinae entirely
impunctate; inner carina acutely pointed at base.
Elytron with longitudinal scarp at base of Stria IV; striae impressed; punctures fine, shallow; male without ventral
tooth on anterior femur; middle calcar very small, obtuse; hind calcar triangular, its apex obtusely pointed.
For a more complete description, see Bell and Bell (1981).
Among the species from the Caroline Islands, this species is easily recognized by the entirely
impunctate pronotal carinae and the acutely pointed base of the inner carina. This combination
of characters also occurs in several species from New Guinea, such as O. planiceps. Such
species lack the basal scarp on Stria IV, and many of them have either postorbital or suborbital
tubercles.
Range. - Known only from the Islands of the Yap Group, in the western Caroline Islands.
Omoglymmius (sensu stricto) oceanicus Bell and Bell NEW COMBINATION
Fig. 175
Omoglymmius oceanicus Bell and Bell 1981: 60-61.
Type material. - HOLOTYPE male, labelled: “Kusiae, Mutunlik, 22 m., 1-31-53, J.F.G.
Clarke” (BPBM). PARATYPES one male, two females, same locality and collector as
holotype (BPBM); one female, Kusaie, Malem, 1 9-XII- 1 937, Teiso Esaki (BPBM). See Bell
and Bell (1981) for complete data.
Description. — Length 5. 0-6. 8 mm. Outer antennal segments scarcely punctate; head slightly longer than wide;
frontal grooves rather narrow; frontal space slightly broader than long, V-shaped, its margins slightly curved;
Quaest. Ent., 1982, 18 (1-4)
232
Bell and Bell
posteriomedial margin curved nearly evenly into posteriolateral margin; occipital angle absent; one temporal seta;
postorbital, suborbital tubercles absent.
Pronotum moderately elongate; length/greatest width about 1.28; marginal groove narrow; outer carina with about
10-14 very fine punctures; inner carina impunctate; base of inner carina more broadly dilated than in O. caelatus.
Elytron with longitudinal scarp at base of Stria IV; striae impressed, coarsely punctate; male with minute ventral
tooth on anterior femur; middle calcar acute, very small; hind calcar with apex truncate, dorsal margin sinuate.
This species differs from O. caelatus in having the outer carina more finely and sparsely
punctate, the inner carinae more broadly truncate at the base, and the posterior margin of the
temporal lobe evenly curved.
Range. - Widely distributed in the Caroline Islands. Known from Kusiae, Ponape, and
Palau. For complete data, see Bell and Bell (1981).
The lindrothi complex
This group consists of the following nine species: Omoglymmius lindrothi, O. modicus, O.
rusticus, O. manni, O. regius, O. princeps, O. gurneyi, O. renutus, and O. scopulinus. These
very similar allopatric forms correspond to the pignoris complex of Kaveinga and could also be
regarded as subspecies of a single species. Since Rhysodini from continental landmasses are
rarely differentiated into subspecies, we prefer to regard the forms from the Solomon Islands as
distinct species. It is, nevertheless, convenient to describe the O. lindrothi complex as a whole
before describing the individual species.
Description. — Length 4. 9-7. 5 mm. Antennal Segments I-X punctate; I- 111 coarsely punctate; distal segments with
punctures very fine; Segment XI impunctate; head slightly longer than broad; orbital groove absent or present; temporal
lobe punctate; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotal carinae subequal at middie of length; both outer, inner carinae punctate; pronotum without setae; prosternum
without precoxal carinae.
Elytron relatively long, narrow; base of Stria IV with longitudinal scarp; in most specimens with one seta at apex of
Stria IV, one in subapical stride, one or both of these setae absent in some specimens; apex of Stria VII with several setae.
Shape of temporal lobes, pronotum, and calcars, punctuation of temporal lobes,
metasternum and abdominal sterna, and depth of stria and lateral pits of the abdomen vary
among the species of this complex. Among species from outside the Solomon Islands, the most
similar species is O. classicus of the Admiralty Islands. It resembles O. princeps in having the
tip of the median lobe pointed, but has a much more quadrate pronotum, with the base much
less narrowed.
Omoglymmius (sensu stricto) lindrothi new species
Fig. 176
Type material. - HOLOTYPE male, labelled: “SOLOMON IS. Guadalcanal, Kukum,
25-11-1963, P. Greenslade, 11.007, B.M. 1966-477” (BMNH). PARATYPES two males,
same data as holotype (BMNH); one female, Kukum, 19-4-1963, 5078; one female, Kukum,
8-12-1962, 3047; two males, one female (on same pin), Kukum, 15-11-1962, 2898; one male,
Honiara, Kukum, 20-2-62, 830; two males, one female (on same pin), Honiara, Kukum, 6-2-62,
669; one female, Honiara, Kukum, 4-11-61, 68;one male, Guadalcanal, Manna, 23-7-1962,
2507; two males (on same pin), Guadalcanal, Mt. Austen, 19-9-672, 1297 (BM 1963-5) (all
the preceding specimens collected by P. J. M. Greenslade, B. M. 1966-477)(all BMNH); one
female, labelled: “Guadalcanal, Tambalias, 30 km. W. Honiara, 22-V-64, sweeping, R.
Straatman” (BPBM). This species is apparently confined to Guadalcanal.
Revision of Omoglymmius Ganglbauer
233
Description. — Length 5. 0-6. 2 mm. Median lobe broad, its tip broadly rounded; frontal space broad, U-shaped, its
lateral margin curved; medial angles rounded, slightly separated; posteriomedial margin only slightly rounded, nearly
oblique; posteriolateral margin evenly curved; occipital angle obsolete; orbital groove represented by short but broad band
of pollinosity ended opposite middle of eye; temporal lobe with about 20 fine punctures.
Pronotum moderately long, length/greatest width 1.27; widest near middle; base slightly narrowed; apex distinctly
narrowed; lateral margins almost straight except near apex; lateral margin not sinuate anterior to hind angle; medial
margin of outer carina straight, not sinuate anterior to base; outer carina broadest well anterior to middle, evenly narrowed
posteriorly; basal part of inner carina narrow, its sides parallel; outer carina with 20-27 fine punctures; inner carina with
four to six very fine punctures.
Elytral striae moderately impressed, not pollinose except for pollinosity within punctures; latter moderately coarse;
female with Sternum IV of abdomen with lateral pit deep, semicircular; male with ventral tooth on anterior femur; female
with small ventral tooth or tooth entirely absent; hind calcar obtuse, subtruncate.
The subquadrate pronotum, with outer carina not at all sinuate near base and inner carina
with base very narrow, are the marks of this species. Of species from nearby islands, O.
modicus of Savo and Nggela, is closest in form of pronotum. However, this latter species has
the pronotum shorter, its lateral margins more distinctly curved, and its frontal space broader.
The notation “sweeping” on the specimen from Tambalis, is probably a mistake, as there are
no other records of Rhysodini from sweeping vegetation.
We dedicate this species to the memory of the eminent carabid taxonomist, Carl H.
Lindroth.
Omoglymmius (sensu stricto) modicus new species'
Fig. 177
Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Savo, 5-4-62, P.J.M.
Greenslade, 1327, B. M. 1966-477” (BMNH). PARATYPES two females (on same pin), one
male, one female (on same pin), same data as holotype, labelled “1390” (BMNH); one male,
same collector and locality as holotype, dated 6-4-62, “1455”(BMNH).
Description. — Length 5. 0-6. 3 mm. Median lobe broad, its tip broadly rounded; frontal space very broad,
U-shaped, its lateral margin deeply, evenly curved; medial angles obtuse, slightly separated; posteriomedial margin
oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove represented by short band of
pollinosity which scarcely extended past anterior margin of eye; temporal lobe with about 20 fine punctures.
Pronotum short, broad; length/greatest width 1.17; widest near middle; base slightly narrowed; apex markedly
narrowed; lateral margins curved; margin shallowly sinuate anterior to hind angle; medial margin of outer carina scarcely
sinuate anterior to base; outer carina broadest near middle, slightly narrowed to base; narrowed almost to point at apex;
basal part of inner carina less narrowed than in O. lindrothi, base narrowly truncate; outer carina with 22-25 fine
punctures; inner carina with six to eight fine punctures.
Elytral stria impressed, pollinosity limited to punctures; latter coarse, mostly elongate; female with deep, semicircular
lateral pits on Sternum IV; male with prominent ventral tooth on anterior femur; female with or without small femoral
tooth; hind calcar small, triangular, its apex obtuse.
The pronotum in this species is shorter and less quadrate than in O. lindrothi and the medial
margin of the temporal lobe is obtusely angulate opposite the median lobe. O. rusticus of
nearby Russell Islands, has the margins of the pronotum much more markedly curved and the
hind calcar is differently shaped. O. regius of Isabella has the outer carina more distinctly
narrowed posteriorly and the frontal space more nearly V-shaped.
Range. - Savo and Nggela Islands in the Solomons. In addition to the type series from Savo,
we have seen the following specimens from Nggela: one female, labelled “Florida Is., Nggela L, Haleta,
0-50 m., 18-X-1964, R. Straatman, light trap” (BPBM); three females (on same pin), labelled “Nggela, Toa, 8-1-62, 524,
P.J.M. Greenslade, B.M. 1966-477” (BMNH).
'see p. 254 for correction
Quaest. Ent., 1982, 18 (1-4)
234
Bell and Bell
Omoglymmius (sensu strict o) rusticus new species
Fig. 178
Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Russell Is., Loani,
31-5-1963, 6174, P. Greenslade, B.M. 1966-477” (BMNH). PARATYPES three females (on
same pin), labelled: “Russell Is., Barika 30-5-1963, 6169”; one male, one female (on same pin),
labelled: “Russell Is., Yandina, in logs, 22-24-11-1967, 13464” two males (on same pin).
Russell Islands, Yandina, same data as preceding entry. All specimens collected by P.J.M.
Greenslade, B.M. 1966-477 (all BMNH).
Description. — Length 5. 0-6. 3 mm. Median lobe broad, its tip rounded; frontal space very broad, U-shaped, its
margins deeply, evenly curved; medial angles narrow, slightly produced, obtuse; narrowly separated; posteriomedial
margin slightly emarginate; posteriolateral margin evenly curved; occipital angle indefinite; orbital groove scarcely
developed, but pollinose preorbital impression present; temporal lobe with about 20 very fine punctures.
Pronotum moderately long; length/ greatest width 1.24; widest near middle, base, apex strongly narrowed; lateral
margins markedly curved; margin not sinuate anterior to hind angle; medial margin of outer carina scarcely sinuate
anterior to base; outer carina widest anterior to middle, both base and apex strongly narrowed; basal part of inner carina
strongly narrowed, base narrowly truncate; outer carina with about 12 fine punctures; inner carina with six to eight fine
punctures.
Elytral striae impressed; punctures of Stria I, II fine, scarcely pollinose; those of outer striae coarser, distinctly
pollinose; female with deep, semicircular lateral pits on Sternum IV; male with small ventral tooth on anterior femur;
female with vestige of femoral tooth; hind calcar triangular, its apex obtuse, its upper margin curved.
This species contrasts with the two preceding ones in the strongly curved lateral margins of
the pronotum. The latter has the base almost as narrow as the apex. Form of the hind calcar
also differs from that of O. modicus of the nearby islands of Nggela and Savo.
Range. - Apparently identical forms are found in the Russell Islands and on San Cristobal.
In addition to the type series, from the Russells, we have studied the following specimens from
San Cristobal: one^male, one female (on the same pin), labelled; “Cristobal, Kinkia, 24-8-1962, P. Greenslade, 2193”
(BMNH). Future study may show that the forms on the two islands owe their similarity to
convergence, rather than close relationship.
Omoglymmius ( sensu stricto) manni new species
Figs. 179, 180
Type material. - HOLOTYPE male, labelled: “Auki, Sol. Isl., W. M. Mann” (MCZ).
PARATYPE female, same data as holotype (MCZ). The type locality is on Malaita Is.
Description. — Length 5. 7-6.0 mm. Median lobe broad, its tip broadly rounded; frontal space broad, its lateral
margin more shallowly curved than in O. modicus; medial angles obtuse, slightly produced, nearly contiguous;
posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle indistinct; orbital groove short but
broad band of pollinosity, ended near middle of eye; temporal lobe 10-15 fine punctures.
Pronotum moderately long, length/greatest width 1.19; widest near or slightly posterior to middle; base distinctly
narrowed; apex markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; medial
margin of outer carina shallowly sinuate anterior to base; outer carina curved, of nearly even width except for narrowed
apex; basal part of inner carina constricted; outer carina with 22-25 fine punctures; inner carina with eight or nine very
fine punctures.
Basal scarp of elytron pollinose; elytral striae impressed; coarsely punctate; pollinosity limited to punctures; both sexes
with shallow lateral pit on Sternum IV; male with prominent ventralTooth on anterior femur; female with minute one; hind
calcar with proximal margin slightly concave, apex obtuse (Fig. 180).
Shape of the pronotum in this species is close to that of O. rusticus , though the apex is more
narrowed. However, the latter species has the hind calcar differently shaped and the lateral pit
of Sternum IV of the female much deeper.
We dedicate this species to the collector, the myrmecologist and former director of the
National Zoological Garden, William M. Mann, who contributed many specimens of tropical
Revision of Omoglymmius Ganglbauer
235
Rhysodini to American museums.
Omoglymmius (sensu stricto) regius new species2
Figs. 182,190
Type material. - HOLOTYPE male, labelled: “SOLOMON IS., Isabel, Regi, 22-8-1963,
P. Greenslade, 9554, B.M. 1966-477“ (BMNH). PARATYPES one female, labelled: “Isabel,
Tatumba, 24-8-1963, P. Greenslade, 9587“ (BMNH); three males, one female (on same pin),
labelled: “Isabel, Tatumba, 2-8-1962; 2432, P. Greenslade, B.M. 1966-477“ (BMNH).
Description. — Length 5. 0-6. 6 mm. Median lobe broad, its tip rounded; frontal space moderately broad, less
U-shaped than in O. manni, its lateral margins more oblique, less emarginate; medial angles obtuse, not produced, well
separated; posteriomedial margin curved; posteriolateral margin curved; occipital angle absent; orbital groove pollinose,
extended to or beyond middle of eye, slightly developed; temporal lobe with 12-15 very fine punctures.
Pronotum moderately long; length/greatest width 1.19; widest slightly posterior to middle; base moderately narrowed
(less so than in O. manni)', apex strongly narrowed; lateral margins moderately curved, less so than in O. manni', margin
not sinuate anterior to hind angle; medial margin of outer carina shallowly sinuate anterior to base; outer carina oblique, of
nearly even width except at apex, where narrowed; basal parts of inner carinae less narrowed than in O. manni', outer
carina with about 20 fine punctures; inner carina with seven or eight fine punctures.
Basal scarp of elytron pollinose; striae impressed, coarsely punctate; pollinosity limited to punctures; female with deep
semi-circular lateral pits on Sternum IV; male pits fairly deep; male with prominent ventral tooth on anterior femur;
female with minute one; hind calcar small, triangular, tip obtuse (Fig. 190).
This species differs from O. manni in the more oblique, less curved margins of the pronotum
and in the more separated medial angles of the temporal lobes. In general, shape is similar to O.
gurneyi of Bougainville and Choiseul, but the latter has much more extensive pollinosity,
including completely pollinose elytral striae.
Omoglymmius (sensu stricto ) gurneyi new species
Figs. 183, 189
Type material. - HOLOTYPE male labelled: “BOUGAINVILLE I., Nov. 1944, A.B.
Gurney” (NMNH). PARATYPES one male, same locality and collector as type, dated
11- VI-44 (NMNH); one male, labelled: “Choiseul, Malangono, 25-8-1963, P. Greenslade,
9141, B.M. 1966-477“ (BMNH); one female, labelled: “Fauro I. Toumoa village, 30 m.,
1 2- 1 V- 1 964, P. Shanihan, light trap” (BPBM).
Description. — Length 6.2-7. 5 mm. Median lobe broad, its tip rounded; frontal space broad, its margin sharply bent
at middle; medial angles obtuse, not produced, well separated; posteriomedial margin curved; posteriolateral margin evenly
curved; occipital angle absent; orbital groove broadly pollinose, continued posteriorly as pollinose band along posterior
margin of temporal lobe; temporal seta located within this band; temporal lobe with 15-25 fine punctures.
Pronotum moderately long; length/greatest width 1.22; widest near middle; base slightly narrowed; apex markedly
narrowed; lateral margins slightly curved except where moderately markedly curved near apex; margin scarcely sinuate
anterior to hind angle; medial margin of outer carina sinuate anterior to base; outer carina slightly broadened at base,
narrowed at apex, otherwise of nearly even width; base of inner carina gradually narrowed; outer carina with 30-35 fine
punctures; inner carina with 12-17 fine punctures.
Basal scarp of elytron pollinose; striae impressed, deep; coarsely punctate; elytral striae, except for Stria V, with
punctures connected by continuous pollinosity; female with deep, semicircular lateral pit on Sternum IV; male with
shallow one; male with prominent ventral tooth on anterior femur; female with small one; hind calcar triangular, its apical
angle nearly rectangular, both proximal, distal margins straight.
This species is separated from the others by continuous pollinosity of the striae, and
extensive pollinosity along the posterior margin of the temporal lobe. It differs from all species
except O. scopulinus of Santa Cruz in the high number of punctures in the pronotum and
temporal lobes. O. princeps is sympatric with this species in Bougainville. It differs markedly in
the shape of pronotum, as well as in having elytral pollinosity confined to the punctures.
2see p. 254 for correction
Quaest. Ent., 1982, 18 (1-4)
236
Bell and Bell
The specimen from Choiseul has the pollinosity of the elytral striae and temporal margins
less developed, though still continuous. Otherwise, it seems identical to those from Bougainville.
We dedicate this species to the distinguished orthopterist, Dr. Ashley B. Gurney, who
collected the type specimen.
Omoglymmius ( sensu stricto) princeps new species
Fig. 184
Type material. - HOLOTYPE male, labelled: “SOLOMON Is., Bougainville (S). Kokure,
nr. Crown Prince Range, 900 m., VI-8-1956, coll. E. J. Ford, Jr” (BPBM).
Description. — Length 6.0 mm. Median lobe obtusely pointed posteriorly; frontal space broad, its margin
moderately curved at middle; medial angles obtuse, separated; posteriomedial margin oblique; posteriolateral margin
evenly curved; occipital angle distinct; orbital groove absent, but pollinose preorbital pit present; temporal lobe with about
20 fine punctures.
Pronotum moderately long; length/greatest width 1.20; widest near middle; base, apex very markedly narrowed;
lateral margins markedly curved; margin markedly sinuate anterior to hind angle; medial margin of outer carina sinuate
well anterior to base; outer carina narrower than inner carina at middle and also at base; outer carina widest anterior to
middle; inner carina constricted near base, latter broadened; outer carina with 10-11 fine punctures; inner carina with
three or four fine punctures.
Basal scarp of elytron not pollinose; striae impressed; strial punctures coarse, separated from one another by less than
diameter of one puncture; male with prominent ventral tooth on anterior femur; hind calcar triangular, its apex obtuse;
female unknown.
The markedly rounded pronotal margins, absence of orbital groove and marginal pollinosity
of the temporal lobe, and the lack of pollinosity between strial punctures separate this species
easily from the sympatric O. gurneyi. The most similar species is O. renutus from Rendova and
Mavovo. The latter species has a pronotum of similar shape, but has fewer punctures on the
temporal lobes and pronotal carinae, while the hind calcar has a different shape.
Omoglymmius (sensu stricto ) renutus new species
Fig. 185
Type material. - HOLOTYPE male, labelled: “SOLOMON IS., New Georgia, Mavovo,
30-12-1964, P. Greenslade (cacao?) 16183, B.M. 1966-477” (BMNH). PARATYPE female,
labelled: “SOLOMON IS., New Georgia, Rendova Is., 17-8-1963, P. Greenslade, B.M.
1966-477” (BMNH).
Description. — Length 5. 0-6. 4 mm. Median lobe obtusely pointed posteriorly; frontal space broad, its margin
moderately curved at middle; medial angles obtuse, nearly contiguous; posteriomedial margin oblique, slightly sinuate;
posteriolateral margin evenly curved; occipital angle obsolete; orbital groove absent, but pollinose preorbital pit present;
temporal lobe with 10-12 fine punctures.
Pronotum long, length/greatest width 1.31; widest near middle; base, apex very markedly narrowed; lateral margins
markedly curved; margin strongly sinuate anterior to hind angle; medial margin of outer carina sinuate anterior to base;
outer carina narrower than inner one at middle and also at base; outer carina widest anterior to middle; inner carina
narrowed anterior to base, latter rather broadly truncate; outer carina with about 10 very fine punctures; inner carina
impunctate.
Basal scarp of elytron not pollinose; striae impressed, rather finely punctate, punctures separated by more than
diameter of one of them; male with prominent ventral tooth off anterior femur; female with smaller one; hind calcar
triangular, its tip slightly lobate, truncate; female with deep semicircular lateral pit on Sternum IV; male with shallower
one.
The markedly rounded pronotal margins separate this species from all others except for O.
princeps of Bougainville. The latter species differs in having the inner carina punctate, the
temporal lobe with more punctures, and the hind calcar not truncate.
Revision of Omoglymmius Ganglbauer
237
Omoglymmius (sensu stricto) scopulinus new species
Fig. 186
Type material. - HOLOTYPE female, labelled: “SOLOMON IS., Santa Cruz, Reef Is.,
30-6-1962, P. Greenslade 2131, B.M. 1966-477” (BMNH).
Description. — Length 7.3 mm. Median lobe obtusely pointed posteriorly; frontal space moderately wide, its
margins moderately curved at middle; medial angles rounded, well separated; posteriomedial margin evenly curved;
posteriolateral margin evenly curved; occipital angle absent; orbital groove obsolete, ended slightly behind anterior margin
of eye; temporal lobe with 30-35 fine punctures.
Pronotum very long; length/greatest width 1.34, widest near middle; base, apex slightly narrowed; lateral margins
slightly curved; margin slightly sinuate anterior to hind angle; medial margin of outer carina not distinctly sinuate anterior
to base; outer carina distinctly narrower than inner one at middle; outer carina with base slightly narrower than that of
inner carina; inner carina constricted just anterior to base; latter slightly broadened; rather narrowly truncate; outer carina
with 18-21 fine punctures; inner carina with about 14 fine punctures.
Elytral striae impressed; strial punctures coarse; Intervals III, V more convex than others, in form of inconspicuous
raised carinae near base; female without ventral tooth on anterior femur; female with deep semicircular lateral pit on
Sternum IV; male unknown.
The long pronotum, the shape of the temporal lobe and its numerous punctures differentiate
this species from all others.
Omoglymmius ( sensu stricto ) mycteroides new species
Fig. 187
Type material. - HOLOTYPE male, labelled: “SOLOMON Is., New Georgia gp.
Kolombangara, hunda 2018, 1963, P. Greenslade, 8906, B.M. 1966-477” (BMNH).
PARATYPES two males (on same pin) labelled: “SOLOMON IS., Guadalcanal, Mt.
Tonapan, 3,500', 5-5-1963, P. Greenslade, 18120, B.M. 1966-477” (BMNH).
Description. — Length 6. 0-6. 7 mm. Antennal Segments 1, II coarsely punctate; Segments Ill-XI impunctate; head
nearly twice as long as wide preocular portion produced; median lobe elongate, its margins convergent in posterior half; its
tip obtusely pointed; frontal space narrow; medial angles obtuse, contiguous, posteriomedial margin slightly emarginate;
posteriolateral margin evenly curved; occipital angle distinct; antennal lobe far from eye; temporal lobe with anterior
portion in form of oblique, pollinose ridge; margin of head in form of deep preorbital impression between eye and antennal
lobe; orbital groove represented by short, broad pollinose impression ending opposite middle of eye; temporal lobe with
eight to 16 fine punctures, mostly near temporal seta; one temporal seta; postorbital, suborbital tubercles absent; eye large,
round.
Pronotum moderately long, length/greatest width 1.20; widest near middle; base slightly narrowed; apex markedly
narrowed; lateral margins curved; margin not sinuate anterior to hind angle; inner carina slightly broader than outer
carina at middle; medial margin of outer carina angulate, sinuate anterior to base; outer carina curved, of nearly even
width except at extreme apex and base; inner carina widest near middle, narrowed to base, latter truncate; outer carina
with 23-25 coarse punctures; inner carina with four or five very fine punctures; pronotum without setae; prosternum
without precoxal carinae.
Elytron rather elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal pollinose
scarp; Intervals III, V raised above others, subcarinate in anterior third; Stria IV with one seta near apex; subapical stride
with one seta; Stria VII with several setae near apex; metasternum coarsely punctate; abdominal Sterna 1I1-V coarsely,
sparsely punctate, punctures in form of irregular transverse row, especially near midline; lateral pit of Sternum IV barely
discernible in either sex; male with ventral tooth on anterior femur; middle calcar acute, nearly as long as spur; hind calcar
small, its apex an obtuse angle; female unknown.
This isolated species differs strikingly from other species of the Solomon Islands in having a
long snout. In this respect, it is superficially similar to O. bucculatus of Sumbawa and O.
nasalis of Buru. In head structure, it is intermediate; the anterior part of the temporal lobe
forms an oblique, pollinose ridge, as in O. nasalis , but there is no oblique groove latered to the
ridge, as in the latter species. It differs from both these species in the subcarinate bases of
Intervals III and V, a character shared with O. scopulinus , of the lindrothi group.
Quaest. Ent., 1982, 18 (1-4)
238
Bell and Bell
One might expect the populations on Guadalcanal and Kolombangara to represent distinct
species, as is so in other groups of Rhysodini in the area. There are minor differences in shape
of body and form of calcars, but these could be within the range of variation of a single
population. Pending collection of more specimens, we consider the forms on the two islands to
be conspecific.
Omoglymmius (sensu stricto) tabulatus new species
Figs. 188, 191
Type material. - HOLOTYPE male, labelled: “SOLOMON Is., Bougainville (S.) Boku-50
m., VI-5-56, coll. J.L. Gressitt” (BPBM). PARATYPE female, labelled: “SOLOMON IS.,
New Georgia gp., Kolombangara, nr. Kusi in log, 2-9-1965, leg. P. Naturaga, pres. P.J.M.
Greenslade, B.M. 1966-477” (BMNH).
Description. — Length 4. 3-4. 7 mm. Antennal Segments I, II coarsely punctate; Segments III-XI impunctate; head
slightly longer than wide; median lobe rather short, its apex obtusely rounded; frontal space small, narrow, U-shaped,
longer than wide; medial angle obtuse, slightly produced, contiguous; posteriomedial margin shallowly sinuate;
posteriolateral margins evenly curved; occipital angles distinct, rather far apart; orbital groove abbreviated, extended to
middle of eye; temporal lobe with 14-15 very coarse punctures; one temporal seta; postorbital, suborbital tubercles absent;
eye large, round.
Pronotum small, short, narrow; length/greatest width 1.31; lateral margins parallel except near base, apex; pronotum
thus hexagonal; outer carina formed of two planes meeting at a straight line, medial plane is sloped toward paramedian
groove, outer plane is nearly vertical; outer carina in dorsal view about 0.67 as broad as inner carina; outer carina with
about 16 coarse punctures; inner carina with three to seven coarse punctures; pronotum without setae; prosternum without
precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctured; base of Stria IV without longitudinal scarp; Stria
II with one to four setae; Stria IV with four or five setae; subapical stride with one seta; apex of Stria VII with several
setae; metasternum coarsely punctate; abdominal Sterna III-V with scattered coarse punctures; Sternum IV with deep,
semicircular lateral pit in both sexes; anterior femur with small, obtuse tooth in male, without tooth in female; middle
calcar obtuse, about 0.5 as long as spur; hind calcar triangular, longer than deep, its apex obtuse, its proximal margin
convexly curved (Fig. 191).
Form of the outer carina in this species is unique, with separate dorsal and lateral surfaces,
separated by a sharp edge. Among species from the Solomon Islands it is unique in its small
size and in having numerous elytral setae.
The specimen from Kolombangara is only provisionally associated with the holotype. It
differs in having fewer punctures on temporal lobe, these limited to margin; inner carina with
fewer punctures, and the second stria with four rather than one seta. These might be specific
differences, but more specimens are needed to confirm it.
Omoglymmius (sensu stricto) classicus new species
Fig. 192
Type material. - HOLOTYPE female, labelled: “ADMIRALTY IS., Los Negros, XI-45,
W. Wagner jr., D. Greether” (BPBM).
Description. — Length 6.0 mm. Antennal Segments I- 1 X punctate, distal segments very finely so; Segments X-XI
impunctate; median lobe short, lance-shaped, its apex obtusely pointed; frontal space large, almost twice as wide as long,
U-shaped, its margins markedly curved; medial angles obtuse, well separated; posteriomedial margin oblique, slightly
sinuate; posteriolateral margin evenly curved; occipital angles very obtuse; trace of orbital groove present, near anterior
margin of eye, posterior to pronounced preorbital pit; temporal lobe with about 23 fine punctures; one temporal seta;
postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately long; length/greatest width 1.24; widest near middle; base slightly narrowed; apex more
markedly narrowed; lateral margins weakly curved; margin markedly sinuate anterior to hind angle; inner carinae slightly
wider than outer ones at middle; medial margin of inner carina markedly sinuate anterior to base; outer carina widest
anterior to middle, markedly narrowed anteriorly; inner carina narrowed just anterior to base; base broadened, truncate,
Revision of Omoglymmius Ganglbauer
239
very finely pollinose; outer carina with about 22 fine punctures; inner carina with eight to 10 fine punctures; pronotum
without setae; prosternum without precoxal carina.
Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp very finely pollinose; base
of Stria IV with pollinose longitudinal scarp; Stria IV with one seta at apex; subapical stride with one seta; Stria VII
with several setae near apex; metasternum with most punctures near margins or midline, disc otherwise nearly
impunctate; abdominal Sterna II1-V with punctures scattered, well separated in medial portion, becoming coalescent
laterally; female with deep, round lateral pit in Sternum IV: female without ventral tooth on anterior femur; male
unknown.
This species resembles members of the lindrothi complex. The sinuation of the lateral
margin of the pronotum suggests O. princeps and O. renutus, but the lateral margins are much
less curved than in the latter two species. Another similar species is O. vicinus of New Guinea.
The latter species differs in the shape of the temporal lobe, and in having the median lobe
punctate.
Omoglymmius ( sensu stricto) vicinus (Grouvelle)
Fig. 193
Rhysodes vicinus Grouvelle 1895a: 157.
Omoglymmius ( sensu stricto) vicinus (Grouvelle) Bell and Bell 1978.
Type material. - HOLOTYPE female, labelled: “NOUV. GUINEE, Dorey, Baie de
Geelvink, Raffray & Maindron 78” (MNHN). Additional specimens listed in Grouvelle
(1903) are not authentic types, but merely additional localities.
Description. — Length 4. 6-6. 2 mm. Antennal Segments 1-IV coarsely punctate; Segments V-1X very finely
punctate; Segments X-Xl impunctate; median lobe short, oval, its apex obtusely rounded; median lobe with three to six
punctures; frontal space twice as broad as long, U-shaped, its margins markedly, abruptly curved; median angles obtuse,
well separated, lobate; posteriomedial margin oblique, emarginate; posteriolateral margin evenly curved; occipital angle
obtuse; orbital groove represented by a narrow line of pollinosity of varied length; antennal lobe separated from temporal
lobe by broader pollinose area than in O. classicus\ temporal lobe with 18-22 fine punctures; one temporal seta; postorbital,
suborbital tubercles absent; eye large round.
Pronotum moderately long, length/greatest width 1.22; widest near middle; base slightly narrowed; apex markedly
narrowed; lateral margins slightly curved; margin sinuate anterior to hind angle; inner carina slightly wider than outer
carina at middle; medial margin of outer carina slightly sinuate anterior to base; outer carina widest anterior to middle,
markedly narrowed to apex; inner carina narrowed just anterior to base; latter narrowly truncate; outer carina with 20-30
very fine punctures; inner carina with six to 12 very fine punctures; pronotum without setae; prosternum without precoxal
carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; base of Stria IV with longitudinal pollinose scarp;
Stria IV with one seta at apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum entirely
punctate; punctures of abdominal Sterna III-V very coarse, scattered, confluent near lateral margin; in some specimens
with very fine punctures medial to and posterior to the coarse principal punctures; female with deep semicircular lateral pit
on Sternum IV; male with shallow one; both sexes with ventral tooth on anterior femur; middle calcar minute, obtuse; hind
calcar triangular, its apex acute, its dorsal margin straight.
Among species from New Guinea, this one is recognized by the distinct sinuation anterior to
the hind angles and the U-shaped frontal space. No other species has both of these characters.
In form it is rather isolated among New Guinean species, and is really closer to O. classicus of
the Admiralty Islands, from which it differs in presence of punctures on the median lobe and in
shape of the temporal lobe.
Range. - North Coast of New Guinea from Geelvink Bay eastward, and on Goodenough
Island, in the D’Entrecasteaux Islands near the eastern end of New Guinea. In addition to the
type material, we have seen the following specimens: two males, Friedrich-Wilh.-hafen, Biro, 96 (MNHN);
one male, one female, Dutch New Guinea, Maffin Bay, IX-1944, coll. E. S. Ross (CAS); one female, Stephansort,
Astrolabe Bay, Biro 1900 (MNHN); two females, Goodenough Is., Gennais 1890, L. Loria (GEN).
Quaest. Ent., 1982, 18 (1-4)
240
Bell and Bell
Omoglymmius(sensu stricto ) oroensis new species
Fig. 194
Type material. - HOLOTYPE female, labelled: “PAPUA, N. G., Oro Bay, Dec. 43-Jan.
44, Darlington” (MCZ).
Description. — Length 7.0 mm. Antennal Segments I- IV coarsely punctate; Segments V-X more finely punctate;
Segment XI impunctate; head distinctly longer than wide; median lobe short, oval, its tip rounded; median lobe with a few
punctures; frontal space longer than broad, narrow, nearly V-shaped, its margins shallowly curved; medial angles obtuse,
markedly separated; posteriomedial margin oblique, short; posteriolateral margin bent near temporal puncture, its
posterior portion oblique; occipital angle very obtuse; orbital groove distinct, ended posterior to middle of eye; anterior
portion of temporal lobe a convex, pollinose ridge; temporal lobe with 14-16 rather fine punctures; one temporal seta;
postorbital, suborbital tubercles absent; eye large, round.
Pronotum rather short, subquadrate; length/greatest width 1.14; base slightly narrowed; apex markedly narrowed;
lateral margins slightly curved; margin scarcely sinuate anterior to hind angle; outer carina approximately 0.67 as wide as
inner carinae at middle; medial margin of outer carina sinuate just anterior to base; outer carina widest anterior to middle,
strongly narrowed at apex; inner carina narrowed to base; latter narrowly truncate; outer carina with 10-20 fine punctures;
inner carina with three or four fine punctures; pronotum without setae; prosternum without precoxal carina.
Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV
with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several
setae near apex; metasternum entirely punctate; abdominal Sterna III-V coarsely punctate; punctures confleunt near
lateral margin; female with lateral pit on Sternum IV shallow; female with minute ventral tubercle on anterior femur.
This species is somewhat similar to O. vicinus except for shape of the pronotum and the
more extensive pollinosity between antennal and temporal lobes.
Variation. - We provisionally include two other specimens with the holotype, although they
differ from it in some respects. One is a male, length 6.0 mm. labelled: “PAPUA, N. G.,
Dobodura, Mar. -July 1944, Darlington” (MCZ), It has a wider median lobe than the holotype.
It has a ventral tooth on the anterior femur. The middle calcar is very small and obtuse, while
the hind calcar is triangular, with an obtuse apex, and with its distal margin distinctly
emarginate. The second specimen is a female, length 7.2 mm., labelled: “PAPUA-Kokoda, 200
ft. VI-1933, L. E. Cheesman, B. M. 1933-456” (BMNH). It differs from the holotype in having
the temporal lobe more evenly rounded posteriorly, and in having two temporal setae on each
side. More material will be necessary to ensure that these specimens are really variants of O.
oroensis.
Omoglymmius (sensu stricto ) fringillus new species
Fig. 195
Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NE) Huon Peninsula,
Finschhafen, 150 m., 1 4-1 V- 1 963, Sedlacek” (BPBM). PARATYPES one male, same data as
holotype (BPBM); one female, labelled; “NEW GUINEA (NE), Eliptamin Valley, 1200-1350
m., July 1-15, 1959, W. W. Brandt” (BPBM); one male, labelled: “Lae, Aug. 1944, F. E.
Skinner” (BPBM); one female, labelled: “Lae 24-8-65”(NMNZ).
Description. — Length 6. 2-7.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-X finely punctate;
Segment XI impunctate; head as broad as long; median lobe short, oval, its apex obtusely rounded; median lobe
impunctate; frontal space broader than long, strongly U-shaped, margins parallel anteriorly, abruptly curved, nearly
transverse posteriorly; medial angles obtuse, nearly rounded, separated, posteriomedial margin curved into posterior
margin; posteriolateral margin curved into posterior margin; occipital angle absent; antennal lobe separated from temporal
lobe by broad pollinose area; orbital groove distinct, extended to posterior margin of eye; temporal lobe with 15-17 fine !
punctures; posterior margin of temporal lobe with narrow fringe of pollinosity; one temporal seta; postorbital, suborbital
tubercles absent; eye large, round.
Pronotum short, quadrate; length/greatest width 1.16; widest near middle; base scarcely narrowed; apex markedly j
narrowed; lateral margins nearly parallel except near apex, where curved, markedly convergent; margin not sinuate
anterior to hind angle; outer carina distinctly wider than inner one at middle; medial margin of outer carina sinuate
Revision of Omoglymmius Ganglbauer
241
anterior to base; outer carina widest anterior to middle; apex markedly narrowed; inner carina narrowed to base, latter
very narrowly truncate; outer carina with 22-27 fine punctures; inner carina with five to seven fine punctures; pronotum
without setae; prosternum without precoxal carinae.
Elytra relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV
with longitudinal pollinose scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with
several setae near apex; metasternum coarsely punctate; abdominal Sterna III-V very coarsely punctate, punctures
confluent near lateral margin on Sterna III-VI; both sexes with shallow lateral pits on Sternum IV; male with
prominent ventral tooth on anterior femur; female with small tooth; middle calcar small, obtuse; hind calcar triangular,
its apex sharp; both distal and proximal margins straight.
This is a species with deep, coarsely punctate striae, a broad head, and a quadrate
pronotum. Among New Guinean species it is closest to O. oroensis, but differs from the latter
in having a shorter, broader head, a more nearly quadrate pronotum and an impunctate median
lobe. Like the two preceding species, it inhabits the lowlands of the north coast.
Omoglymmius (sensu stricto ) puncticornis new species
Fig. 196
Type material. - HOLOTYPE male, labelled: “NEW GUINEA:PAPUA, Kiunga, Fly
River, VII- 1 5-2 1 , Wm. W. Brandt” (BPBM). PARATYPES one male, labelled: “Nuova
Guinea, Fly River, 1876-77, L. M. D'Albertis” (MNHN), one female same label as paratype
(GEN).
Description. — Length 5. 8-6. 5 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head
distinctly longer than wide; median lobe short, its tip obtuse; frontal space broader than long, U-shaped, its lateral margins
curved; medial angles obtuse, narrowly separated; posteriomedial margin curved into posteriolateral margin; occipital
angle absent; preorbital pollinose impression distinct, but orbital groove absent; antennal lobe separated from temporal
lobe by rather broad pollinose area; temporal lobe with 22-27 punctures; one temporal seta; postorbital, suborbital
tubercles absent; eye large, round.
Pronotum rather short, length/greatest width 1.13; widest near middle; base moderately narrowed; apex markedly so;
lateral margins curved; margin not sinuate anterior to hind angle; inner carina slightly wider than outer carina at middle;
medial margin of outer carina scarcely sinuate anterior to base; outer carina widest anterior to middle, markedly narrowed
to apex; inner carina constricted anterior to base, latter pollinose; outer carina with about 23 rather coarse punctures; inner
carina with seven to 10 finer punctures; pronotum without setae; prosternum without precoxal carinae.
Elytron relatively long, narrow; striae impressed, coarsely punctate; transverse basal scarp pollinose; base of Stria IV
with pollinose longitudinal scarp; Stria IV with one seta near apex; subapical striole with one seta; Stria VII with several
setae near apex; metasternum entirely punctate; abdominal Sterna III-V with coarse, scattered punctures; both sexes with
deep semicircular lateral pits on Sternum IV; male with prominent ventral tooth on anterior femur; middle calcar very
small, obtuse; hind calcar triangular, its apex obtuse, its proximal margin straight.
The very coarse punctures on the outer antennal segments are distinctive. This species
differs from other similar species from New Guinea in having the lateral margins of the
pronotum curved, but without a sinuation anterior to the hind angle. Curved margins eliminate
O. fringillus and O. oroensis, while the absence of a sinuation eliminates O. vicinus , as do the
coarse punctures of the outer antennal segments. O. viduus of the Kei Islands, is the most
similar extralimital species. It differs in having the posteriomedial margin of the temporal lobe
sinuate, and elytral punctures much finer.
Variation. - A pair (male GEN, female MNHN) labelled “Ighibirei, New Guinea,
VII-VIII 90, Loria”, resemble O. puncticornis in most respects, so we provisionally include
them in the species. However, the inner carinae of the pronotum are entirely impunctate; and
the abdominal punctures tend to form a single transverse row on each sternum. We have been
unable to locate Ighibirei. These specimens may represent another species, but final judgment
must await the collection of more specimens.
Quaest. Ent., 1982, 18 (1-4)
242
Bell and Bell
Omoglymmius (sensu stricto) trepidus new species
Fig. 197
Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NE), Wau, Morobe
Dist., 1200 m„ 1 5-30-IX- 1 962, J. Sedlacek” (BPBM).
Description. — Length 6.0 mm. Antennal Segments 1- 1 V coarsely punctate; Segments V-XI very finely punctate;
head distinctly longer than wide; median lobe short, impunctate, its tip very obtuse, almost rounded; frontal space slightly
broader than long, nearly V-shaped, its margins only shallowly curved; medial angles obtuse, slightly lobate;
posteriomedial margin emarginate; posteriolateral margin evenly curved; occipital angles very obtuse; preorbital pollinose
impression very prominent, continued posteriorly as very short orbital groove; antennal lobe separated from temporal lobe
by short, narrow antennal groove; temporal lobe with 21-24 punctures; one temporal seta; postorbital, suborbital tubercles
absent; eye large, round.
Pronotum rather short, length/greatest width 1.17; widest near middle; base moderately narrowed; apex markedly
narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; inner carina slightly wider than
outer carina at middle; medial margin of outer carina shallowly sinuate anterior to base; outer carina anterior to middle,
very markedly narrowed to apex and to base; latter rather narrowly truncate; inner carina constricted near base, latter
broadly truncate; outer carina with 16-18 very fine punctures; inner carina impunctate; pronotum without setae;
prosternum without precoxal carinae.
Elytron rather short, broad; striae shallowly impressed, strial punctures rather fine; especially in inner striae;
transverse basal scarp iridescent, not pollinose; base of Stria IV with very short longitudinal pollinose scarp; Stria IV
without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum largely finely
punctate, but with impunctate area at either side of midline; abdominal Sterna III-V with scattered punctures; with deep,
semicircular lateral pit on Sternum IV; with ventral tooth on anterior femur; middle calcar scarcely evident; hind calcar
triangular, its apex rounded; female unknown.
The large preorbital pit and the rounded median lobe separate this species from O. patens
and O. cavea , while the presence of a small basal carina on Stria IV and rounded, rather than
oblique lateral pits of Sternum IV, separate it from O. sectatus.
Omoglymmius ( sensu stricto ) patens new species
Fig. 198
Type material. - HOLOTYPE female, labelled: “Maffin Bay, Dutch New Guinea,
IX- 1944, E.S. Ross” (CAS).
Description. — Length 5.2 mm. Antennal Segments I-IV coarsely punctate; Segments V-X very finely punctate;
Segment XI impunctate; head distinctly longer than wide; median lobe short, with six scattered punctures; median lobe
lance-shaped, its tip obtusely pointed; frontal space broader than long, nearly V-shaped, its lateral margins shallowly
curved; medial angles obtuse, well separated; posteriomedial margin oblique; anteriomedial margin evenly curved; occipital
angle indistinct; preorbital pollinose impression small; orbital groove absent; antennal groove short, narrow; temporal lobe
with 23-27 fine punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately long, length/greatest width 1.22; widest near middle, base moderately narrowed; apex more
markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; marginal groove dilated,
much broader than in O. trepidus ; outer carina narrow, about 0.5 as wide as inner carina at middle; medial margin of outer
carina scarcely sinuate anterior to base; outer carina widest anterior to middle; slightly narrowed at base; markedly
narrowed at apex; inner carina constricted anterior to base; latter broad, truncate; outer carina with 18-22 fine punctures;
inner carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron rather long, narrow; striae impressed; strial punctures coarse; transverse basal scarp glabrous; base of Stria IV
with pollinose longitudinal scarp rather long, its length equal to width of two elytral intervals; Stria IV without seta near
apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum coarsely punctate near margins,
otherwise impunctate; abdominal Sterna III-V with punctures coarse, sparse, in form of irregular transverse row on each
sternum; not confluent near lateral margins; female with deep, semicircular lateral pits on Sternum IV; female with
prominent ventral tooth on anterior femur; male unknown.
Outer carinae of the pronotum are narrower than in any other member of the subgenus in
New Guinea. This easily separates this species from the two most similar species, O. trepidus
and O. cavea.
Revision of Omoglymmius Ganglbauer
243
Omoglymmius ( sensu stricto) cavea new species
Fig. 199
Type material. - HOLOTYPE female, labelled: “NEW GUINEA, Ramoi II., Beccari,
1875” (GEN).
Description. — Length 5.9 mm. Antennal Segments I- 1 1 coarsely punctate; Segments 1II-XI very finely punctate;
head distinctly longer than broad; median lobe short, impunctate, lance-shaped, its tip obtusely pointed; frontal space
broader than long, nearly V-shaped, its lateral margins shallowly curved; medial angles obtuse, well separated;
posteriomedial margin oblique, shallowly emarginate; posteriolateral margin evenly curved; occipital angle obtuse;
preorbital pollinose impression small; orbital groove absent; antennal groove short, narrow; temporal lobe with 17-18 fine
punctures; one temporal seta; postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately long, length/greatest width 1.30; widest near middle; base moderately narrowed; apex markedly
narrowed; lateral margins markedly curved; margin shallowly sinuate anterior to hind angle; marginal groove not dilated;
outer carina slightly narrower than inner one at middle; medial margin of outer carina scarcely sinuate anterior to base;
outer carina widest anterior to middle, slightly narrowed at base, markedly narrowed at apex; inner carina constricted
anterior to base; latter broad, truncate; outer carina with 16-18 rather coarse punctures; inner carina impunctate;
pronotum without setae; prosternum without precoxal carinae.
Elytron rather long, narrow; striae impressed; strial punctures moderate, finer than in O. patens ; transverse basal scarp
glabrous; base of Stria IV with rather short pollinose longitudinal scarp; Stria IV with seta near apex; subapical stride
with one seta; Stria VII with several setae near apex; metasternum entirely punctate; abdominal Sterna III-V rather
densely, finely punctate, punctures scattered; female with deep triangular lateral pits on Sternum IV, these strongly
narrowed posteriorly; with ventral tooth on anterior femur; male unknown.
This species resembles O. trepidus and O. patens in having inner carinae impunctate, base of
Stria IV carinate, and anteriomedial margin of the temporal lobe shallowly sinuate. It differs
from the former in having the tip of the median lobe pointed and preorbital pollinose impression
very small, and from the latter species in having the outer carinae broad, marginal groove not
dilated and lateral pits of Sternum IV triangular. Shape of lateral pits is intermediate between
the semicircular one of most Omoglymmius and the narrow oblique ones of O. sectatus.
Omoglymmius (sensu stricto ) sectatus new species
Figs. 200, 201
Type material. - HOLOTYPE male, labelled: “NEW GUINEA:(NE) Mt. Missim,
950-1000 m., 10-VIII-64, J. Sedlacek” (BPBM). PARATYPE female, labelled: “NEW
GUINEA (NE), Wau. Morobe Dist., 1200 m., 2-VI-62, J. & M. Sedlacek” (BPBM).
Description. — Length 5.8 mm. Antennal Segments I-IV coarsely punctate; Segments V-XI finely, irregularly
punctate; head distinctly longer than broad; median lobe short, impunctate, lance-shaped, its tip narrowly rounded; frontal
space longer than broad, V-shaped, its margin shallowly curved; medial angles obtuse, well separated; posteriomedial
margin oblique; posteriolateral margin evenly curved, occipital angle scarcely evident; antennal groove narrow, its lateral
end scarcely broadened; orbital groove absent; temporal lobe with 15-21 fine punctures; one or two temporal setae;
postorbital, suborbital tubercles absent; eye large, round.
Pronotum moderately long, length/greatest width 1.25; widest slightly anterior to middle; base moderately narrowed;
apex markedly narrowed; lateral margins markedly curved; margin not sinuate anterior to hind angle; outer carina
subequal to inner carina at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina widest at
anterior third of length, moderately narrowed, subtruncate at apex; inner carina constricted anterior to base; latter broad,
truncate; outer carina with 10-12 very fine, irregularly scattered punctures; inner carina with two or three very fine
punctures; pronotum without setae; prosternum without precoxal carinae.
Elytron rather narrow, elongate; striae impressed; strial punctures coarse; base of Stria IV without longitudinal scarp;
Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum
coarsely punctate in midline and near margins; disc otherwise impunctate; abdominal Sterna III-V with coarse, scattered
punctures; both sexes with narrow, oblique, slit-like lateral pit on Sternum IV; male with ventral tooth on anterior femur;
female without such tooth; middle calcar very small, acute (Fig. 201) hind calcar triangular, its apex acute, its proximal
margin straight, distal margin emarginate.
This small, narrow species is easily identified by the oblique, slit-like lateral pits on Sternum
IV.
Quaest. Ent., 1982, 18 (1-4)
244
Bell and Bell
Omoglymmius (sensu stricto) ephemeris new species
Figs. 202, 209
Type material. - HOLOTYPE male, labelled: “NEW GUINEA (NW), Nabire, S.
Geelvink Bay, 0-30 m., 2-9-VII-1962, J. L. Gressitt coll,” (BPBM). PARATYPE female,
labelled: “NEW GUINEA: Papua, Kiunga, Fly River, VII-23-25- 1 957. coll. Wm. W. Brandt”
(BPBM).
Description. — Length 5.0 mm. Antennal Segments I-IV coarsely punctate; Segments V-XI finely punctate; head
1.3 longer than broad; median lobe short, coarsely punctate, lance-shaped, its tip obtusely pointed; frontal space broader
than long, V-shaped, lateral margins shallowly curved; medial angles obtuse, well separated; posteriomedial margin
oblique; posteriolateral margin evenly curved; occipital angle distinct; antennal groove narrow, its lateral end not
broadened; orbital groove absent; temporal lobe with 25-30 rather coarse punctures; one temporal seta; postorbital,
suborbital tubercles absent; eye large, round.
Pronotum moderately long, length/greatest width 1.21; widest near middle; base moderately narrowed; apex strongly
so; lateral margins markedly curved; margin not sinuate anterior to hind angle; outer carina subequal to inner carina at
middle; medial margin of outer carina not sinuate anterior to base; outer carina widest anterior to middle, moderately
narrowed, rounded anteriorly; median margin of outer carina indistinct opposite basal impression; inner carina constricted
anterior to base, latter very broadly truncate, nearly closing paramedian groove posteriorly; outer carina with 13-15
punctures; inner carina with five or six rather fine punctures; pronotum without setae; prosternum without precoxal
carinae.
Elytra narrow, elongate; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria IV with
one seta near apex; subapical stride with one seta; Stria VII with one seta near apex; metasternum coarsely punctate;
abdominal Sterna III-V with coarse scattered punctures; both sexes with deep lateral pits on Sternum IV; lateral pits
nearly semicircular, but anteriomedial margin oblique; male with distinct ventral tooth on anterior femur; female with very
small, obtuse one; middle calcar acute, nearly as long as spur (Fig. 209); hind calcar triangular, its apex acute, slightly
lobate, its proximal margin shallowly sinuate; its distal margin nearly straight.
This is a small, narrow species which lacks the basal scarp on Stria IV. It differs from O.
sectatus having the lateral pits of the abdomen nearly round and the hind calcar much longer
and more acute. Although the two specimens come from distant localities, they are markedly
similar, suggesting that this is a widely distributed lowland species.
Omoglymmius (sensu stricto) gracilicornis (Grouvelle)
Figs. 203,210
Rhysodes gracilicornis Grouvelle 1895a: 157.
Omoglymmius ( sensu stricto ) gracilicornis (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE male, labelled: “Baie du Geelvink, Raffray et Maindron 78”
(MNHN).
Description. — Length 4.5 mm. Antennal punctures not recorded; head slightly longer than broad; head nearly as
broad as pronotum; median lobe short, impunctate, its tip obtuse; frontal space as long as broad, V-shaped, its lateral
margins only slightly curved; medial angles nearly rectangular, well separated; posteriomedial margin oblique;
posteriolateral margin evenly curved; occipital angle distinct; antennal groove narrow, its lateral end not broadened; orbital
groove absent; temporal lobe with 10-12 punctures restricted to lateral half; one temporal seta; postorbital, suborbital
tubercles absent; eye round, rather small.
Pronotum moderately elongate; length/greatest width 1.30; widest near middle; base slightly narrowed; apex
moderately narrowed; lateral margins slightly curved; margin not sinuate anterior to hind angle; outer carina equal in
width to inner carina at middle; medial margin of outer carina not sinuate anterior to base; outer carina widest anterior to
middle; subtruncate apex; inner carina constricted anterior to base; latter narrowly truncate; inner and outer carina
entirely without punctures; pronotum without setae; prosternum without precoxal carinae.
Elytra rather elongate, narrow; striae impressed, coarsely punctate; base of Stria IV without longitudinal scarp; Stria
II with one seta near apex; Stria IV with two to four setae; subapical striole without seta; Stria VII with several setae near
apex; metasternum with coarse punctures near lateral margins, a few coarse punctures in midline, otherwise impunctate;
abdominal Sterna III-V with coarse punctures in form of transverse row on each sternum; with deep semicircular lateral
pit on Sternum IV: without ventral tooth on anterior femur; middle calcar very small, obtuse; hind calcar prominent, its
apex broadly rounded; female unknown.
Revision of Omoglymmius Ganglbauer
245
This small, narrow species has more elytral setae than any similar species from New
Guinea, and is also recognized by impunctate pronotal carinae and broadly rounded hind calcar
(Fig. 210). A second specimen (MNHN), from Astrolabe Bay, labelled as R. gracilicornis by
Grouvelle, is a different species, O. vicinus (Grouvelle).
Omoglymmius (sensu stricto) follis new species
Figs. 204,212
Type material. - HOLOTYPE female, labelled: “NEW GUINEA (NE), Wau, Morobe
Distr., 1700-1800 m., 7-X-1962, J. & M. Sedlacek” (BPBM). PARATYPES one male, one
female, labelled: “N. GUINEA. Okapa, Putosa, 2-11-1964, R. Hornabrook” (NMNZ).
Description. — Length 7. 8-8. 8 mm. Antennal Segments I, 11 coarsely punctate; Segments 1I1-X very finely,
sparsely punctate; Segment XI impunctate; head slightly longer than broad; median lobe short, broad, nearly
quadrangular; its tip broadly truncate, with narrow pollinose posterior border; frontal space broader than long, its
anteriomedial margin curved at middle; medial angles obtuse, well separated; posteriomedial margin oblique; posterior
margin transverse, bordered with pollinosity, meeting posteriolateral margin at obtuse angle near temporal seta; antennal
groove broad, dilated at lateral end in form of pollinose preorbital impression; latter extended posteriorly medial to eye for
short distance, in form of vestige of orbital groove; temporal lobe with seven or eight extremely minute punctures, visible
only under high magnification; one temporal seta; postorbital tubercle small, about 0.25 as long as eye, visible only in
lateral view; eye large, round.
Pronotum short, length/greatest width 1.04, widest near middle; base slightly narrowed, apex very markedly
narrowed; lateral margins slightly curved in basal half, becoming markedly curved near apex; margin shallowly sinuate
anterior to hind angle; outer carina slightly narrower than inner carina at middle; outer carina broadest at base, slightly
constricted anterior to base, slightly broader to anterior third, tapered to point at apex; inner carinae long oval, tapered to
point at base; outer carina with 12-15 very minute punctures, visible only at high magnification; inner carina with about 12
equally minute punctures; pronotum without setae; prosternum without precoxal carinae.
Elytra rather narrow, elongate; striae not impressed; strial punctures coarse; base of Stria IV with longitudinal scarp;
transverse basal scarp of elytron pollinose from base of Stria IV to suture; Stria IV without seta at apex; subapical stride
with one or no setae; Stria VII with few setae near apex; metasternum punctate near lateral and posterior margins, its disc
impunctate; abdominal Sterna III-V with punctures scattered, fine near midline, coarse, coalescent laterally; both sexes
with deep, semicircular lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without this tooth;
middle calcar minute, obtuse; hind calcar triangular, its apex obtuse, distance from tibial spur to proximal end of calcar
more than twice distance from tip of calcar to its base.
This large species has the striae coarsely punctate but not impressed, and has a small
postorbital tubercle, visible only in lateral view. The most similar species is O. iridescens. The
latter species has the median lobe sharply pointed posteriorly, rather than truncate, outer
carinae not suddenly dilated at base, and medial angles of temporal lobe contiguous.
Omoglymmius (sensu stricto ) iridescens new species
Figs. 205,211
Type material. - HOLOTYPE male (left outer pronotal carina malformed), labelled:
“NIEUW GUINEA Exp., KNAG 1939, Araboebivak, 19-X-1939” (LEI). PARATYPE
female, labelled: “Nieuw Gunea Exp., KNAG 1939, Koteboe Vallei ann Paniai meer,
24- VIII- 1939” (LEI).
Description. — Length 6. 6-6. 9 mm. Antennal Segments I- 1 1 coarsely punctate; Segments III-X very finely
punctate; Segment XI impunctate; head longer than broad; median lobe short, broad, rhomboid; posteriolateral margins
oblique, tip angulate; frontal space slightly broader than long, V-shaped, its lateral margins oblique, not sinuate; medial
angles nearly rectangular, contiguous; posteriomedial margin slightly sinuate; posteriolateral margin evenly curved;
occipital angle absent; posterior margin of temporal lobe narrowly bordered with pollinosity; antennal groove narrow, not
dilated laterally; orbital groove absent; temporal lobe with five or six punctures near eye; one temporal seta; postorbital
tubercle small, about 0.20 as long as eye, visible only in lateral view; eye large, round.
Pronotum short; length/greatest width 1.05, widest at base, lateral margins oblique, slightly convergent except near
curved apex, markedly convergent; apex markedly narrowed; outer carina slightly narrower than inner carina at middle;
Quaest. Ent., 1982, 18 (1-4)
246
Bell and Bell
outer carina broadest at base, gradually evenly narrowed from there nearly to more abruptly narrowed apex; inner
carina long, oval, tapered to obtuse point at base; pronotal carinae impunctate; pronotum without setae; prosternum
without precoxal carinae.
Elytra rather narrow, elongate; striae not impressed; strial punctures fine, separated by several times the diameter
of one puncture, especially fine in lateral striae and at bases of all striae; elytral intervals flat, iridescent; base of Stria
IV with longitudinal scarp; transverse basal scarp of elytron pollinose from base of Stria IV to suture; Stria IV with one
seta near apex; subapical stride with one seta; apex of Stria VII with several setae; metasternum with blue reflections,
impunctate except for one row of punctures along each lateral margin, cluster of punctures in posterior part of midline;
abdominal Sterna III-V with scattered punctures which are not coalescent laterally; both sexes with moderately deep
semicircular lateral pit on Sternum IV; male with ventral tooth on anterior femur; female without this tooth; middle
calcar acute, about 0.33 as long as spur; hind calcar triangular, its apex angulate, obtuse; distance from tribal spur to
proximal end of calcar about 1.5 longer than distance from tip of calcar to its base.
A large species with a broad pronotum, resembling O. follis in having a small postorbital
tubercle, visible only in lateral view, but differing in having the pronotum broadest at base, the
median lobe pointed posteriorly, the striae finely punctate, and in many other details.
Omoglymmius ( sensu stricto ) massa new species
Figs. 206, 213
Type material - HOLOTYPE male, labelled: “NEW GUINEA (NE), Eliptamin Valley,
1200-1350 m., July 1-15, 1959, W. W. Brandt coll.” (BPBM). PARATYPES one female, same
locality and collector as type but dated Aug. 16-30, 1959 (BPBM); one female, Wau, Morobe
Dist., 1200 m., 14-IX-1961, Sedlacek & native (BPBM); one female, Koibuga, 1500 m.,
3- VII- 1 963, coll. H. W. Clissold (BPBM); one female, 13 km. SE Okapa, 1650-1870 m.,
26-VIII-1964, J. & M. Sedlacek(BPBM).
Description. — Length 6. 5-8. 2 mm. Antennal Segments I, II coarsely punctate; Segments III-X very finely
punctate; Segment XI impunctate; head slightly longer than wide; median lobe short, oval, its apex broadly rounded;
frontal space broader than long, V-shaped, its margins oblique, very shallowly sinuate; medial angles obtuse, very narrowly
separated; posteriomedial margin oblique; posterior margin transverse, bordered with pollinosity, meeting posteriolateral
margin at obtuse angle near temporal seta; antennal groove rather broad, connected to very short orbital groove which ends
just posterior to anterior margin of eye; temporal lobe with six to eight very fine punctures near medial margin of eye; one
temporal seta or no temporal seta, in some specimens present unilaterally; postorbital tubercle about 0.4 as long as eye,
clearly visible in dorsal view; eye large, round.
Pronotum, length/greatest width 1.10; widest near middle, base slightly narrowed; apex markedly narrowed; lateral
margins slightly curved except near markedly curved apex, convergent; lateral margin scarcely sinuate anterior to hind
angle; outer carina slightly narrower than inner carina at middle; outer carina broadest at base, narrowed just anterior to
base, then nearly parallel-sided to anterior 0.3, tapered to apex; inner carina long, oval, tapered to point at base; both pairs
of carinae impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytra rather narrow, elongate; stria not impressed; strial punctures moderately coarse, round; base of Stria IV with
longitudinal pollinose scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria II; Stria IV
without seta near apex; subapical striole with one seta; apex of Stria VII with several setae; metasternum with row of
punctures along each lateral margin; disc of metasternum impunctate, with bluish opalescence; abdominal Sterna III-V
with coarse punctures in form of irregular transverse row on each sternum, punctures coalescent near lateral margin;
female with deep oval lateral pit on Sternum IV; male with shallow, ill-defined lateral pit on Sternum IV; male with
ventral tooth on anterior femur; female without this tooth; middle calcar very minute, obtuse; hind calcar small, triangular,
its apex obtusely angled, its proximal margin longer than distal margin.
This species and O. denticulatus have the postorbital tubercle of moderate size, larger than
that of the two preceding species, and clearly visible from above, but smaller than in O.
auratus. This species differs from O. denticulatus in having the median lobe broadly oval, a
slightly larger postorbital tubercle and more produced medial angles on the temporal lobe.
Range. - Mountains of northeastern New Guinea. In addition to the type material we
provisionally include the following specimens: two males, four females, labelled “Keefu, Okapa, Eastern
Highlands, New Guinea, 17-4-1965, coll. R. Hornabrook” (NMNZ); one female, labelled “Morae, 6000, Eastern
Highlands, Kuku Kuku, 1-3-64, R. Hornabrook” (NMNZ).
Revision of Omoglymmius Ganglbauer
247
Omoglymmius ( sensu stricto) denticulatus new species
Figs. 207,214
Type material. - HOLOTYPE female, labelled: “NEW GUINEA (NW), Wissel Lakes,
Enarotadi, 1900-2000 m., 2-1 l-VII-62, coll. J. Sedlacek” (BPBM). PARATYPE female,
labelled: “N. Guinea, Hatam VII, Beccari, 1875, Rhysodes pulvinatus ” (GEN). It is not
conspecific with male lectotype of R. pulvinatus.
Description. — Length 7. 7-8. 5 mm. Antennal Segments I, II coarsely punctate; Segments III-X very finely,
sparsely punctate; Segment XI impunctate; head slightly longer than wide; median lobe moderately narrow, its tip
subtruncate; frontal space longer than wide, V-shaped; lateral margins oblique; medial angles rather sharp, well separated;
posteriomedial margin curved, extended to slightly oblique posterior margin; both margins bordered by pollinosity;
posteriolateral margin in form of obtuse angle with posterior margin; antennal groove rather broad, slightly dilated at
lateral end in form of preorbital impression; orbital groove absent; temporal lobe with six or seven fine punctures in lateral
half; one temporal seta; postorbital tubercle about 0.2 as long as eye, clearly visible in dorsal view; eye large, round.
Pronotum rather short, length/greatest width 1.14; broadest at base; apex strongly narrowed; lateral margins oblique
in posterior half, curved, markedly convergent in anterior half; lateral margin not sinuate anterior to hind angle; outer
carina slightly narrower than inner carina at middle; outer carina markedly sinuate just anterior to base, broadest at base,
narrowed to point at apex; inner carina long oval, tapered to point at base; both pairs of carinae entirely impunctate;
pronotum without setae; prosternum without precoxal carinae.
Elytra rather narrow, elongate; striae not impressed; strial punctures moderately coarse, round; base of Stria IV with
longitudinal pollinose scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria I; Stria IV
without seta near apex; subapical stride with one seta; Stria VII with several setae near apex; metasternum with row of
punctures along each lateral margin; disc of metasternum impunctate, with bluish opalescence; abdominal Sterna III-V
with coarse, scattered punctures; punctures coalescent near lateral margin; female with deep, oval lateral pit on Sternum
IV; male unknown.
This species is most like O. massa of the eastern part of New Guinea, but differs in the size
of the postorbital tubercle and the shape of the medial angle of the temporal lobe. It is also
similar to O. iridescens, which is sympatric or nearly sympatric with it. This species differs
from O. iridescens in having a larger postorbital tubercle. O. auratus , also of West Irian, is also
similar, but has the pronotum narrowed at the base and has much larger postorbital tubercles.
Omoglymmius (sensu stricto) auratus new species
Figs. 208,215
Type material. - HOLOTYPE male, labelled: “NEW GUINEA:Neth., Swart Val: W.
ridge 1800-2000 m., Nov. 19, 1958, J. L. Gressitt” (BPBM).
Description. — Length 7.0 mm. Antennal Segment I punctate, pollinose; Segments II-IV impunctate; Segments
V-X finely punctate; Segment XI impunctate; head, including postorbital tubercles, wider than long; median lobe short,
broadly rounded; frontal space V-shaped, longer than wide, margins oblique, scarcely sinuate; medial angles obtuse,
separated; posteriomedial margin curved evenly into posteriolateral margin; posterior margin bordered with pollinosity;
antennal groove rattier broad, slightly dilated at lateral end; orbital groove absent; temporal lobe with nine or 10 very fine
punctures in lateral half; one temporal seta; postorbital tubercle as deep as eye, about 0.9 as long as eye; postorbital
tubercles prominent in dorsal view, head width across them greater than width across eyes; eye large, round.
Pronotum moderately short, length/greatest width 1.13; widest near middle; base slightly narrowed; apex more
markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina slightly narrower than
inner one at middle; outer carina broadest near middle, very slightly narrowed at base, strongly narrowed anteriorly, apex
obliquely truncate; medial margin of outer carina not sinuate anterior to base; inner carina long-oval, tapered to point at
base; both pairs of carinae impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytra rather narrow, elongate; striae not impressed; strial punctures very fine, elliptical; base of Stria IV with short
pollinose longitudinal scarp; transverse basal scarp of elytron pollinose from base of Stria IV to base of Stria II; Stria IV
with one seta at apex; subapical striole with one seta; Stria VII with several setae near apex; metasternum nearly
impunctate, with a few fine punctures near lateral margins, microsculptured but not opalescent; abdominal Sterna III-V
with one transverse row of coarse punctures on each; those of III, IV widely interrupted at midline; that of V not
interrupted; with shallow lateral pit on Sternum IV; with very obtuse ventral tooth on anterior femur; middle calcar small,
obtuse; hind calcar small triangular, its apex obtuse, slightly lobate; female unknown.
Quaest. Ent., 1982, 18 (1-4)
248
Bell and Bell
This species is easily recognized by the large postorbital tubercles.
Omoglymmius (sensu strict o) sus new species
Figs. 216, 222
Type material. - HOLOTYPE male, labelled: “NUOVA GUINEA, Katau, L.M.
D’Albertis” (GEN). PARATYPES one male, two females, same label as holotype (GEN); one
female, labelled: “Nuova Guinea, Fly River, L.M. D’Albertis” (GEN); one damaged male with
this label is probably conspecific, but we think it best not to designate it as a paratype
(MNHN).
Description. — Length 6. 8-8.0 mm. Antennal Segments I- VI 1 1 coarsely punctate; Segments IX, X more finely
punctate; Segment XI impunctate; head distinctly longer than wide; median lobe lance-shaped, its tip very obtuse; frontal
space nearly as long as wide, V-shaped, its margins only shallowly curved; medial angles obtuse, well separated;
posteriomedial margin emarginate; posteriolateral margin evenly curved, occipital angle obtuse; antennal groove rather
broad, slightly dilated at lateral end; orbital groove absent; temporal with 17-18 fine punctures; one temporal seta; orbital
tubercle about 0.3 as long as eye, visible in dorsal view; eye large, round.
Pronotum elongate; length/greatest width 1.30; widest near middle; base slightly narrowed; apex very markedly
narrowed; lateral margins slightly curved posteriorly, markedly curved anteriorly; margin sinuate anterior to hind angle;
outer carina slightly narrower than inner one at middle; medial margin of outer carina angulate, not sinuate just anterior
to base; outer carina of nearly even width in posterior half, markedly narrowed near apex; inner carina narrowly
subtruncate at base; outer carina with 24-29 fine punctures; inner carina with 19-25 fine punctures; pronotum without
setae; prosternum without precoxal carinae.
Elytron rather narrow, elongate; striae impressed; strial punctures coarse; base of Stria IV with short longitudinal
pollinose scarp; basal transverse scarp pollinose from Stria IV to suture; one seta near apex of Stria IV, one seta in
subapical stride; about four setae near apex of marginal stria; metasternum entirely punctate; abdominal Sterna III-V
with coarse, scattered punctures; female with deep, narrowly oval, oblique lateral pit on Sternum IV; male with shallow,
ill-defined pit on Sternum IV; male with ventral tooth on anterior femur; female with obtuse angle in this position; middle
calcar very small, almost absent; hind calcar of moderate size, broadly rounded.
Punctures on both inner and outer carinae and the elongate pronotum separate this species
from others with a medium-sized postorbital tubercle.
Omoglymmius ( sensu stricto) planiceps new species
Figs. 217, 223
Type material. - HOLOTYPE female, labelled: “NEW GUINEA, Morobe District, Wau,
2-3-X-1969, James E. Tobler” (CAS).
Description. — Length 7.0 mm. Antennal Segments I-X coarsely punctate; Segment XI with few punctures; head
distinctly longer than wide; median lobe short, broad, its apex rounded; frontal space slightly wider than long; its margins
curved; medial angles nearly rectangular, but blunt; well separated; posteriomedial margin oblique; posteriolateral margin
evenly curved; occipital angle distinct; antennal groove short, connected laterally to shallow preorbital impression; very
narrow orbital groove traceable almost to posterior margin of eye; temporal lobe with about 12 punctures, most in form of
row medial to eye; one temporal seta; postorbital tubercle very short but deep, visible though inconspicuous in dorsal view;
temporal lobe markedly flattened in lateral view; eye large, round.
Pronotum short, length/greatest width 1.12; widest posterior to middle; base very slightly narrowed; apex strongly
narrowed; lateral margins nearly straight posteriorly, curved, convergent anteriorly; margin slightly sinuate anterior to
hind angle; marginal groove dilated; in dorsal view, outer carina appearing 0.6 as wide as inner carina at middle; outer
carina convex, directed dorsolaterad, apparently narrower in dorsal view than in dorsolateral view; medial margin of outer
carina shallowly sinuate just anterior to base; outer carina widest at base; inner carina obtusely pointed posteriorly; both
pairs of carinae entirely impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytra moderately long; striae not impressed, represented only by rows of fine, shallow, elongate punctures; base of
Stria IV with longitudinal pollinose scarp; Stria IV without seta; subapical striole with one seta; Stria VII without setae (or
perhaps broken off); metasternum entirely punctate; metasternum, prosternum with faint bluish opalescence; abdominal
Sterna III-V with fine, scattered punctures; female with moderately deep semicircular lateral pit on Sternum IV; female
without ventral tooth on anterior femur; male unknown.
Revision of Omoglymmius Ganglbauer
249
Flattened temporal lobes are distinctive in this species. The dilated marginal groove and
narrow, convex outer carina also separate it from any other species from New Guinea. If the
very short postorbital tubercle were overlooked, this species would trace to O. trepidus in the
key to species from New Guinea. It differs from the latter species in having the temporal lobe
with fewer punctures, the outer carina impunctate and narrow and the pronotum not narrowed
at the base.
Omoglymmius (sensu stricto ) lent us new species
Figs. 218, 224
Type material. - HOLOTYPE male, labelled: “D. N. Guinea, Sattelberg” (MNHB).
Description. — Length 7.2 mm. Antennal Segments 1-X coarsely punctate; Segment XI with one or two coarse
punctures; head 1.3 longer than wide; median lobe broad, its apex obtuse; frontal space much broader than long, nearly
U-shaped, its margins abruptly curved; medial angles obtuse, separated; posteriomedial margin oblique, curved gradually
into posterior margin; latter nearly transverse laterally, bent abruptly into lateral margin; occipital angle absent; width
across temporal lobes greatest just posterior to eyes, margins convergent anteriorly; antennal groove moderately broad, its
lateral end slightly dilated; orbital groove absent; temporal lobe with seven or eight punctures near eye; temporal seta
apparently absent, but possibly rubbed off in the holotype; postorbital tubercles about 0.33 as long as eye, visible in dorsal
view, width across them slightly greater than that across eyes; latter large, round, more protuberant than usual in
subgenus.
Pronotum short, length/greatest width 1.07; widest near middle; base moderately narrowed; apex very markedly
narrowed; lateral margins markedly curved; margin sinuate anterior to hind angle; outer carina 0.6 as wide as inner carina
at middle; medial margin of outer carina sinuate just anterior to base; outer carina widest at base, abruptly narrowed
anterior to base, from there nearly even width, abruptly narrowed near apex; inner carina with base narrowed, rounded;
outer carina with two or three fine punctures; inner carina impunctate; median groove markedly narrowed at middle;
pronotum without setae; prosternum without precoxal carinae.
Elytron with bluish opalescence; striae not impressed, represented by rows of fine, shallow, elongate punctures; base of
Stria IV with oblique pollinose scarp; transverse scarp not pollinose; Stria IV without seta near apex; subapical stride with
one seta; Stria VII with several setae near apex; metasternum with bluish opalescence, entirely punctate; abdominal Sterna
III-V with fine, scattered punctures; male with shallow lateral pits on Sternum IV; male with ventral tooth on anterior
femur; middle calcar acute, small, about 0,25 as long as spur; hind calcar triangular, its apex obtuse, its proxijnaJ margin
straight; female unknown.
The species closest in appearance to this one is O. massa. In addition to the absence of
punctures on the metasternal disc, the latter species differs in the shape of the temporal lobes,
the shorter, more rounded median lobe, and the very broad base on the pronotum. The broad
based pronotum and flattened temporal lobes distinguish O. planiceps from O. lentus.
Omoglymmius (sensu stricto ) capito (Grouvelle)
Figs. 219, 225
Rhysodes capito Grouvelle 1895a: 157-158.
Omoglymmius (sensu stricto ) capito (Grouvelle) Bell and Bell 1978
Type material. - HOLOTYPE male, labelled: “Nouv. Guinee, Dorey, Baie du Geelvink,
Raffray & Maindron 78” (MNHN).
Description. — Length 8.3 mm. Antennal Segments V-X coarsely punctate; Segment XI missing in holotype; head
as broad as long; median lobe lance-shaped, tip obtuse; frontal space broader than long; nearly V-shaped, margins oblique,
shallowly sinuate; medial angles rounded, well separated; posteriomedial margin curved into posteriolateral margin;
occipital angle absent; antennal groove moderately broad, not expanded at lateral end; orbital groove absent; temporal lobe
with three or four coarse punctures between eye and temporal seta, also about 10 fine punctures more anteriorly, medially;
one temporal seta; postorbital tubercles about 0.8 as long as eye, prominent in dorsal view; width across them much greater
than across eyes; postorbital tubercles narrower, more divergent than in O. auratus ; eye large, round.
Pronotum short, length/greatest width 1.09, widest near middle; base slightly narrowed; apex more markedly
narrowed; lateral margin curved, moderately so in basal half, markedly so near apex; margin not sinuate anterior to hind
angle; outer carina slightly narrower than inner one at middle; medial margin of outer carina sinuate anterior to base;
Quaest. Ent., 1982, 18 (1-4)
250
Bell and Bell
outer carina broadest just anterior to base, abruptly narrowed just anterior to it, then slightly broadened to anterior 0.33
of length; apex narrowed; inner carina with base narrowed, rounded; outer carina with 20-30 fine punctures; inner
carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron rather broad, surface not opalescent, but opaque, faintly microsculptured; striae not impressed, represented
by rows of fine, elongate punctures; base of Stria IV with longitudinal pollinose scarp; Stria IV with one seta at apex;
subapical stride without seta; Stria VII with several setae near apex; metasternum completely finely, shallowly
punctate; abdominal Sterna 11I-V with scattered punctures; male with shallow lateral pit on Sternum IV; male with
acute prominent ventral tooth on anterior femur; middle calcar acute, about 0.2 as long as spur; hind calcar triangular,
apex nearly rectangular, proximal margin straight; female unknown.
The prominent postorbital tubercles of this species are comparable to those of O. auratus.
The latter species has postorbital tubercles broader, outer carinae impunctate, the paramedian
grooves much narrower, and disc of the metasternum impunctate.
Range. - Known only from the type locality, on the Vogelkop Peninsula at the west end of
New Guinea. Grouvelle (1903) later cited additional localities. We have not located these
specimens, and do not know whether they really belong to this species.
Omoglymmius (sensu stricto) cheesmanae (Arrow)
Figs. 220, 226
Rhysodes cheesmanae Arrow 1942: 180-181
Omoglymmius (sensu stricto ) cheesmanae (Arrow) Bell and Bell 1978
Type material. - HOLOTYPE male, labelled: “DUTCH N. GUINEA, Cyclops Mtns.,
Sabron, 930 ft., V-1936, L. E. Cheesman, BM 1936-271” (BMNH).
Description. — Length 5. 3-6.0 mm. Antennal Segments I-XI punctate; head distinctly longer than wide; median
lobe oval with two to six coarse punctures, tip obtusely pointed; frontal space wider than long; margins moderately curved;
medial angles acute, contiguous; posteriomedial margin oblique; posteriolateral margin evenly curved; occipital angle
absent; antennal groove narrow; orbital groove rudimentary, ended just posterior to anterior margin of eye; temporal lobe
with 15-26 fine punctures; one temporal seta; postorbit flat, pollinose; suborbital tubercle inconspicuous, about 0.2 as long
as eye; eye very large, round, prominent.
Pronotum moderately elongate, length/greatest width 1.20; widest near middle; base distinctly narrowed; apex
markedly narrowed; lateral margins curved; margin not sinuate anterior to hind angle; outer carina slightly narrower than
inner one at middle; medial margin of outer carina scarcely sinuate anterior to base; outer carina broadest at anterior 0.33,
narrowed to point anteriorly; inner carina with base narrow, attenuate; outer carina with 25-30 fine punctures; inner carina
with about six fine punctures; pronotum without setae; prosternum without precoxal carinae.
Elytron with striae shallowly impressed; strial punctures coarse, shallow; base of Stria IV with longitudinal pollinose
scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex;
metasternum completely punctate; abdominal Sterna 111-V with coarse, scattered punctures, these slightly confluent near
lateral margin; male with shallow lateral pit on Sternum IV; male with ventral tooth on anterior femur; middle calcar
minute, obtuse, almost absent; hind calcar triangular, its apex acute, both margins straight; female unknown.
Among the species which have suborbital tubercles, this one has the smallest tubercle. O.
pulvinatus and O. asetatus have the tubercles conspicuously larger, and have the posterior
margin of the temporal lobe margined with pollinosity, while O. sedlaceki has a much broader
pronotum which is not narrowed at the base.
Range. - North coast of New Guinea at low elevations. In addition to the holotype, we have
Studied the following specimens: one male, labelled “NEW GUINEA (NW), Ifar, Cyclops Mts, 300-500 m„
23-25-VI-1962, J. Sedlacek” (BPBM); one male, labelled “Maffin Bay, Dutch New Guinea, V 1 1-8- 1 944,coll. E. S. Ross”
(CAS).
Omoglymmius (sensu stricto ) asetatus new species
Figs. 221, 227
Type material. - HOLOTYPE male, labelled: “Madang, N. Guinea, Nov. 196 ,
Hornabrook coll.” (NMNZ).
Revision of Omoglymmius Ganglbauer
251
Description. — Length 6.0 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head distinctly
longer than wide; median lobe impunctate, oval, tip obtuse; frontal space twice as wide as long; nearly U-shaped, its
margins markedly curved; medial angles obtuse, nearly contiguous; posteriomedial margin oblique, posteriolateral margin
evenly curved; occipital angle absent; posterior margin of temporal lobe bordered by pollinosity; antennal groove rather
broad; orbital groove short, ended opposite anterior 0.3 of eye; temporal lobe with 22-27 very fine punctures; temporal seta
absent; postorbit convex, pollinose; suborbital tubercle both longer and deeper than those of O. cheesmanae , length 0.4 of
that of eye; eye very large, round, prominent.
Pronotum moderately long, length/greatest width 1.20; widest slightly anterior to middle; base slightly narrowed; apex
markedly narrowed; lateral margins curved; margin sinuate anterior to hind angle; outer carina about 0.6 as wide as inner
Carina at middle; outer carina with medial margin distinctly sinuate just anterior to base; outer carina with width just
anterior to base and width at anterior 0.33 equal; inner carina long, oval, its base narrowly rounded; outer carina with
12-13 fine punctures; inner carina with three to four fine punctures; pronotum without setae; prosternum without precoxal
carinae.
Elytron with striae shallowly impressed; strial punctures coarse, shallow; base of Stria IV with longitudinal pollinose
scarp; Stria IV with one seta near apex; subapical stride with one seta; Stria VII with several setae near apex;
metasternum completely punctate; abdominal Sterna III-V coarsely punctate, punctures in irregular transverse row on
each sternum; punctures markedly coalescent near lateral margins, in form of indistinct transverse sulci; that of Sternum
IV dilated to form indistinct lateral pit; with small obtuse ventral tooth on anterior femur; middle calcar almost absent;
hind calcar small, triangular, its apex obtuse; female unknown.
This species is similar to O. cheesmanae , but differs in having the posterior face of the
temporal lobe pollinose, the suborbital tubercle larger, the temporal seta absent, and the lateral
margin of the pronotum sinuate just anterior to the hind angle.
Omoglymmius (sensu stricto) biroi new species
Figs. 228, 234
Type material. - HOLOTYPE female, labelled: “N. GUINEA Biro’ 1899. Sattelberg,
Huon Gulf’ (MNHN). The specimen is labelled as a type of R. pulvinatus , but is not from the
type locality of the latter species, Hatam, and is not mentioned in the original description. It is
not conspecific with Grouvelle’s type series. PARATYPE female, labelled: “D. N. Guinea,
Sattelberg”, collector and data not indicated (MNHB).
Description. — Length 6.0-7. 7 mm. Antennal Segments I-X coarsely punctate; Segment XI impunctate; head
distinctly longer than wide; median lobe impunctate, suboval, tip pollinose, obtusely rounded frontal space broader than
long; margins curved at middle of length; medial angles obtuse, slightly lobate; moderately separated; posteriomedial
margin oblique; posteriolateral margin rather suddenly bent near temporal seta, posterior portion nearly transverse;
posterior margin of temporal lobe margined with pollinosity; occipital angle obtuse; antennal groove broad; orbital groove
broad, ending near middle of eye; temporal lobe with 15-20 fine punctures; one temporal seta; postorbit flat, pollinose;
suborbital tubercle small, about 0.3 as long as eye; eye large, round, prominent.
Pronotum moderately elongate; length/greatest width 1.21; widest slightly anterior to middle; base slightly narrowed;
apex moderately narrowed; lateral margins weakly curved; margins shallowly sinuate anterior to hind angle; outer carina
about 0.6 as wide as inner carina at middle; medial margin of outer carina curved, not sinuate anterior to base; outer
carina widest at anterior 0.25, narrowed to apex, narrowed near base, but base widened; inner carina with base narrowly
truncate; outer carina with 30-35 fine punctures; inner carina with 10-16 fine punctures; pronotum without setae;
prosternum without precoxal carinae.
Elytron with striae shallowly impressed, intervals flat; strial punctures fine, shallow; base of Stria IV with pollinose
longitudinal scarp; transverse basal scarp pollinose from Stria IV to Stria I; Stria IV with one seta near apex; subapical
stride with one seta; Stria VII with several setae near apex; metasternum with very fine punctures along lateral margins
and in midline, otherwise impunctate; abdominal Sterna III-V with coarse punctures in form of irregular transverse row on
each sternum; female with shallow semicircular lateral pit on Sternum IV; female with obtuse ventral tooth on anterior
femur; male unknown.
Among those species with suborbital tubercles, this one is recognized by the elongate,
subquadrate form of the pronotum, and nearly truncate tip of the median lobe.
Quaest. Ent., 1982, 18 (1-4)
252
Bell and Bell
Omoglymmius (sensu stricto ) pulvinatus (Grouvelle)
Figs. 229, 235
Rhysodes pulvinatus Grouvelle 1903: 115-116.
Omoglymmius (sensu stricto ) pulvinatus (Grouvelle) Bell and Bell 1978
Type material. - LECTOTYPE (here designated) male, labelled: “Hatam, VII, Beccari,
1875, typus, 6881” (GEN). It bears a label “ Rhysodes pulvinatus ” in Grouvelle’s handwriting.
PARALECTOTYPE: one male, same label as lectotype (GEN). A female, with the same
labels as the lectotype (GEN), is not this species, but is the species described herein as O.
denticulatus new species.
Description. — Length 6.3 mm. Antennal Segments I-IX coarsely punctate; Segments X, XI more finely punctate,
head distinctly longer than wide; median lobe impunctate, suboval, tip obtusely rounded; frontal space broader than long;
margins curved at middle of length; medial angles obtuse, moderately separated; posteriomedial margin oblique;
posteriolateral margin evenly curved, margined with pollinosity; occipital angle obtuse; antennal groove broad; orbital
groove absent; temporal lobe with 11-13 fine punctures; one temporal seta; postorbit flat, nearly glabrous; suborbital
tubercle small, about 0.3 as long as eye; eye large, round, less prominent than in O. biroi.
Pronotum rather short; length/greatest width 1.13; widest near middle; base slightly narrowed, apex markedly
narrowed; lateral margins rather strongly curved; margin sinuate anterior to hind angle; outer carina about 0.4 as wide as
inner carina at middle; medial margin of outer carina curved, scarcely sinuate anterior to base; marginal groove strongly
dilated; inner carina narrowed to base; outer carina with 10-15 moderately coarse punctures; inner carina impunctate;
pronotum without setae; prosternum without precoxal carinae.
Elytron with striae shallowly impressed, intervals flat; strial punctures fine, shallow; base of Stria IV with pollinose
longitudinal scarp; transverse basal scarp pollinose from Stria IV to Stria I; Stria IV with one seta near apex; subapical
stride with one seta; Stria VII with several setae near apex; metasternum punctate near margins and in midline, otherwise
impunctate; abdominal Sterna 1II-V with numerous scattered punctures; with small lateral pit on Sternum IV; anterior
femur with ventral tooth; middle calcar minute; hind calcar triangular, obtuse; female unknown.
This species differs from others with suborbital tubercles in having the outer carina very
narrow, and the marginal groove dilated. O. biroi appears to be the closest related species, but
it differs from the latter in having the abdominal punctures numerous and scattered, and in
having the pronotum shorter, with more rounded lateral margins.
Omoglymmius (sensu stricto ) sedlaceki new species
Figs. 230-233, 236, 237
Type material. - HOLOTYPE male, labelled: “NEW GUINEA:(NE), Wau, Morobe
Distr. 1200 m., 14-IX-61, coll. J. Sedlacek & native collector” (BPBM). PARATYPES one
male, two females, same data as holotype. The following paratypes all have the same locality
data as holotype but with the following dates, elevations and collectors: two males, 1400 m.,
27-VIII-1961, J. Sedlacek; two males, 1300 m., 27-VII-1961 and 22-XII-1961, J. & J. H.
Sedlacek; one female, 1050-1100 m., 1 5-XII- 1 96 1 , G. Monteith, J. Sedlacek; one female, 1650
m., 5-XII- 1961, J. Sedlacek; one female, 1700-1800 m., 17-XI-1961, J. Sedlacek; one male,
1200 m., 1-1-1962, J. & M. Sedlacek; one female, 1200-1700 m., 1 0- VIII- 1 962, J. Sedlacek;
one male, 1200-1300 m., 6-IV-1963, J. Sedlacek (ALL BPBM); one female, New Guinea
(NE): Wau, Mt. Kaindi, 1500-1800 m., 12-14- VI-1963, coll. P. Shanahan (BPBM); one
female, New Guinea (NE), Wau, Kunai Creek, 1500 m., 28-30-V-1963, coll. P. Shanahan
(BPBM).
Description. — Length 6. 8-8. 4 mm. Outer antennal segments finely punctate; head distinctly longer than wide;
median lobe moderately to very broad, its tip obtusely rounded to slightly pointed; frontal space nearly as long as broad; its
margin shallowly curved to nearly oblique; medial angles nearly rectangular, well separated; posteriomedial margin deeply
emarginate; posteriolateral margin curved; occipital angle distinct, in some specimens lobate; posterior margin between
occipital angle and posteriolateral margin oblique in some specimens, emarginate in others; antennal groove moderately
Revision of Omoglymmius Ganglbauer
253
broad; orbital groove absent; temporal lobe with nine to 13 fine punctures, mostly near lateral margin; one temporal
seta; postorbit flat, glabrous; suborbital tubercle inconspicuous, 0.3 of length of head; eye large, round, prominent.
Pronotum rather short, length/greatest width about 1.10; widest near middle, subquadrate; base slightly narrowed,
extreme apex markedly narrower; lateral margin feebly curved except near apex, where markedly curved; margin feeble
to markedly sinuate anterior to hind angles; width of outer carina subequal to inner one at middle; outer carina sinuate
on medial margin just anterior to base; outer carina with two equal points of greatest width, one just anterior to base,
another anterior to middle; inner carina with base narrow, attenuate, pointed; outer carina varying in punctuation; inner
carina impunctate; pronotum without setae; prosternum without precoxal carinae.
Elytron with striae shallowly impressed; strial punctures shallow, rather coarse; base of Stria IV with longitudinal
pollinose scarp; transverse basal scarp pollinose between bases of Stria 1-1 V; Stria IV with one seta near apex; subapical
stride with one seta; Stria VII with several setae near apex; metasternum with punctures near margins, its disc
impunctate; punctures of abdominal Sterna II1-V coarse, sparse, tending to form one transverse row on each Sternum,
interrupted at midline; semicircular lateral pit present on Sternum IV in both sexes, deeper in female; male with
prominent ventral tooth on anterior femur; female without such tooth; middle calcar scarcely developed; hind calcar
triangular, obtuse, its proximal margin nearly straight, its distal margin emarginate between tip of calcar and spur.
In this species, the punctures of the metasternum are limited to margins, the postorbit is flat
and glabrous, and the outer carina is as wide just anterior to the base as it is at the anterior
third.
Variations. - We provisionally include in this species a series of allopatric samples which
differ from one another in shape of the pronotum, median lobe, and posterior margin of the
temporal lobe, as well as in the number of punctures on the pronotum. Possibly these forms are
subspecifically or specifically distinct, although it is also possible that there are clinal variations
in these characters.
We believe that we can distinguish five allopatric forms. Form A, from Wau and vicinity,
includes the type series. The posterior margin of the temporal lobe is sinuate near the medial
angle but oblique lateral to it, or with sinuation barely suggested. The outer carina has about 20
punctures in its lateral half. Form B, from Kassem, has the temporal lobe as in the preceding,
but lacks punctures in the outer carina. Form C, from the Eastern Highlands Province
(Kamira, Okapa, Doulo Pass, Mount Michael), has the posterior margin of the temporal lobe
twice sinuate, with a lobate occipital angle. The outer carina has only two or three punctures.
Form D, from Mount Otto, differs from all others in having a very broad median lobe with a
distinct angle at its tip. It has the posterior margin of the temporal lobe twice sinuate, and the
outer carina with many punctures. Form E, from Sepalakambang, has the median lobe
narrower than in any other form. The posterior margin of the temporal lobe is once sinuate, and
the pronotol carinae are impunctate.
Range. - Highlands of Northeast New Guinea. In addition to the type series, we
provisionally include the following specimens: (FORM B) one male, one female, Kassem Pass, E. Highlands,
24-2-74, coll. R. Hornabrook (NMNZ); (FORM C) two females, Doulo Pass, Asato-Chimbu Divide, 16-1-72 & 13-3-72,
coll. R. Hornabrook (NMNZ; one female, Mt. Michael, Lufa, Feb. -71, coll. R. Hornabrook; one male, one female, Okapa,
Kamira, 2-9-1964 & 24-6-1965, coll. R. Jornabrook (NMNZ); (FORM D) one female, Mt. Otto, 2200 m„ June 24-1955,
coll. J. L. Gressitt (BPBM); (FORM E) one female, Sepalakambang, Salawaket Range, 1920 m., IX- 1 5-56, E. J. Ford, jr.
(BPBM).
SUBSTITUTIONS FOR PREOCCUPIED GENERIC NAMES
Two of the generic names proposed by us in Part I. (Bell and Bell 1978) are preeoccupied.
Tangaroa Bell and Bell 1978 is preoccupied by Tangaroa Lehtinnen 1967. We propose the
substitute name Tangarona New Name. The type species and only member of the genus is
Tangarona pensus (Broun) NEW COMBINATION.
Kupea Bell and Bell 1978 is preoccupied by Kupea Philpott 1930. We propose the substitute
name Kupeus New Name. The type species and only member of the genus is Kupeus arcuatus
Quaest. Ent., 1982, 18 (1-4)
254
Bell and Bell
(Chevrolat) NEW COMBINATION.
We thank Dr. Norman Platnick for pointing out the first homonymy, and Dr. P. Basilewsky
for pointing out the second one.
ACKNOWLEDGEMENTS
We wish to thank the numerous curators and collectors whose aid has made this work
possible. Many of them were mentioned in Parts I and II. We are especially indebted to Dr.
Roberto Poggi for detailed drawings and descriptions of some of the types in the Genoa
Museum, and for the loan of valuable specimens from the latter museum. Dr. F. Hieke of
Humboldt University (Berlin) also made valuable types available to us. We are grateful to Dr.
O. Kryzhanovskij of the Academy of Sciences, Leningrad, for sending us samples of species
from the U.S.S.R., and for the loan of specimens from other regions. We also acknowledge our
debt to Mrs. Joyce Murray for her care in the long and difficult task of typing this manuscript.
SUPPLEMENTARY REFERENCES3
Bell, R. T. and J. R. Bell. 1981. Coleoptera: Rhysodidae. Insects of Micronesia 15(2): 51-67.
Nakane, T. 1978. New or little-known Coleoptera from Japan and its adjacent regions, XXIX
(In English), Reports of the Faculty of Science, Kagoshima University (Earth
Sciences and Biology) 11: 129-134.
NOTE ADDED IN PROOF
Because two specimens designated as holotypes were badly damaged in the mail,
replacements are herewith designated:
Omoglymmius modicus: HOLOTYPE male, labelled “6-4-62-1455, Savo, Solomon
Islands” (BMNH). This is listed on p. 233 as a paratype.
Omoglymmius regius: HOLOTYPE female, labelled “Isabela, Tatumba. 24-8-1963, P.
Greenslade, 9587” (BMNH). This is listed on p. 235 as a paratype.
3This supplements the list of references found in Part I and Part II.
Revision of Omoglymmius Ganglbauer
255
INDEX TO NAMES OF TAXA
(Synonyms in italics)
FAMILY GROUP TAXA
Buprestidae, 206
Cerambycidae, 206
Clavicornia, 206
Omoglymmiina, 131, 164
Rhysodidae, 221
Rhysodini, 162, 232, 233, 235, 238
GENERA AND SUBGENERA
Boreoglymmius new subgenus, 132, 133,
140
Caeconavitia new subgenus, 132, 166, 167,
168
Carinoglymmius new subgenus, 132, 176,
177
Dhysores Grouvelle, 164
Hemiglymmius Bell and Bell, 132, 135,
137, 138, 140, 141, 161, 164, 177
Indoglymmius new subgenus, 132, 157,
161, 167
Kupea Bell and Bell, 253
Kupea Philpott, 253
Kupeus New Name, 253
Laminoglymmius new subgenus, 132, 133,
146, 156, 157, 162, 164, 167
Navitia Bell and Bell, 132, 164, 167
Nitiglymmius Bell and Bell, 132, 166, 167,
168, 180
Omoglymmius (sensu stricto), 140, 167,
169, 180, 207,208,211,224, 225
Omoglymmius Ganglbauer, 130, 131, 132,
137, 146, 157, 162, 164, 243
Orthoglymmius Bell and Bell, 132, 133,
161, 164, 169, 177, 180
Pyxiglymmius Bell and Bell, 130, 131,
132, 143, 146, 154, 155, 157
Rhyzodiastes Fairmaire, 177, 227
Tangaroa Bell and Bell, 253
Tangaroa Lehtinnen, 253
Tangarona New Name, 253
SPECIES AND SUBSPECIES
actae new species, Omoglymmius, 132,
146, 156, 157, 160, 161
africanus (Grouvelle), Omoglymmius,
133, 135
africanus Grouvelle, Rhysodes , 132
alticola (Grouvelle), Omoglymmius, 170,
173, 175
alticola Grouvelle, Rhysodes , 173
americanus (Castelnau), Omoglymmius,
133, 141, 143, 144, 206
americanus Castelnau, Rhysodes , 140,
144
americanus Reitter, Rhysodes , 194
amplus new species, Omoglymmius, 183,
189, 211, 212
aratus Chevrolat, Rhysodes , 143, 194
aratus Newman, Rhysodes , 144
arcuatus (Chevrolat), Kupeus, 254
armatus (Arrow), Omoglymmius, 147,
150, 153, 154
armatus (Arrow), Pyxiglymmius , 154
armatus Arrow, Rhysodes , 1 54
asetatus new species, Omoglymmius, 186,
193,250
aterrimus (Chevrolat), Omoglymmius,
153
aterrimus Chevrolat, Rhysodes , 153, 154
auratus new species, Omoglymmius, 187,
193,246, 247,250
batchianus (Arrow), Omoglymmius, 184,
190, 223,225,226, 228
batchianus Arrow, Rhysodes , 223
bicarinatus new species, Omoglymmius,
181, 189, 230
biroi new species, Omoglymmius, 186,
193,251,252
borneensis (Grouvelle), Omoglymmius,
133,214
bouchardi new species, Omoglymmius,
184, 189,212,216, 221,222
bucculatus (Arrow), Omoglymmius, 181,
190, 229, 237
Quaest. Ent., 1982, 18 (1-4)
256
Bell and Bell
bucculatus Arrow, Rhysodes, 229
caelatus Bell and Bell, Omoglymmius,
183, 194, 232
capito (Grouvelle), Omoglymmius, 187,
194, 249
capito Grouvelle, Rhysodes , 249
carinatus (Grouvelle), Omoglymmius,
177, 179
carinatus Grouvelle, Rhysodes , 176, 179
cavea new species, Omoglymmius, 184,
192, 242, 243
cavifrons (Grouvelle), Omoglymmius, 170,
172, 174, 175
cavifrons Grouvelle, Rhysodes , 174
cheesmanae (Arrow), Omoglymmius, 186,
193, 250, 251
cheesmanae Arrow, Rhysodes , 250
classicus new species, Omoglymmius, 183,
192, 232, 238,239
coelebs new species, Omoglymmius, 185,
189, 209, 215
consors new species, Omoglymmius, 185,
189,214,216, 221
continuus new species, Omoglymmius,
180, 190, 226
coomani (Arrow), Omoglymmius, 170,
174, 175, 176, 179
coomani Arrow, Rhysodes , 176
crassicornis new species, Omoglymmius,
183, 188,211
crassiusculus (Lewis), Omoglymmius, 146,
150, 151
crassiusculus (Lewis), Pyxiglymmius, 151
crassiusculus Lewis, Rhysodes , 143, 151
crenatus (Grouvelle), Omoglymmius, 170,
175, 176
crenatus Grouvelle, Rhysodes , 175
cristatus new species, Omoglymmius, 146,
150, 152, 153
data new species, Omoglymmius, 185, 188,
209,217
denticulatus new species, Omoglymmius,
187, 193,246, 247, 252
duplex new species, Omoglymmius, 180,
184, 189, 209,219, 220
ephemeris new species, Omoglymmius,
185, 192, 244
evasus new species, Omoglymmius, 185,
189,212,213,214,215
exaratus Erichson, Rhysodes , 194
exaratus Serville, Rhysodes , 144
feae (Grouvelle), Omoglymmius, 170, 172,
174, 176
feae Grouvelle, Rhysodes , 172
follis new species, Omoglymmius, 186,
193,245,246
fraudulentus new species, Omoglymmius,
185, 189,212,214,215,217,222
fringillus new species, Omoglymmius, 182,
193,224, 227, 240, 241
fulgens Bell and Bell, Omoglymmius, 168
germaini (Grouvelle), Omoglymmius, 133,
136, 137, 138
germaini Grouvelle, Rhysodes , 137
germari (Ganglbauer), Omoglymmius,
141, 180, 187, 194, 206
germari Ganglbauer, Rhysodes , 131, 180,
194
gorgo new species, Omoglymmius, 155,
158, 160, 163
gracilicornis (Grouvelle), Omoglymmius,
185, 192, 230, 244
gracilicornis Grouvelle, Rhysodes , 244,
245
greensladei Bell and Bell, Omoglymmius,
168
gurneyi new species, Omoglymmius, 182,
191,227,232, 235,236
hamatus (Leconte), Omoglymmius, 133,
141, 143, 145
hamatus Leconte, Rhysodes , 143
hemipunctatus new species,
Omoglymmius, 133, 136, 137, 138
hesperus new species, Omoglymmius, 147,
150, 153, 154
hexagonus (Grouvelle), Omoglymmius,
177, 178, 179
hexagonus Grouvelle, Rhysodes , 178
hiekei new species, Omoglymmius, 185,
189, 209,218
Quaest. Ent., 1982, 18 (1-4)
Revision of Omoglymmius Ganglbauer
257
hornabrooki Bell and Bell, Omoglymmius,
168
humeralis (Grouvelle), Omoglymmius,
184, 190, 224
humeralis Grouvelle, Rhysodes , 224
ichthyocephalus Lea, Rhysodes , 180
impletus Bell and Bell, Omoglymmius,
186, 194, 231
imugani new species, Omoglymmius, 184,
188, 209,210,211,218
inaequalis new species, Omoglymmius,
132, 157, 158, 160, 161, 167, 177, 178
ineditus (Dajoz), Omoglymmius, 135, 136,
138, 139, 140
ineditus Dajoz, Rhysodes , 138
inermis new species, Omoglymmius, 135,
139, 140
insularis (Grouvelle), Omoglymmius, 146,
157, 160, 162
insularis (Grouvelle), Pyxiglymmius , 162
insularis Grouvelle, Rhysodes , 156, 162
intrusus (Grouvelle), Omoglymmius, 164,
166
intrusus Grouvelle, Rhysodes , 164, 166
iridescens new species, Omoglymmius,
187, 193,245,247
javanicus (Grouvelle), Omoglymmius,
133, 136, 137, 138
krikkeni new species, Omoglymmius, 146,
150, 153, 156
laticeps Bell, Omoglymmius, 181, 188, 207
lederi (Lewis), Omoglymmius, 146, 150,
152, 157
lederi (Lewis), Pyxiglymmius , 152
lederi Lewis, Rhysodes , 152
lentus new species, Omoglymmius, 187,
194, 249
lewisi (Nakane), Omoglymmius, 133, 141,
143, 151,206
lewisi Nakane, Rhysodes , 141
lindrothi new species, Omoglymmius, 183,
192, 231,232, 233
lineatus (Grouvelle), Omoglymmius, 167
lineatus Grouvelle, Rhysodes , 167
longiceps (Grouvelle), Omoglymmius,
170, 173, 174, 176
longiceps Grouvelle, Rhysodes , 173
lustrans Bell and Bell, Omoglymmius, 168
malabaricus (Arrow), Omoglymmius, 181,
187, 207,208
malabaricus Arrow, Rhysodes , 208
malaicus (Arrow), Omoglymmius, 185,
187,214,216
malaicus Arrow, Rhysodes , 215
manni new species, Omoglymmius, 183,
191,232, 234, 235
massa new species, Omoglymmius, 187,
193,246, 247, 249
microtis new species, Omoglymmius, 170,
172, 175
modicus new species, Omoglymmius, 183,
191,232, 233,234
modiglianii new species, Omoglymmius,
183, 189,212
morditus new species, Omoglymmius, 183,
190, 228
mycteroides new species, Omoglymmius,
185, 191,237
nasalis new species, Omoglymmius, 184,
190, 228,237
nemoralis new species, Omoglymmius,
185, 189,213,214,215,222
nicobarensis (Grouvelle), Omoglymmius,
133, 177
nicobarensis Grouvelle, Rhysodes , 161,
111
oberthueri (Grouvelle), Omoglymmius,
133, 157, 160, 162
oberthueri Grouvelle, Rhysodes , 162
occultus new species, Omoglymmius, 133,
136, 138
oceanicus Bell and Bell, Omoglymmius,
184, 194, 231
offafinus Bell and Bell, Omoglymmius,
168
opticus new species, Omoglymmius, 184,
190, 225,226
oroensis new species, Omoglymmius, 182,
193,224, 240, 241
patens new species, Omoglymmius, 186,
192, 231,242, 243
258
Bell and Bell
pectoralis new species, Omoglymmius,
185, 189,216,218,219
pensus (Broun), Tangarona, 253
philippensis (Chevrolat), Omoglymmius,
184, 188, 209, 210, 211, 212, 213, 215,
216,217, 220, 231
philippensis (Chevrolat), Rhysodes, 209
philippensis Chevrolat, Rhyzodes, 209
philippinensis (Chevrolat), Rhysodes , 209
pilosus (Grouvelle), Omoglymmius, 146,
150, 153, 155, 156, 161
pilosus (Grouvelle), Pyxiglymmius, 155
pilosus Grouvelle, Rhysodes , 155
planata Fab., Uleiota, 206
planiceps new species, Omoglymmius, 187,
194, 231,248,249
politus new species, Omoglymmius, 181,
184, 188, 189, 210, 218, 230
princeps new species, Omoglymmius, 183,
191, 232, 235, 236, 239
pulvinatus (Grouvelle), Omoglymmius,
186, 193,250, 252
pulvinatus Grouvelle, Rhysodes , 251, 252
puncticornis new species, Omoglymmius,
182, 192, 241
quadraticollis (Arrow), Omoglymmius,
186, 190, 222, 225, 226
quadruplex new species, Omoglymmius,
180, 185, 189,219, 220
regius new species, Omoglymmius, 183,
192, 232, 233,235
renutus new species, Omoglymmius, 184,
191,232, 236, 239
repetitus new species, Omoglymmius, 186,
190, 222, 223
rimatus new species, Omoglymmius, 133,
136, 139
rojasi Chevrolat, Clinidium, 137
rugosus (Grouvelle), Omoglymmius, 146,
147, 157, 160, 163
rugosus (Grouvelle), Pyxiglymmius , 160
rugosus Grouvelle, Rhysodes , 160
rusticus new species , Omoglymmius, 192
rusticus new species, Omoglymmius, 183,
232, 233,234
sakuraii (Nakane), Omoglymmius, 181,
188,207
sakuraii Nakane, Rhysodes , 207
scopulinus new species, Omoglymmius,
182, 191, 232, 235, 237
sectatus new species, Omoglymmius, 185,
192, 242, 243,244
sedlaceki new species, Omoglymmius, 186,
193,250, 252
semioculatus new species, Omoglymmius,
168, 169
semperi new species, Omoglymmius, 182,
188,217,218
silvatica L., Fagus, 206
solitarius (Arrow), Omoglymmius, 132,
178, 186, 187,208
solitarius Arrow, Rhysodes , 208
strabus (Newman), Omoglymmius, 146,
147, 150, 151, 152, 153, 154
strabus (Newman), Pyxiglymmius , 153
strabus Newman, Rhysodes , 146, 153
stylatus new species, Omoglymmius, 164,
165, 166'
subcaviceps (Grouvelle), Omoglymmius,
146, 150, 151
subcaviceps (Grouvelle), Pyxiglymmius ,
150
subcaviceps Grouvelle, Rhysodes , 1 50
sulcicollis (Lewis), Omoglymmius, 170
sulcicollis Germar, Dechomus, 206
sulcicollis Lewis, Rhysodes , 169, 170
summissus new species, Omoglymmius,
181, 188,212,216,217
sus new species, Omoglymmius, 187, 194,
248
tabulatus new species, Omoglymmius,
186, 191, 238
thoracicus new species, Omoglymmius,
185, 189,216,219
toxopei Bell and Bell, Omoglymmius, 168
trepidus new species, Omoglymmius, 184,
192, 242, 243,249
trisinuatus new species, Omoglymmius,
158, 160, 163, 164
vadosus new species, Omoglymmius, 181,
190, 227,228
Quaest. Ent., 1982, 18 (1-4)
Revision of Omoglymmius Ganglbauer
259
vicinus (Grouvelle), Omoglymmius, 182,
192, 239, 240, 241,245
vicinus Grouvelle, Rhysodes, 239
viduus new species, Omoglymmius, 182,
190, 225, 226, 241
wittmeri new species, Omoglymmius, 182,
190, 227
zimmermani Bell and Bell, Omoglymmius,
166, 167, 168
CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY OF THE NORTH
AMERICAN SPECIES OF SIPHONA MEIGEN (DIPTERA: TACHINIDAE)
J.E. O’Hara
Department of Entomology
University of Alberta
Edmonton, Alberta, Canada
T6G 2E3
Quaestiones Entomologicae
18:261-380 1982
ABSTRACT
Members of the cosmopolitan genus Siphona are small tachinid flies with a long geniculate
proboscis. Their known hosts are Tipula spp. (Tipulidae) and Microlepidoptera ( especially
Noctuidae ).
Siphonopsis Townsend, with the following three species, is removed from synonymy with
Siphona: type-species Siphonopsis plusiae (Coquillett) ( California ), Siphonopsis brasiliensis
Townsend ( Brazil ) and Siphonopsis conata Reinhard (California). Two species, Crocuta
alticola Mesnil (Burma) and Crocuta crassulata Mesnil (Burma), are placed in Ceranthia s.
lat. Siphona diluta Wulp (Mexico) is transferred to Clausicella. Crocuta malaisei Mesnil
(Burma) and Crocuta pellex Mesnil (Burma) are placed in Actia. Phantasiosiphona Townsend
is synonymized with Siphona, with type-species P. tropica Townsend (Mexico) and P. kuscheli
Cortes (Juan Fernandez Islands).
Twenty-one Siphona species of North America (America north of Panama-Colombia
border) are recognized and described, including 11 new species. Three species described from
the Palearctic Region represent new New World records: S. cristata ( Fabr .), S. hokkaidensis
Mesnil and S. maculata Staeger. A species described from western United States, S. lurida
Reinhard, is a new Palearctic record (from Japan). Two keys are provided for identification of
adult specimens, one to species of America north of Mexico and the other to Middle American
species. Male and female head profiles and male genitalia are illustrated for all species.
Known distributions of North American species are mapped.
Six species groups are recognized in North America on the basis of a cladistic analysis of
15 characters of adults. A cladogram is presented for each species group, showing
hypothetical phylogenetic relationships among North American species. The S. maculata
group is inferred to be the most primitive of the North American species groups. Available
data are insufficient to determine phylogenetic relationships among the other species groups.
Analysis of Palearctic Siphona species illustrates the general applicability of North American
species groups to species in other regions.
The following six species groups with their included species are: (1) S. maculata group: S.
floridensis n. sp. (type-locality Orlando, FA), S. intrudens (Curran), S. lurida Reinhard, S.
maculata Staeger and S. pacifica n. sp. (type-locality Dartford, WA), (2) S. cristata group: S.
cristata (Fabr.), S. lutea (Townsend), S. multifaria n. sp. (type-locality mi. 87, Dempster Hwy.,
Y.T.), S. oligomyia n. sp. (type-locality Keremeos, B.C.) and S. pisinnia n. sp. (type-locality
21km. n. Silver City, NM), (3) S. geniculata group: S. hokkaidensis Mesnil, S. geniculata (De
Geer) and S. medialis n. sp. (type-locality Lockeport, N.S.), (4) S. futilis group: S. brunnea n.
262
O’Hara
sp. ( type-locality Zontehuitz, Chiapas, Mex.j, S. futilis Wulp and S. illinoiensis Townsend, (5)
S. tropica group: S. akidnomyia n. sp. ( type-locality Zontehuitz, Chiapas, Mex.), S. longissima
n. sp. ( type-locality lOmi. ne. San Cristobal de las Casas, Chiapas, Mex.), S. rizaba n. sp.
( type-locality Orizaba, Veracruz, Mex.), and S. tropica (Townsend), and (6) S. macronyx
group: S. macronyx n. sp. ( type-locality Pullman, WA).
S. tenuis Curran is synonymized with S. lutea (Townsend), S. ceres (Curran) with S. futilis
Wulp and S. silvarum Herting with S. hokkaidensis Mesnil. Lectotypes are designated for S.
futilis Wulp, S. illinoiensis Townsend and S. lutea (Townsend).
Three general distribution patterns are evident within species groups of Siphona: (1)
America north of Mexico and the Palearctic Region - S. maculata, S. cristata and S.
geniculata groups, (2) America north of Mexico and Middle America - S. futilis group, and (3)
Middle America and Afrotropical Region - S. tropica group. Within North America,
distribution patterns of Siphona species are classed as follows: (1) transcontinental - S.
cristata, S. hokkaidensis, S. intrudens, S. lutea, S. maculata, S. medialis and S. multifaria, (2)
western - S. geniculata ( introduced ), S. lurida, S. macronyx, S. oligomyia, S. pacifica and S.
pisinnia, and (3) Mexican - S. akidnomyia, S. brunnea, S. futilis, S. longissima, S. rizaba and S.
tropica.
A historical zoogeographic analysis seeks an explanation to these distribution patterns.
The patterns are considered in the context of current paleoclimatic and paleogeographic
hypotheses to arrive at several conclusions about the historical zoogeography of Siphona
species: (1) Siphona is hypothesized to have originated in Africa between the Late Cretaceous
and Late Eocene, (2) a S. tropica group taxon was first to enter the New World, following a
northern paratropical route between Eurasia and North America, no later than the Late
Eocene, (3) the first taxa of the S. cristata, S. geniculata and S. maculata groups migrated to
North America through Beringia after the Tertiary climatic optimum, probably no earlier
than the Miocene, (4) exchange between Old and New Worlds continued within the S. cristata,
S. geniculata and S. maculata groups into the Pleistocene, and (5) three species pairs represent
Pleistocene vicariant events: S. maculata and S. floridensis, S. lutea and S. oligomyia, and S.
futilis and S. illinoiensis.
RESUME
Le genre Siphona comprend de petites mouches tachinides caracterisees par un long proboscis genicule. Les holes connus
incluent des Tipula spp. (Tipulidae) et des Microlepidopteres (principalement des Noctuidae).
Siphonopsis Townsend, contenant les trois especes suivantes, n’est plus considere comme synonyme de Siphona:
Siphonopsis plusiae ( Coquillett ) (de Calif ornie), F espece-type du genre, Siphonopsis brasiliensis Townsend ( du Bresil), et
Siphonopsis conata Reinhard (de Californie). Les especes Crocuta alticola Mesnil et Crocuta crassulata Mesnil (toutes
deux de Birmanie ) sont placees dans le genre Ceranthia s. lat. Siphona diluta Wulp (du Mexique) est transferee dans le
genre Clausicella. Crocuta malaisei Mesnil et Crocuta pellex Mesnil (toutes deux de Birmanie ) sont placees dans le genre
Actia. Phantasiosiphona Townsend, incluant P. tropica Townsend (du Mexique ), V espece-type, et P. kuscheli Cortes (des
lies Juan Fernandez ), est considere comme synonyme de Siphona.
Vingt-et-une especes de Siphona, dont II nouvelles, sont reconnues et decrites pour FAmerique du Nord (au nord de
la frontiere panamo-colombienne). Trois especes connues precedemment de la region palearctique represented de
nouvelles additions a la fauna nearctique: S. cristata (Fabr.), S. hokkaidensis Mesnil et S. maculata Staeger. S. lurida
Reinhard, connue de Vouest des Etats-Unis, est nouvellement mentionnee dans la region palearctique (Japon). Deux cles
permettent Tidentification des adultes, une pour les especes se trouvant au nord du Mexique, et V autre pour les especes
d’Amerique Centrale. Le profit de la tete des males et des femelles, de me me que Torgane genital des males sont illustres
pour chaque espece. Des cartes illustrant la repartition geographique connue des especes nord-americaines sont fournies.
Une analyse cladistique portant sur 15 caracteres des adultes permet de reconnoitre six groupes d’especes
nord-americaines. Pour chaque groupe, les relations phyletiques hypothetiques entre les especes constituantes sont
representees par un cladogramme. Le groupe de S. maculata represente probablement le group d’especes
North American species of Siphona Meigen
263
nord-americaines le plus primitif. Les donnees disponibles sont insuffisantes pour etablir les relations phyletiques
entres les autres groupes d’especes. L' analyse des especes palearctiques de Siphona illustre I'applicabilite generate des
regroupements bases sur les especes nord-americaines aux especes des autres regions.
Les six groupes, ainsi que leurs especes constituantes, sont: (1) le groupe de S. maculata, comprenant S. floridensis,
nouvelle esp'ece (localite du type: Orlando, Floride), S. intrudens (Curran), S. lurida Reinhard, S. maculata Staeger, et
S. pacifica, nouvelle espece ( localite du type: Dartford, Washington); (2) le groupe de S. cristata, incluant S. cristata
(Fabr.), S. lutea (Townsend), S. multifaria, nouvelle espece (localite du type: mille 87, autoroute Dempster, Yukon), S.
oligomyia, nouvelle espece (localite du type: Keremeos, Colombie-Britannique), et S. pisinnia, nouvelle espece (localite
du type: 21 km. au nord de Silver City, Nouveau- Mexique); (3) le groupe de S. geniculata, comprenant S. hokkaidensis
Mesnil, S. geniculata (De Geer), et S. medialis, nouvelle espece (localite du type: Lockeport, Nouvelle-Ecosse); (4) le
groupe de S. futilis, comprenant S. brunnea, nouvelle espece (localite du type: Zontehuitz, Chiapas, Mexique), S. futilis
Wulp, et S. illinoiensis Townsend; (5) le groupe de S. tropica, incluant S. akidnomyia, nouvelle espece (localite du type:
Zontehuitz, Chiapas, Mexique), S. longissima, nouvelle espece (localite du type: 10 mi. au nord-est de San Cristobal
de las Casas, Chiapas, Mexique), S. rizaba, nouvelle espece (localite du type: Orizaba, Veracruz, Mexique), et S.
tropica (Townsend); (6) le groupe de S. macronyx, incluant S. macronyx, nouvelle espece (localite du type: Pullman,
Washington).
S. tenuis Curran est consideree comme synonyme de S. lutea (Townsend), S. ceres (Curran), comme synonyme de
S. futilis Wulp, et S. silvarum Herting, comme synonyme de S. hokkaidensis Mesnil. Des lectotypes sont designes pour
S. futilis Wulp, S. illinoiensis Townsend et S. lutea (Townsend).
Les groupes d’especes de Siphona presentent trois patrons de distribution: (I) des groupes repartis en Amerique du
Nord au nord du Mexique et dans la region palearctique, c’est-a-dire ceux de S. maculata, S. cristata et S. geniculata;
(2) le groupe de S. futilis, reparti en Amerique du Nord et en Amerique Centrale; (3) le groupe de S. tropica, reparti en
Amerique Centrale et dans la region afrotropicale. En Amerique du Nord, les patrons de distribution des especes de
Siphona se repartissent comme suit: (I) especes transcontinentales, incluant S. cristata, S. hokkaidensis, S. intrudens, S.
lutea, S. maculata, S. medialis et S. multifaria; (2) especes de I'Ouest, avec S. geniculata (introduite), S. lurida, S.
macronyx, S. oligomyia, S. pacifica et S. pisinnia; (3) especes mexicaines, avec S. akidnomyia, S. brunnea, S. futilis, S.
longissima, S. rizaba et S. tropica.
Une analyse historique de la biogeographie tente d’expliquer ces patrons de distribution. En examinant ceux-ci a
la lumiere des hypotheses cour antes sur les paleoclimats et la paleogeographie, Thistoire biogeographique des especes
de Siphona semble avoir inclu les evenements suivants: (I) Siphona supposement eut son origine en Afrique entre le
Cretace superieur et I’Eocene superieur; (2) un taxon du groupe de S. tropica atteignit le premier le Nouveau Monde
en suivant un itineraire septentrional a climat paratropical entre VEurasie et I'Amerique du Nord, et ce, avant
I’Eocene superieur; (3) les premiers taxons des groupes de S. cristata, S. geniculata et S. maculata atteignirent
I’Amerique du Nord via Beringia apres la periode climatique la plus chaude du Tertiaire, soit probablement au plus
tot durant le Miocene; (4) les groupes de S. cristata, S. geniculata et S. maculata continuerent d’echanger des elements
entre I’Ancien et le Nouveau Monde jusque durant le Pleistocene; (5) finalement, trois paires d’especes, soit S.
maculata et S. floridensis, S. lutea et S. oligomyia, et S. futilis et S. illinoiensis, evoluerent a la suite de phenomenes de
vicariance qui eurent lieu durant le Pleistocene.
Table of Contents
Introduction 264
Materials and Methods 265
Materials 265
Methods . . . 267
Genus Siphona Meigen 272
World list of Siphona species 277
Keys to adults of North American Siphona species 281
Key to adults of Siphona species of Canada and United States 281
Key to adults of Siphona species of Middle America 284
The S. maculata group 286
The S. cristata group 292
The S. geniculata group 299
The S. futilis group 305
The S. tropica group 309
The S. macronyx group 312
Quaest. Ent., 1982, 18 (1-4)
264
O’Hara
Phylogenetic interpretation 314
Zoogeographic considerations 327
Acknowledgements 336
Literature cited 337
Figures 345
Index 377
INTRODUCTION
Specimens of Siphona species are small (3-6mm long) tachinids of cosmopolitan
distribution. Like most other tachinids, they spend their three larval instars as internal
parasitoids of other insects. Chiefly parasitic on larval Microlepidoptera in general and the
Noctuidae in particular (if the meagre published data are any indication), a few species
parasitize their distant dipteran relatives, Tipula species (Tipulidae).
As parasitoids of other insects, tachinids are highly regarded for their role in the natural
environment and their potential for use as control agents against pest insects. When the
introduced pest Tipula paludosa Meigen began damaging lawns and meadows in parts of
Newfoundland, Cape Breton Island and the lower Fraser Valley of British Columbia, a
Palearctic parasitoid was sought for its control. Siphona geniculata (De Geer) was
recommended for introduction and became part of a biological control program on both coasts
(Wilkinson, 1971). Though little success has been achieved in limiting T. paludosa populations
by S. geniculata (Wilkinson, per. comm.), the program illustrates the desirability of an
alternative to long term chemical control of a pest species.
Whether New World Siphona species are economically important, or have the potential to
be so, is unknown. Until now even taxonomic aspects have been neglected, so a framework has
not been developed upon which to build a more comprehensive knowledge of the group. This
paper represents a step in that direction by providing a revision to North American1 Siphona
species.
The history of North American Siphona classification has been primarily one of individual
species descriptions under varied generic names. Curran (1932) tried to improve the situation
through publication of a key to North American Siphona species, but evidently based the key
mainly on descriptions. As a result, an assemblage of species belonging to three genera were
keyed according to untrustworthy and insignificant characteristics. Furthermore, two described
Siphona species were apparently overlooked, for they were not treated in the key.
Judged from a historical perspective, the list of eight Siphona species compiled by Sabrosky
and Arnaud (1965) represented a significant contribution. It was from this source that my first
impression of the number of species and constitution of the genus in the Nearctic Region was
gleaned.
I was soon to discover the inadequate state of Siphona classification - a result of lack of
attention, not expertise. Entwined with species problems was the more basic question of how to
define the genus. In answering this question my revision was extended to include Middle
America, for it was chiefly through study of Siphonopsis species and allied genera from
America north of Panama-Colombia border
North American species of Siphona Meigen
265
southwestern United States and Mexico that I recognized the fundamental difference between
those taxa and Siphona.
A major portion of this paper is concerned with classificatory aspects of North American
Siphona species. Also considered are phylogenetic and zoogeographic aspects, with reference to
species from other regions. Due to the excellent taxonomic contributions of Dr. L.P. Mesnil
(1960, 1961, 1964 and 1965), it has been possible to study interrelationships between North
American (particularly Nearctic) and Palearctic Siphona species.
Species groups are recognized for the first time in Siphona. Based on cladistic relationships,
it is hoped that these North American species groups will have general applicability.
MATERIALS AND METHODS
Materials
This study was based on examination of approximately 4000 adult specimens of Siphona
and related siphonines, borrowed from the following institutions and individuals. Abbreviations
are those used in the text to indicate deposition of specimens.
Institutional loans. -
AMNH Department of Entomology, American Museum of Natural History, Central
Park West at 79th Street, New York, NY 10024; P. Wygodzinsky.
BMNH Department of Entomology, British Museum (Natural History), Cromwell
Road, London, England SW7 5BD; J.P. Dear.
CAES Connecticut Agricultural Experiment Station, 123 Huntington Street, Box
1 106, New Haven, CT 06504; K.A. Welch.
CAS Department of Entomology, California Academy of Sciences, Golden Gate
Park, San Francisco, CA 94118; P.H. Arnaud, Jr.
CFA Coleccion de la Facultad de Agronomia, Universidad de Chile, Casilla 1004,
Santiago, Chile; R. Cortes.
CNC Biosystematics Research Institute, Central Experimental Farm, K.W. Neatby
Building, Ottawa, Ontario K1A 0C6; D.M. Wood.
CSU Department of Zoology and Entomology, Colorado State University, Fort
Collins, CO 80523; H.E. Evans.
CUI Department of Entomology, Cornell University, Comstock Hall, Ithaca, NY
14853; L.L. Pechuman.
FSCA Florida State Collection of Arthropods, Division of Plant Industry, Florida
Department of Agriculture, Gainesville, FL 32602; H.V. Weems, Jr.
INHS Illinois Institute of Natural Resources, Section of Faunistic Surveys and Insect
Identification, Illinois Natural History Survey, 172 Natural Resources
Building, Urbana, IL 61801; D.W. Webb.
KSU Department of Entomology, Kansas State University, Waters Hall, Manhattan,
KS 66506; H.D. Blocker.
KUR Biological Laboratory, College of General Education, Kyushu University,
Ropponmatsu, Fukuoka 810, Japan; H. Shima.
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138;
M.K. Thayer.
MSU Department of Entomology, Michigan State University, East Lansing, MI
48824; R.L. Fischer.
Quaest. Ent., 1982, 18 (1-4)
266
O’Hara
MSUB Department of Biology, Montana State University, Bozeman, MT 59717; S.
Rose.
NCSU Department of Entomology, Entomology Museum, North Carolina State
University, Box 5215, Raleigh, NC 27650; C.S. Parron.
NDSU Department of Entomology, North Dakota State University, Fargo, ND 58102;
E.U. Balsbaugh, Jr.
NSM Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia, B3H 3A6;
B. Wright.
OKSU Department of Entomology, Oklahoma State University, 501 Life Sciences
West, Stillwater, OK 74074; W.A. Drew.
OSU Department of Entomology, Oregon State University, Corvallis, OR 97331;
M.D. Schwartz.
OUCO Department of Entomology, Ohio State University, 103 Botany and Zoology
Building, 1735 Neil Avenue, Columbus, OH 43210; C.A. Triplehorn.
PSU Frost Entomological Museum, Department of Entomology, Pennsylvania State
University, University Park, PA 16802; S.W. Frost.
ROM Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6; G.B.
Wiggins.
SMNS Staatl. Museum fur Naturkunde, Stuttgart, D-714 Ludwigsburg, Arsenalplatz
3, West Germany; B. Herting.
UAF Department of Entomology, University of Arkansas, Fayetteville, AR 72701; R.
Chenowith.
UASM Department of Entomology, Strickland Museum, University of Alberta,
Edmonton, Alberta T6G 2E3; G.E. Ball.
UCD Department of Entomology, University of California, Davis, CA 95616; R.O.
Schuster.
UCR Department of Entomology, University of California, Riverside, CA 92521; S.I.
Frommer.
UGA Department of Entomology, University of Georgia, Athens, GA 30602; C.L.
Smith.
UGG Department of Environmental Biology, University of Guelph, Guelph, Ontario
NIG 2W1; S. Marshall.
UKL Department of Entomology, Snow Entomology Museum, University of Kansas,
Lawrence, KS 66045; C.D. Michener and G.W. Byers.
UMW Department of Entomology, University of Manitoba, Winnipeg, Manitoba R3T
2N2; T.D. Galloway.
USNM Systematic Entomology Laboratory, SEA, U.S. Department of Agriculture,
United States National Museum, Washington, DC 20560; C.W. Sabrosky.
USP Museu de Zoologia, Universidad de Sao Paulo, Avenida Nazare, 481, Caixa
Postal 7172, 04263, Sao Paulo, SP, Brazil; J.H. Guimaraes.
WSUP Department of Entomology, James Entomological Collection, Washington State
University, Pullman, WA 99164; W.J. Turner.
ZMUC Zoologisk Museum, Universitetsparken 15, Dk 2100, Copenhagen, Denmark; S.
Andersen.
Loans from private collections. -
DMW D.M. Wood, c/o CNC.
North American species of Siphona Meigen
267
WLD W.L. Downes, c/o P.L. Fischer, MSU.
JEOH designates specimens in my personal collection.
Methods
Criteria for ranking of taxa. - My concept of a bisexual species is essentially that of Mayr
(1969: 26), whereby reproductive isolation of populations is the basic criterion upon which the
concept is based. Seldom is it practical, or even possible, for a systematist to field test this
hypothesis with phena under revision. Rather, reproductive isolation is inferred from other
data.
I used two criteria to aid in recognition of species. One was distributional data and the other
phenetic divergence.
Widely sympatric phena were assumed to represent species, for otherwise interbreeding
would presumably have led to deterioration of phenetic difference (barring ecological or
temporal barriers). For phenetically similar species like S. cristata and S. multifaria , and S.
maculata and S. intrudens, this criterion was especially useful.
The degree of phenetic divergence exhibited among sympatric species provided an indication
of interspecific variation within Siphona. Along with studies of intraspecific variation, it
became possible to distinguish which allopatric phena were sufficiently unique phenetically as
to represent species.
The subspecific category (as discussed by Mayr, 1969: 187-197) has not been used in this
work. It was considered for the polymorphic populations of S. hokkaidensis, but rejected as
inappropriate for reasons given under “Variation” of that species. Geographical distributions
and structural characteristics of other phena were such that the subspecific category was not a
viable alternative.
Species sharing one or more derived states and representing apparently monophyletic
(Ashlock’s (1971) holophyletic) lineages were arranged under the informal category of species
groups.
Decisions at the generic level do not figure prominently in this work, so criteria governing
generic decisions are only briefly mentioned here.
Monophyletic genera are preferred, though paraphyletic genera are considered acceptable.
Polyphyletic genera are considered unacceptable.
The following are regarded as important generic considerations: (1) distinctiveness from
other taxa, (2) degree of intrageneric morphological difference, and (3) number of taxa
included (see Mayr, 1969, for discussion). These three parameters are believed to influence the
acceptability of paraphyletic taxa in a classification. Cladistic classifications such as those
expounded by Eldridge and Cracraft (1980) and Nelson and Platnick (1981) are not used here.
Specimen examination. - Pinned, dried specimens were examined with a Wild M5
stereoscopic microscope with standard light source, at magnifications up to 50X. Genitalia were
studied with a Leitz SM-LUX compound microscope at magnifications up to 400X.
Measurements. - Body length was measured dorsally at SOX through a Wild M5
stereoscopic microscope with an ocular micrometer (100 divisions) calibrated to a slide
micrometer (100 divisions to the millimeter). Body length was taken as length from pedicel to
scutellum plus abdominal length from attachment to thorax to apex of tergum 5. This
measurement eliminated differences due to position of antennae and abdomen relative to rest of
body.
Quaest. Ent., 1982, 18 (1-4)
268
O’Hara
Head measurements of eye and head height and length of first flagellomere were taken in
profile, as shown in Fig. 1. Proboscis length was taken as combined length of prementum and
labella. Prementum was measured basally from ventral edge of sclerotization to most distal
point. Labellar length was measured from dorsal base of labella to labellar tip. Head
measurements were taken at 50X with an ocular micrometer, and expressed in text and
Hubbs-Hubbs diagrams in ratios.
Illustrations are provided with scale bars, which were obtained by projecting an image of a
slide micrometer through a camera lucida (see “Illustrations” below) onto the drawing.
Hubbs-Hubbs diagrams. - Hubbs-Hubbs diagrams representing intraspecific variation in
height of compound eye, lengths of first flagellomere and proboscis (Figs. 13-15), and ratio of
length of first flagellomere: eye height (Fig. 16) were prepared. Male specimens were chosen
for representation in Hubbs-Hubbs diagrams because their eye and first flagellomere display
greater interspecific difference than females. Proboscis length is not sexually dimorphic so
measurements from female specimens were included in Fig. 15 for species with few male
specimens.
Hubbs-Hubbs diagrams are arranged as follows. Species names are along the left margin,
abbreviated to their first four letters and listed in order of proboscis length, from shortest to
longest. S. hokkaidensis is divided into samples “a” and “b”, as discussed under variation of
that species. Number of specimens measured is given along the right margin. The appropriate
ratio is along the X-axis.
Illustrations. - All illustrations were made with the aid of a camera lucida, as detailed
below, and later inked to mylar drafting film for reproduction.
Rough head drawings were made with the aid of a camera lucida attached to a Wild M5
stereoscopic microscope, at a magnification of 50X. Rough drawings were redrawn before
inking so parts could be reoriented and details altered to best represent the species, and to show
characteristics of each species in comparable positions.
For male head figures, setae of the ocellar triangle and postcranium were not drawn. To
simplify the task of drawing male and female heads of all species, setae were drawn on male
head illustrations only.
With large variation in eye height and first flagellomere and proboscis lengths within
species, head figures should be consulted in conjunction with the Hubbs-Hubbs diagrams to
avoid reliance on the figured characteristics of each species.
The rostrum and proboscis are shown in a variety of positions. The rostrum is externally
membranous so its shape has no taxonomic value. The prementum of the proboscis is shown as
a straight cylindrical structure, for in life it is inflexible. This is not true of the labella, the distal
half of which is highly manoeuverable in live specimens. This flexibility is evident in its twisted
nature in a few drawings, but generally the labella is shown fairly straight to keep drawings
standardized. Proboscises as drawn are meant to convey relative differences in length, not shape
or structure.
Genitalia were temporarily mounted on slides in glycerine jelly2 for illustrative purposes.
Once drawings were completed genitalia were washed free of jelly in hot water and returned to
glycerine for storage.
2formula of Pantin (1969).
North American species of Siphona Meigen
269
Genitalia should not be left for extended periods in glycerine jelly or their complete
extraction becomes difficult; ie. the jelly loses its ability to mix freely with hot water and
remains coated to the specimen.
A Leitz SM-LUX compound microscope with drawing tube was used at a magnification of
100X for rough drawings of all genitalic figures. Rough drawings were redrawn and reoriented
for better representation of parts, as mentioned for heads.
I did not detect setal differences in male and female genitalia useful for distinguishing
species, so setae were not illustrated.
All sclerites of female Siphona genitalia from sternum 5 to cerci are shown in Fig. 12. Note
that the dorso-longitudinally interrupted tergum of segment 8 is all that remains of terga 6 to
10. Sternum 9 is either lost, or fused with sternum 10.
Figures of male genitalia show only those structures associated with segments 9 to 11. In
addition to vestiture, the ejaculatory apodeme and paramere (postgonite) are excluded from
illustrations.
I do not recommend comparison of figured male genitalia on the basis of structures
illustrated but not discussed in the text, for differences may be misleading. For instance,
hypandrial and aedeagal apodemes and epandrium are considerably varied intraspecifically, so
their figured appearance cannot be used for identification purposes. Genitalia were drawn as
shown to represent taxonomically useful characters in their proper relative positions as parts of
a functional complex.
Thoracic and abdominal dorsa (Figs. 2, 7-8) were drawn as described for heads. Hairs,
which sparsely cover both regions, were not drawn.
Acropods (Figs. 4-6) and the wing (Fig. 3) were drawn with the aid of a drawing tube
attached to a SM-LUX compound microscope. Acropods were suspended in glycerine jelly and
drawn at a microscope magnification of 400X. Short hairs on claws, pulvilli and tarsomere 5
are not shown.
The wing (Fig. 3) was mounted in Canada balsam and drawn at a microscope magnification
of 100X. Costal setulae and wing hairs were not illustrated.
Terms. - A variety of terms apply to the same structures of adult flies. This developed partly
from gradual introduction of new terms by successive generations of dipterists, and partly by
establishment of terms unique to particular groups.
With the recent publication of the “Manual of Nearctic Diptera, Volume 1”, an attempt
was made to standardize terms for structures of adult Diptera (McAlpine, 1981). The depth
and detail that pervade this work prompted me to abandon the more familiar terms for
tachinids of Crosskey (1973, 1976a) and others in favour of it. Certain of McAlpine’s terms are
controversial, and for that reason unacceptable to all, yet acceptance of his work in principle
would lead to establishment of a unified system of names for structures.
Most structures to which I refer in the text are labelled in Figs. 1-12. Where possible,
abbreviations correspond to those in the “Manual of Nearctic Diptera”. The Manual should be
consulted for structures not figured. In a few instances I included a familiar term in parenthesis
after a Manual term if the feature was not figured.
One of my two deviations from terms in the Manual involves a few thoracic setae. I retained
use of “humeral” and “presutural” setae. The former was termed “postpronotal” in the
Manual, but this seems incongruous without a replacement name for the posthumeral seta,
which was not mentioned in the Manual. I see no reason to incorporate the presutural seta into
the supra-alar row, so I disregarded this change.
Quaest. Ent., 1982, 18 (1-4)
270
O’Hara
Certain descriptive terms in the text require explanation to avoid misinterpretation. First of
these is colouration, which is difficult to describe qualitatively. This is compounded in Siphona,
as in most calyptrate flies, by the effect pruinosity has on ground colour; where pruinosity
(pollinosity) refers to “a nap-like covering to any part of the body formed by closely aggregated
ultramicroscopic pubescence” (Crosskey, 1973: 19), and ground colour simply refers to the hue
of sclerites.
Both pruinosity and ground colour contribute to Siphona colouration, but separately
described impart little meaning. Therefore I describe “surface colour”, which I define as the
hue or hues perceived as a result of light reflected from pruinosity as seen against ground
colour. This is essentially synonymous with “colour” of most authors, but here permits
distinction between ground and surface colour. In the text, where neither surface nor ground
colour is specified, surface colour is implied.
Wing colour is varied specifically, but appears very faint from dorsal aspect. To obtain wing
colour used in descriptions, I viewed wings from an acute angle to increase colour density and
permit differentiation of yellow, brown and cinereous hues.
Less ambiguous than colour but requiring explanation are terms used in describing
macrotrichia. Here I diverge from the Manual again. I classify macrotrichia as hairs, setulae
and setae (bristles). Hairs are macrotrichia of approximately equal diameter along their
lengths. Setulae and setae are macrotrichia with thick bases and slender tips, inserted in
conspicuous sockets. Setulae and setae are qualitatively separated on basis of size; setulae being
small setae.
The Manual equated macrotrichia with setae, reserving bristles, setulae and hairs as
subordinate categories. This is untenable on two counts. Firstly, “seta” is Latin for “bristle”, so
it is incorrect to consider the latter subordinate to the former. Similarly, a setula is by definition
a small seta, so a setula should in no instances be larger than a seta. (In the sense of the
Manual, a seta can refer to a hair, which is smaller than a setula.)
I make use of the terms “weak” and “strong” setae in descriptions. This avoids repeated
reference to the length and thickness aspects of setal size, both of which are implied by these
terms.
Leg setae are named for their positions relative to body axis when a leg is fully extended to
the side. Abbreviations used in description of the genus are as follows:
a - anterior
ad - antero-dorsal
av - antero-ventral
d - dorsal
p - posterior
pd - postero-dorsal
pv - postero-ventral
v - ventral
Notes on descriptive format. - Genus and species descriptions are presented in a uniform
format, with same style and headings. Differences concern degree of coverage. The aim of the
genus description was to describe in detail the attributes of Siphona , without differentiation
between character states shared with other genera and states unique to Siphona.
Siphona species descriptions do not repeat non-varied attributes of the genus. In most
instances a character state or set of states is described in full for each species unless it is shared
with a closely related species. For a few characters in which one state is shared by most species.
North American species of Siphona Meigen
271
that state is described as average, and average is described in the genus description.
Certain characteristics are more useful than others for identification purposes, so a diagnosis
is presented prior to each description under the heading “Recognition”, summarizing those
attributes by which the species is most reliably recognized. Also included under this heading is
a comparison of the described species with other species with which it may be confused.
Label data for holotypes and allotypes of new species, and examined type material of
described species are presented in a uniform manner. Labels are cited in full, following closely
the style of Arnaud (1979). Under this system labels are listed from the top down, with data
from each label enclosed in quotation marks. Original spelling and punctuation are preserved,
and label lines are delimited by a slash mark(/). Square brackets signify information not
included on label. Deposition of each specimen is given in parenthesis.
Paratype data of new species were treated less stringently than the above. A standardized
format was adopted to increase readability and reduce likelihood of errors. Countries and
provinces (or states) are arranged alphabetically; counties, cities, etc. are not. Counties (if
given on label) are followed by locality, date of collection, collector and number of males
and/or females examined. Specimen repositories are given at the end of each paratype list.
Date of collection is given in order of day, month, year, with month in Roman numerals and
year abbreviated to the last two digits (though written in full if earlier than 1900). Specimens
collected in the same locality on different dates by the same collector are cited together, with
commas between dates of collection. Inclusive dates of collection are indicated by dashes. For
example, 5,7-9. VII.81 signifies that specimens were collected on the 5th of July and again
between the 7th and 9th of July, 1981.
I have not presented data about examined, non-type specimens of described species,
considering their value insignificant in light of the distribution maps and chart of adult seasonal
occurrence. For the sake of posterity a list of these specimens was prepared and deposited in the
Archives of the University of Alberta. The list format follows that used for paratypes of new
species.
Distribution maps. - Localities represented by specimens I personally examined are
indicated by dots on the maps. I excluded published locality records because of their
unreliability as to species collected.
Only New World records of species described herein are mapped. Holarctic species are
indicated as such in figure captions and under heading “Distribution” following descriptions.
Genitalia dissections. - Dissections were required to permit examination of Siphona male
and female genitalia. The technique described below refers to dissection of male genitalia from
dried specimens, though it works as well with female and fresh material, with obvious minor
changes. This method differs slightly from those reported elsewhere (eg. Beneway, 1963;
Wilder, 1979), and was developed to its present form during the course of this study.
Dissection of male genitalia involved removal and slight clearing of the abdomen, extraction
and clearing of the pregenital ring (sternum 5 and segments 6-8) and genitalia (segments 9 to
1 1), separation of ring and genitalia, and replacement of abdomen on specimen. Details of this
procedure follow.
Removal of an abdomen from a dried specimen was relatively easy because it naturally
tends to break at its connection to the thorax when gently pushed from below. Gentle prodding
from above and from side to side helped detach the abdomen cleanly and evenly.
The abdomen was placed in a solution of about 10% NaOH until the cuticle became flexible
(about 10 minutes, but varied from specimen to specimen). Air trapped in the abdominal cavity
Quaest. Ent., 1982, 18 (1-4)
272
O’Hara
was released during this time (by gentle squeezing) so that membranes around the genitalia
would soften from within as well as from without. This was done carefully and after some
flexibility had been attained, for otherwise preabdominal sclerites (segments 1-5) pulled apart.
The abdomen was then transferred to 20% acetic acid to neutralize the base.
Dissection of the genitalia was done in the acetic acid solution. The bent and blunted tip of
an insect pin, attached to a handle, was used to hold the abdomen ventral side up against the
dish bottom. In this position only a few minutes were required to slice around syntergosternum
7 + 8 and sternum 5, and pull them and enclosed genitalia free. This operation worked well with
a very sharp tungsten steel needle. If tissues were not sufficiently cleared, the preabdominal
terga were easily torn; if overcleared there was danger of the terga separating.
The genitalia were returned to the NaOH solution until attached membranous tissue
changed from translucent to transparent. This step would take from 5 to 20 minutes, depending
on the specimen.
The preabdomen was taken through 10 minute washes of 70% and 95% ethyl alcohol to
toluene. After 20 minutes it was removed, air dried (a matter of seconds) and affixed with
shellac glue to the thorax of the specimen from whence it came. Slight shrivelling of the
preabdomen usually occurred, but I have been unable to devise a procedure that eliminates this.
The toluene step (for which similar solvents can be substituted) yielded better results than the
traditional method of alcohol steps only.
Sufficiently cleared genitalia were transferred through 5-10 minute washes of 20% acetic
acid, 70% and 95% alcohol to glycerine.
The high viscosity of glycerine facilitated separation of the epandrium from
syntergosternum 7 + 8. This accomplished, the genitalia were easily removed from the ring-like
pregenital segments.
During the course of this study more than 400 male and about 25 female genitalia of
Siphona and related siphonines were dissected. Since study of male genitalia figured
prominently in development of my Siphona species concepts, they needed to be readily
accessible. To suit this end small vials 1cm in diameter were cut to a height of 7mm and stored
in specially constructed trays measuring about 6. 5x6. 5x1. 5cm (outside dimensions). Each tray
was fitted with a lid and held 20 vials in 4 rows. To the pin of each dissected specimen a
numbered label was attached, and a corresponding number placed with the genitalia in the vial
of glycerine. Genitalia were not placed in microvials and stored with their respective specimens
until the study was completed.
GENUS SIPHONA MEIGEN
Aspects of life history
Siphona immatures. - Larvae of Siphona species live as internal parasitoids of certain other
larval insects. Adult females do not produce many eggs (50-100 in those for which this
information is known), which are retained in an elongate, coiled uterus until they have
transformed into active larvae {S. [l]cristata, Roubaud, 1906 and Pantel, 1910; undetermined
species, Townsend, 1934; S. multifaria and S. maculata , personal observation). They are then
deposited onto hosts. Larvae bore through the host integument and feed within the haemocoel,
meanwhile procuring air via a posterior connection to a host trachial tube (Roubaud, 1906;
Pantel, 1910). When fully mature, Siphona larvae abandon their hosts to seek pupation sites in
the ground.
North American species of Siphona Meigen
273
Hosts. - Hosts of Siphona species are inadequately known. S. geniculata has been recorded
from Tipula paludosa Meigen and T. oleracea L. in Europe (Rennie and Sutherland, 1920).
Also recorded from tipulid larvae are S. hokkaidensis ( T . irrorata Meigen, Mesnil, 1965) and
S. cristata ( T . maxima Poda [as T. gigantea Schrank], Roubaud, 1906). In addition to the
tipulid host, S. cristata has been recorded from ca. 15 noctuid species (see Herting, 1960).
However, S. cristata has been confused with S. flavifrons Staeger (and possibly others), so all
records are suspect.
Siphona maculata has been reared from larvae of Agrotis obelisea and A. candelisequa
(Herting, 1960). Also recorded from noctuids are the European species S. collini and S.
confusa (Mesnil, 1965), the Mexican species S.futilis (as Bucentes ceres (Curran, 1932)), and
S. pseudomaculata Blanchard from South America (Blanchard, 1963).
A parasite-host list for European Siphona species is given in Andersen (in press “a”).
Adult seasonal occurrence. - A chart of adult Siphona seasonal occurrence is presented in
Fig. 17. Most species span three or more months, though I suspect this is in most instances an
artifact of supraposition of data from the entire geographic range of each species, rather than
an indicator of long adult life or overlapping generations. Indeed, populations near Edmonton
with which I am familiar have one generation per year, and adults are seen for only about one
month (S. cristata , S. maculata and S. multifaria) . S. geniculata , the only species for which
generation time has been reported, is bivoltine (Rennie and Sutherland, 1920) or trivoltine
(Andersen, in press “a”).
Fig. 17 indicates trends; that is, which species are likely to be found in early spring, the fall,
etc. As such the chart doubles as a check of identifications, although its value in this respect is
limited for species with inadequate data.
Reclassification
Siphona is remarkably distinct for a genus with more than 80 described species. The long
geniculate proboscis is generally a reliable generic characteristic that permits Siphona
specimens to be identified as such with relative ease. However, possession of a Siphona- like
proboscis has led to inclusion of a few species in Siphona that belong elsewhere. In this section I
discuss removal from Siphona of eight species that superficially resemble Siphona but are more
closely related to other taxa3, and I synonymize one genus with Siphona.
In 1890 Wulp described Siphona diluta and S. futilis. The syntype series of both species
were examined, and S. futilis is retained as a valid Siphona species. S. diluta Wulp is only
distantly related to Siphona , belonging to Clausicella Rondani.
I remove the genus Siphonopsis Townsend from Siphona , with type-species S. plusiae
(Coq.), and two other species, S. brasiliensis Tnsd. and S. conata Rnh. (both originally
described under Siphonopsis but later transferred to Siphona by Sabrosky and Arnaud (1965)
and Guimaraes (1971)). The proboscises of members of Siphonopsis are slightly elongate, with
labella folded back against, but shorter than, prementum, and shorter than eye height. The
labella bear pseudotracheae to their base, unlike the Siphona condition wherein the basal half
of the labella is fused into a cylindrical sheath.
Excluded from consideration are Siphona species not examined during this study; see world list
below for species examined.
Quaest. Ent., 1982, 18 (1-4)
The male genitalia of Siphonopsis species are similar to Siphona except the aedeagus is long
and thin and the antero-lateral arms of sternum 5 are hooked inward. A very few Siphona
species have similar male fifth sterna ( S . oligomyia n. sp. (Fig. 11), S. cuthbertsoni Curran,
and perhaps others I did not examine), but these are almost certainly apotypic and developed
independently of Siphonopsis.
Recognition of Siphonopsis is expedient. The three described species are closely related, but
the generic description does not easily accommodate numerous Mexican and Neotropical
species that are yet undescribed. The problem lies not with separation of Siphonopsis and
Siphona , but in elucidation of the relationships among Siphonopsis, Pseudosiphona Tnsd.,
Ceranthia R.D., Aphantorhapha Tnsd. and Asiphona Mesnil. Especially in the New World
this group is inadequately known and in need of revision. Further study may reveal that
Siphonopsis and perhaps others are best regarded as subgenera of Ceranthia.
Two species described from Burma, Crocuta malaisei Mesnil and Crocuta pellex Mesnil,
belong to a group of Actia species whose adults possess a long geniculate proboscis (including
A. siphonosoma Malloch, A. jocularis Mesnil, A. triseta Mesnil and perhaps others). In this
group the length and shape of the proboscis of adults is amazingly like that in Siphona ,
representing an excellent example of parallel evolution. In other features these Actia species are
characteristic of that genus.
Mesnil recognized his error and briefly mentioned C. malaisei and C. pellex as Actia species
in his description of A. jocularis (1957: 47), and again in his treatment of Palearctic Actia
(1963: 813).
Crosskey examined the types of both C. malaisei and C. pellex and placed both species back
into Siphona (1976a: 214). I have examined a male and female specimen of C. malaisei and I
agree with Mesnil’s placement of this species in Actia. From the description of C. pellex and
Mesnil’s remarks concerning it, I am confident it, too, belongs in Actia.
Two other species described from Burma, Crocuta alticola Mesnil and Crocuta crassulata
Mesnil, are placed in Ceranthia s. lat. Adults of both species have prementum and labella
elongate, as in Siphona species, but the labella bear pseudotracheae along their lengths, as in
Siphonopsis adults. The male genitalia are distinctly Ceranthia-Wke, with broad surstyli
(slender in Siphona species) and a single short posteriorly directed seta on each gonopod (long
in other examined Ceranthia spp., absent from all examined Siphona spp.).
I refer to Ceranthia in the wide sense because the genus has traditionally been comprised
only of adults with an average-sized proboscis and very reduced (apotypic) palpus (ie.
Ceranthia s. str.). The palpi of C. alticola and C. crassulata adults are average-sized, but I
consider the male terminalia with its complex structures more suitable for generic decisions. I
include Asiphona in Ceranthia s. lat., for I am unaware of any synapotypies that unite
Asiphona species. Asiphona is apparently comprised of species that are phylogenetically close
to, but do not belong in, Ceranthia s. str., (cf. Andersen, in press “b”).
To Siphona I add one genus, Phantasiosiphona Tnsd., with type-species P. tropica Tnsd.
and one other species, P. kuscheli Cortes. Given the spectrum of specific attributes within
Siphona, P. tropica is not extraordinary, and does not rank even subgeneric status. P. kuscheli
also conforms to the generic concept of Siphona.
In summary, Siphonopsis is removed from synonymy with Siphona , along with species
Siphonopsis plusiae (Coq.) (1895: 125) (the type-species), S. brasiliensis Tnsd. (1929: 374)
and S. conata Rnh. (1959: 162). New combinations for other species removed from Siphona
are as follows:
North American species of Siphona Meigen
275
Siphona alticola (Mesnil), 1953: 1 10 ( Crocuta (Siphona)), Burma = Ceranthia alticola.
Siphona crassulata (Mesnil), 1953: 112 ( Crocuta (Siphona)), Burma = Ceranthia crassulata.
Siphona diluta Wulp, 1890: 126 (Siphona), Mexico = Clausicella diluta.
Siphona malaisei (Mesnil), 1953: 1 10 (Crocuta (Siphona)), Burma = Actia malaisei.
Siphona pellex (Mesnil), 1953: 1 1 1 (Crocuta (Siphona)), Burma = Actia pellex.
Generic synonymy of Siphona Meigen
A complete list of references to each of the following names was not attempted. Rather, only
major catalogues and literature with keys or descriptions pertaining to New World Siphona
species are cited. Published state records are largely inaccurate and are not included.
Crocuta Meigen, 1800: 39. Type-species, Musca geniculata De Geer (Coquillett, 1910: 528). Suppressed by I.C.Z.N.,
1963: 339 (Opinion 678).- Townsend, 1919: 584.
Siphona Meigen, 1803: 281. Type-species, Musca geniculata De Geer, 1776, by designation of I.C.Z.N., 1974: 157
(Opinion 1008) (see Sabrosky, 1971, for discussion of the history of Siphona nomenclature).- Wulp, 1890: 1 25. —
Townsend, 1891: 368.- Coquillett, 1897: 75.- Aldrich, 1905: 444.- Curran, 1933: 10.— Curran, 1934: 455.- Aldrich,
1934: 108.- Townsend, 1936: 150.- Townsend, 1940: 292.- Reinhard, 1943: 20.- Sabrosky and Arnaud, 1965:
1063.- Cole, 1969: 519, 570.- Cortes and Hichins, 1969: 57.- Cortes and Campos, 1970: 98.- Guimaraes, 1971:
170.- Arnaud, 1978: 458.
Bucentes Latreille, 1809: 339. Type-species, B. cinereus Latreille cinereus Latreille, Bucentes (monotypy) = S. geniculata
(DeGeer).— Curran, 1932: 13.
Phantasiosiphona Townsend, 1915: 93. Type-species P. tropica Townsend (original designation).— Townsend, 1936: 150,
1940: 286.- Cortes, 1952: 1 10.— Guimaraes, 1971: 169. New synonymy.
Generic description of Siphona Meigen
The following generic diagnosis and description are based on study of adult specimens of
North American Siphona species. A few species from other regions do not fit this generic
description in all characteristics.
Recognition. - Small flies with narrow clypeus, subquadrangular head in profile and a long,
geniculate proboscis. Labella slightly longer than prementum, in most species longer than eye
height. Second aristomere elongate, at least 2.5X longer than wide. Anal vein extended to wing
margin at least as fold. R4+5 setulose dorsally between bifurcation of R2+3 and R4+5 and
crossvein r-m (beyond crossvein in S. lutea). Three katepisternal (sternopleural) setae,
antero-ventral as strong as or stronger than antero-dorsal. Most species with light coloured
pruinosity, particularly on preabdomen, few species darker but not black. Femur and tibia
predominantly yellow in most species, tarsi brown to black. Preabdomen without discal setae,
T1+2 with or without one pair each median and lateral marginal setae. Female genitalia
unmodified (Fig. 12). Male genitalia as in Figs. 1,60-80.
Description. - Length: 3. 0-6. 5mm.
Male. Head (Figs. 1,18-40). Front: wider than high, 1.1 -1.3 head height; clypeus narrowed, parallel-sided; distance
between vibrissae less than half head width (0.33-0.45).
Profile: subquadrangular; higher than wide, width at antennal axis 0.60-0.75 head height; frons slightly convex; face
vertical to slightly retreated, lower margin protruded beyond vibrissal angle; angle at scape between frons and facial ridge
in most species sharply defined, in few species broadly rounded; width of parafacial and gena wide or narrow, inversely
proportional to eye size.
Colouration: frontal vitta orange or yellow to testaceous, brown or reddish-brown; face, parafacial, gena and lower
third or less of postcranium light in ground colour, surface white or light yellow to light brown; ground colour of
fronto-orbital plate and upper postcranium light to dark, surface colour varied markedly, from yellow or gold to brown or
dark brown, darkest on orbital plate; ocellar triangle dark in ground colour, surface brown to dark brown; antenna highly
varied, yellow to black; palpus entirely yellow in most species, apex infuscate in few species; proboscis highly varied, yellow
to black.
Eye: bare or nearly so; size highly varied, small to large, slender to broad, widest at or above center, narrowed below
center or evenly rounded along anterior margin; eye height 0.627-0.91 1 head height (Fig. 13), average height for genus
between ca. 0.73-0.82 head height.
Quaest. Ent., 1982, 18 (1-4)
276
O’Hara
Macrotrichia: five frontal setae, medio-reclinate, anteriormost seta lateral to insertion of first flagellomere; 2nd and
4th frontal setae stronger than 1st, 3rd and 5th; 2 reclinate orbital setae in line with and posterior to frontal setae,
anterior seta intermediate in strength between largest frontal seta and inner vertical seta, posterior seta latero-reclinate,
in strength intermediate between shortest and longest frontal setae; 2 proclinate orbital setae, subequal in size, not
larger than largest frontal seta, lateral to frontal and reclinate orbital setae; anterior reclinate orbital seta from halfway
between proclinate orbital setae to lateral to posterior proclinate orbital seta (position intraspecifically varied); inner
vertical seta well-developed, subequal in size to vibrissa; outer vertical seta latero-reclinate, subequal in size to lower
reclinate orbital seta [appears shorter in most figures because of its inclination]; ocellar triangle with 1 pair
latero-proclinate ocellar setae and several hairs; postocellar setae divergent or parallel; paravertical seta subequal in size
to setulae of postocular row; postcranium uniformly and sparsely haired; upper parafacial and fronto-orbital plate
sparsely haired with less than 10 hairs in most species, to 15 or 20 in S. lurida and S. geniculata\ vibrissal angle with
2-10 setae in addition to vibrissa, and 1 to several hairs, average for genus 1 seta lateral to or above vibrissa and 1
below, and 2 or 3 hairs; 1 subvibrissal seta on antero-ventral margin of gena; aristomere 3 micropubescent; palpus with
2 or 3 setulae on lower surface, and several hairs; proboscis sparsely haired.
Antenna: first flagellomere extremely varied in length among species, 0.403-0.740 head height (Fig. 14); shape of
first flagellomere in profile from short and slender or broad, to long and slender or broad; length of aristomere 2
proportional to size of first flagellomere, in most species between 2. 5-6.0 times longer than wide, up to 8 times longer
than wide in species with very long first flagellomeres (eg. S. tropica, S. pisinnia and S. lurida)-, length of aristomere 3
0.50-0.65 head height in most species, shorter (0.35-0.45) in most specimens of S. hokkaidensis, S. geniculata and S.
medialis ; in all species except S. hokkaidensis, aristomere 3 not thickened beyond basal half, distally tapered to a fine
tip, in S. hokkaidensis aristomere 3 thickened almost to tip in most specimens.
Mouthparts: maxillary palpus slightly clavate, length 0.35-0.55 head height; labella elongate, straight at least on
basal half, apical 0.25 (on species with a long proboscis) to 0.5 (species with a short proboscis) flexible in live
specimens, twisted on most dried specimens; proboscis length (combined lengths of prementum and labella) varied from
1.3-3. 2 head height (Fig. 15), average for genus between ca. 1.8-2. 5 head height.
Thorax (Fig. 2). Colouration: average colouration for genus: ground colour black, with humeral callus and apex of
scutellum light in ground colour in most specimens; dorsum blue-gray or gray anteriorly, mixed with brown on posterior
prescutum, scutum and scutellum; amount of brown varied within and among species, in most species 3 faint to distinct
brown vittae visible on posterior prescutum and scutum; ground colour more apparent on pleura than dorsum, such that
light reflected from light gray, blue-gray, yellow or light brown pruinosity and black ground colour produces a sheen -
pleura alternately appearing light or dark as specimen rotated. A few species differ from average colouration in
possessing yellow thorax (in ground and surface colour). Wing tegula yellow to black; wing membrane yellowish,
cinereous or brownish hyaline. Legs, average for genus: femora yellow, or predominantly yellow and distally
reddish-brown on mid- and hindlegs and varied amounts of reddish-brown on dorsal and posterior surfaces of foreleg
(intraspecifically varied); tibia yellow, or predominantly yellow and basally reddish-brown when femur reddish-brown
distally. A few species with greater than average amounts of reddish-brown on femora and tibia, or fuscous overall.
Tarsi brown to black.
Macrotrichia: setae on prescutum and scutum: acrostichal (weak) 2-3 (presutural), 3-4 (postsutural); dorsocentral
2- 3, 3-4; intra-alar 1,3; supra-alar 3; humeral 2; posthumeral 1; presutural 1; notopleural 2; postalar 2. Scutellum:
strong basal, lateral and subapical, weak apical and discal. Pleura: proepisternal (propleural) 1; proepimeral
(prostigmatic) 1; anepisternal (mesopleural) row; anepimeral (pteropleural) 1 (weak); katepisternal (sternopleural) 3,
postero-dorsal very strong, others weaker, antero-ventral slightly longer than antero-dorsal; meral (hypopleural) row of
3- 4; katepimeron (barette) and anatergite (mediotergite) bare. Sterna: prosternum with a single pair of setulae in most
specimens, bare in few (or single setula on one side only). Foreleg: femur, pd row of 3-5 setae; pv surface with 1 to
several setae, strongest seta subadjacent to distal pd seta. Tibia, ad and pd surfaces each with row of 2-5 setae; p seta
ca. 0.4 tibial length from apex; preapical setae, well-developed d and pv, pd and v weak or absent. Midleg: femur, 2
adjacent a setae at midpoint; ca. 0.1 femoral length from apex, well-developed pd and p setae, weak ad seta; sparse row
of setulae on pv surface, with 1 strong setula or seta 0.25 femoral length from base. Tibia, ad seta slightly distad of
midpoint; 2 pd setae, ca. 0.3 and 0.6 tibial length from base; v seta ca. 0.6 tibial length from base; preapical setae,
well-developed ad, v and pv, other setae, if present, weak. Hindleg: femur, ad row of setae, increasing in strength
apically; very strong av seta ca. 0.25 femoral length from apex; 2 v setae basad of midpoint; pd seta ca. 0.1 femoral
length from apex. Tibia, ad and pd surfaces each with row of 4-6 setae, alternating weak and strong; 3-5 av setae from
ca. 0.4-0. 7 tibial length from base; preapical setae, d, ad, pd and av.
Acropod (Figs. 4-6): claws and pulvilli varied in size, subequal on all legs; length of each claw and pulvillus from
ca. width of tarsomere 5 (ie. averaged-sized, Figs. 4,6) to ca. length of tarsomere 5 (ie. large, Fig. 5).
Wing (Fig. 3): short and broad; Sc incised adjacent to subcostal break; apices of R4+5 and M closely approximated
or contiguous in wing margin, slightly anterior to wing tip; anal vein (A,) reaching wing margin at least as fold; R,
bare dorsally except for few species with 1 or 2 setulae on bend of R,; R4+5 dorsally with row of 3-10 setulae between
bifurcation of R2+3 and R4+5 and crossvein r-m; setulae extended beyond crossvein r-m in most specimens of S. lutea\
CuA, bare dorsally except for 1-3 setulae in very few specimens of S. lutea-, single setula ventrally at bifurcation of
R2+3 and R4+5.
North American species of Siphona Meigen
277
Preabdomen (Figs. 7-8): Ovoid to slightly elongate in dorsal view; middorsal depression extended to middle of T1+2.
Colouration: highly varied intra- and interspecifically; most species dorso-medially vittate, average pattern T4 and
T5 black in ground colour, vitta on T]+2 and T3 (width averaging distance between median marginals on T3), rest of
preabdomen yellow in ground colour; areas black in ground colour with surface blue-gray, gray, brownish-gray or
brown, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow, light
brown or tawny. Preabdomen entirely black in ground colour in a few species, at least dorsally and laterally, or with
very reduced areas of yellow ground colour on extreme lateral edge of T] + , and antero-lateral edge of T3; preabdomen
entirely yellow in ground and surface colour in a few other species.
Macrotrichia: discals absent; T1+2 without median marginals in most species, 1 seta on one side or 1 pair present in
few species; T,+2 with 1 pair strong lateral marginals in most species (ca. 0.75 length of lateral marginals on T3 or
longer), weak pair in few species (less than 0.5 length of lateral marginals on T3), 2 pair strong lateral marginals in few
specimens of few species; T3 with 1 pair each strong median and lateral marginals in most species, few specimens of few
species with 2 pair lateral marginals; T4 with 3 pair marginals; T5 with 3-5 pair marginals.
Genitalia (Figs. 9,60-80). Epandrium in profile saddle-shaped, slightly higher than long, antero-lateral margin with
medially-angled flange; hypandrial apodeme articulated with antero-ventral margin of epandrium; gonopod
sub-triangular, with a flange produced ventro-laterally; paramere lobe-like, ca. twice as long as wide; surstylus narrow,
apex even with or extended beyond apex of cerci; cerci fused medially along lower half, long and narrow, smoothly
tapered with slight inflexion at midpoint, apex curved anteriorly; epiphallus not developed.
Aedeagus: distiphallus in profile triangular to quadrangular, surface Finely dentate, antero-ventral margin from
dentate to bearing well-developed hooks; angle of distiphallus from even with angle of basiphallus (eg. S. illinoiensis) to
sharply bent downward (eg. S'. intrudens)\ apex in profile from narrow (eg. S. tropica) to broad (eg. S. medialis).
Sternum 5 (Figs. 10-11): posterior margin U-shaped (not V-shaped as in most Actia species); notched
postero-medially; all North American species except S. oligomyia with postero-lateral arms smoothly curved around
posterior margin, S. oligomyia with arms hooked inward.
Female. As described for male except as follows.
Head (Figs. 41-59). Eye and first flagellomere smaller than in male; female palpus of few species longer than in
male.
Thorax. Acropod: smaller than in males; most species with large claws and pulvilli in male have only slightly larger
than average claws and pulvilli in female.
Preabdomen. Colouration: same range as described for male; female darker than male in most species; females of
species with vittate males in most species with more extensive areas of black ground colour than males, vitta shorter and
broader than in males, or absent.
Genitalia (Fig. 12). Unmodified; structurally very similar among Siphona females; terga 6, 7, 9 and 10 lost; tergum
8 dorso-longitudinally interrupted; sternum 9 lost, or fused with sternum 10 (ref. Herting, 1957); cerci well developed;
two spermathecae.
World list of Siphona species
This list is inclusive of all Siphona species described before August, 1981, plus those
described as new herein4. The genus in which each species was described is given only if not
Siphona. A capital letter to left of species name denotes the region from which the species was
described with, for a few species, a second letter denoting presence of the species in another
region.
Species of which specimens were examined during the course of this revision are so indicated
in the list. Unless otherwise stated, specimens examined were not type material. Status of
species not examined follows recent revisionary or catalogue placements. New species,
synonymies, combinations and lectotype designations are noted.
Types of nominal species of Siphona described by Robineau-Desvoidy (1830, 1850) are lost
(Herting, per. comm.), and although most were probably conspecific with S. geniculata, their
status cannot be verified so the names are treated as nomina dubia.
Tour new species of European Siphona are described by Andersen (in press “a”). They are not
listed here to ensure the new names are first published in his paper.
Quaest. Ent., 1982, 18 (1-4)
278
O’Hara
Regional code:
A Afrotropical, sensu Crosskey and White, 1977.
N North America (America north of Panama-Colombia border).
O Oriental, sensu Crosskey, 1976a.
P Palearctic, as limited by Afrotropical and Oriental regions in their present
interpretation.
S South America.
I have examined specimens of a new species of Siphona from Australia, but as yet no
Siphona have been described from that region. The tribe Siphonini is apparently not
represented in New Zealand (Dugdale, 1969).
A abbreviata (Villeneuve), 1915: 199 ( Bucentes ) [examined],
N akidnomyia new species.
A albocincta (Villeneuve), 1942: 55 ( Bucentes ) [examined],
A amoena (Mesnil), 1952: 12 ( Crocuta ) [examined].
A amplicornis Mesnil, 1959: 21.
syn. nigrohalterata Mesnil, 1959: 22 (as ssp.).
A angusta Mesnil, 1959: 22.
A antennalis (Mesnil), 1952: 9 ( Crocuta ) [examined],
A atricapilla Mesnil, 1959: 20.
A bevisi Curran, 1941: 7 [holotype examined].
A bilineata (Mesnil), 1952: 10 ( Crocuta ) [examined].
P boreata Mesnil, 1960: 190 [holotype examined],
N brunnea new species.
A capensis Curran, 1941: 7.
P collini Mesnil, 1960: 188 [holotype examined].
P confusa Mesnil, 1961: 201 [holotype examined].
A cothurnata (Mesnil), 1952: 17 ( Crocuta ) [examined].
A creberrima (Speiser), 1910: 142 {Crocuta).
P,N cristata (Fabricius), 1805: 281 ( Stomoxys ) [lectotype examined],
syn. chaetolyga Rondani, 1865: 194.
palpina Zetterstedt, 1859: 6064.
A cuthbertsoni Curran, 1941: 7 [holotype examined].
P efflatouni Mesnil, 188 [holotype examined].
P flavifrons Staeger in Zetterstedt, 1849: 321 1 [examined].
N floridensis new species.
O foliacea (Mesnil), 1953: 1 13 {Crocuta {Siphona )) [examined].
A fuliginea Mesnil, 1977: 77.
ssp. cerina Mesnil, 1977: 76.
ssp. rubea Mesnil, 1977: 77.
N futilis Wulp, 1890: 125 [lectotype designated].
syn. ceres (Curran), 1932: 14 ( Bucentes ) [holotype examined, new
synonymy].
O gedeana Wulp, 1896: 109.
syn. nigripalpis (de Meijere), 1924: 223 {Bucentes).
P,N geniculata (De Geer), 1776: 20 {Musca) [examined],
syn. analis Meigen, 1824: 157.
North American species of Siphona Meigen
279
cinerea Meigen, 1824: 156.
cinerea (Latreille), 1809: 339 ( Bucentes ).
meigenii (Lepeletier and Serville in Latreille, et al.), 1828: 501
{Bucentes).
minuta (Fabricius), 1805: 282 ( Stomoxys ).
nigrovittata Meigen, 1824: 157.
tachinaria Meigen, 1824: 156.
urbanis (Harris), 1780: 153 ( Musca ).
A gracilis (Mesnil), 1952: 13 ( Crocuta ) [examined],
P grandistyla Pandelle, 1894: 108 [examined].
P,N hokkaidensis Mesnil, 1957: 36 [holotype examined],
syn. silvarum Herting, 1967: 9 [holotype examined, new synonymy].
N illinoiensis Townsend, 1891: 368 [lectotype designated].
N intrudens (Curran), 1932: 14 ( Bucentes ) [holotype examined],
A janssensi (Mesnil), 1952: 4 ( Crocuta ) [examined].
S kuscheli (Cortes), 1952: 1 10 ( Phantasiosiphona ) [examined, new combination].
A laticornis Curran, 1941: 9.
A lindneri Mesnil, 1959: 22.
N longissima new species.
N,P lurida Reinhard, 1943: 20 [holotype examined],
N lutea (Townsend), 1919: 584 ( Crocuta ) [lectotype designated].
syn. tenuis Curran, 1933: 10 [holotype examined, new synonymy].
N macronyx new species.
P,N maculata Staeger in Zetterstedt, 1849: 3212 [holotype examined].
ssp. griseola Mesnil, 1970: 1 18 [holotype examined].
P maculipennis Meigen, 1830: 365 (unrecognized).
N medialis new species.
A melania (Bezzi), 1908: 58 {Bucentes).
A melanura Mesnil, 1959: 23.
N multifaria new species.
A munroi Curran, 1941: 6.
A murina (Mesnil), 1952: 15 {Crocuta) [examined].
P nigricans (Villeneuve), 1930: 100 {Bucentes) [holotype examined].
A nigroseta Curran, 1941: 8.
O nobilis (Mesnil), 1953: 1 12 {Crocuta {Siphona)).
A obesa (Mesnil), 1952: 8 {Crocuta) [examined],
A obscuripennis Curran, 1941: 8 [holotype examined].
N oligomyia new species.
N pacifica new species.
P paludosa Mesnil, 1960: 188 [examined].
A patellipalpis (Mesnil), 1952: 10 {Crocuta) [examined],
P pauciseta Rondani, 1865: 193 [examined].
syn. delicatula Mesnil, 1960: 190.
oculata Pandelle, 1894: 108.
A phantasma (Mesnil), 1952: 7 {Crocuta) [examined].
A pigra Mesnil, 1977: 78.
Quaest. Ent., 1982, 18 (1-4)
280
O’Hara
N pisinnia new species.
S pseudomaculata Blanchard, 1963: 233.
A reducta (Mesnil), 1952: 18 ( Crocuta ) [examined].
ssp. ludicra Mesnil, 1977: 78.
N rizaba new species.
P rossica Mesnil, 1961: 202 [examined].
A rubrapex Mesnil, 1977: 79.
A rubrica (Mesnil), 1952: 11 ( Crocuta ) [examined].
A setinerva (Mesnil), 1952: 16 ( Crocuta ) [examined],
P setosa Mesnil, 1960: 191 [examined],
P seyrigi Mesnil, 1960: 189 [holotype examined].
A simulans (Mesnil), 1952: 18 {Crocuta).
S singulars (Wiedemann), 1830: 335 ( Tachina ) (unrecognized).
A sola Mesnil, 1959: 21 [examined].
A spinulosa (Mesnil), 1952: 12 ( Crocuta ) [examined],
A trichaeta (Mesnil), 1952: 18 ( Crocuta ) [examined].
N,S tropica (Townsend), 1915: 93 ( Phantasiosiphona ) [paratype examined, new
combination].
A unispina (Mesnil), 1952: 14 {Crocuta) [examined].
syn. infuscata (Mesnil), 1952: 14 ( Crocuta , as ssp.).
A vittata Curran, 1941:8 [holotype examined].
A vixen Curran, 1941: 9 [holotype examined].
A wittei (Mesnil), 1952: 5 {Crocuta) [examined].
Nomina Dubia
analis Robineau-Desvoidy, 1830: 92.
clausa Robineau-Desvoidy, 1850: 209.
consimilis Robineau-Desvoidy, 1850: 205.
fuscicornis Robineau-Desvoidy, 1850: 205.
humeralis Robineau-Desvoidy, 1850: 207.
melanocera Robineau-Desvoidy, 1850: 206.
pusilla Robineau-Desvoidy, 1830: 92.
syn. persilla. Misspelled reference to pusilla Robineau-Desvoidy
(Coquillett, 1897: 76).
quadrinotata Robineau-Desvoidy, 1850: 203.
syn. quadricincta. Misspelled reference to quadrinotata
Robineau-Desvoidy (Rondani, 1859: 10).
silvatica Robineau-Desvoidy, 1850: 208.
testacea Robineau-Desvoidy, 1850: 207.
tristis Robineau-Desvoidy, 1850: 203.
Nomina Nuda
anthomyformis Lynch Arribalzaga in Brauer, 1898: 505 (1898: 13).
dorsalis Brauer and Bergenstamm, 1891: 410.
pauciseta Mesnil, 1964: 856 (as ssp. of geniculata).
taiwanica (Baranov in Hennig), 1941: 195 {Crocuta).
North American species of Siphona Meigen
281
Keys to adults of North American Siphona species
Siphona species are not sufficiently different and are too intraspecifically varied to permit
construction of a key based solely on external, diagnostic characteristics. Therefore I found it
necessary to frequently employ mensural characteristics (based on data presented in
Hubbs-Hubbs diagrams Figs. 13-16) and features of male genitalia in the following keys to
North American Siphona species. As a consequence, female specimens do not key as accurately
as males, and should only be used if male associates are unavailable. Even males of S. cristata
and 5*. multifaria cannot always be separated (couplet 19"), while a few specimens of other
species may be outside the recorded range of variation.
Given certain conditions it should be possible to identify most Siphona specimens. Firstly,
males are more reliably keyed than females. Secondly, it is advisable to check characteristics of
several male specimens of the same species against values given in the Hubbs-Hubbs diagrams
to confirm a species identification. Thirdly, the “Recognition” section under each species
treatment has been included to assist identifications by summarizing key features of a
description and contrasting similar species. To simplify keying of North American Siphona
specimens, species were divided into two geographical groups and keyed separately: (1) species
of Canada and the United States, and (2) species of Middle America. S. pisinnia is shared
between regions and is keyed twice. S. multifaria is recorded almost to the Mexican border in
the United States, so it is included in the key to Middle American species.
The Siphona fauna of Middle America is inadequately known. As more specimens of known
species become available the key will probably require modification to incorporate greater
intraspecific variation. Similarly, the key will have to be modified for inclusion of currently
undiscovered species.
Most terms are as used in the “Manual of Nearctic Diptera, Volume 1” (McAlpine, 1981).
See “Terms” section and genus description for information regarding Siphona characters.
Figure numbers of male and female head profiles are cited following each species name.
Male genitalia of all species are shown in Figs. 9, 60-80. The geographical distribution of each
species is given as an aid to identification of specimens.
Key to adults of Siphona species of Canada and United States
1 T1+2 with one pair strong median marginal setae (Fig. 8) 2
V T1+2 without median marginal setae (Fig. 7), or one strong seta on one side
only, or one pair scarcely differentiated from adjacent marginal hairs 6
2 (1) Surstylus with apex even with or extended slightly beyond apex of cerci (Fig.
72); distiphallus slightly or sharply bent downward from angle of basiphallus;
specimens from Washington, Idaho, Oregon and California with femora and
tibiae reddish-brown to fuscous, specimens from other areas with dorsal and
posterior surfaces of fore femur reddish-brown to fuscous in few specimens,
other specimens with legs average (ie. femoral-tibial articulation reddish-brown,
rest of femur and tibia yellow); transcontinental, most widely recorded from
western USA (Fig. 94) S. medialis n. sp. (Figs. 29, 52), p. 303
2' Surstylus with apex extended beyond apex of cerci by more than preapical width
of surstylus (Figs. 70-71); slope of distiphallus even with that of basiphallus; leg
colouration average 3
Quaest. Ent., 1982, 18 (1-4)
282
O’Hara
3 (2') Male 4
3' Female 5
4 (3) Distiphallus parallel-sided in profile, slender in anterior view (Fig. 70);
aristomere 3 tapered to fine tip; first flagellomere short and narrow, length
0.42-0.48 head height; introduced to, and only recorded from lower, Fraser
Valley of British Columbia (Fig. 91)
S. geniculata (De G.) (Fig. 28), p. 299
4' Distiphallus slightly broader at base than apex in profile, average Siphona
width in anterior view (Fig. 71); aristomere 3 thickened to near tip in most
specimens; first flagellomere varied, from short and narrow to long and broad,
length 0.43-0.67 head height; transcontinental (Fig. 81)
S. hokkaidensis Mesn. (in part) (Figs. 30-32), p. 301
5 (3') First flagellomere with apex truncate in most specimens; transcontinental (Fig.
81) S. hokkaidensis Mesn. (in part) (Fig. 51), p. 301
5' First flagellomere with apex rounded along ventral margin; introduced to, and
only recorded from, lower Fraser Valley of British Columbia (Fig. 91)
S. geniculata (De G.) (Fig. 50), p. 299
6 (T) Body pale yellow overall in ground colour 7
6 ' Body reddish-brown to black in ground colour at least on thoracic dorsum 8
7 (6) Male sternum 5 with antero-lateral arms not hooked inward (Fig. 10); proboscis
shorter than 2.2X head height; most specimens with R4+5 setulose beyond
crossvein r-m on at least one side; transcontinental (Fig. 89)
S’, lutea (Tnsd.) (Figs. 26,48), p. 296
1' Male sternum 5 with antero-lateral arms hooked inward (Fig. 11); proboscis
longer than 2.3X head height; R4+5 not setulose beyond crossvein r-m; western
North America (Fig. 90); female unknown . S. oligomyia n. sp. (Fig. 27), p. 297
8 (60 Proboscis shorter than 1.8X head height 9
8' Proboscis longer than 1.8X head height 13
[S. intrudens keys through both halves of couplet]
9 (8) Vibrissal angle with 5-10 setae in addition to vibrissa and several hairs; eye very
small, in male less than 0.68 head height; T1+2 with strong lateral marginal
setae; western USA (Fig. 86) S. lurida Rnh. (Figs. 21, 41), p. 290
9' Vibrissal angle with 2-3 setae in addition to vibrissa and several hairs; male eye
larger than 0.70 head height; T1+2 with weak or strong lateral marginal setae 10
10 (9') Four postsutural dorsocentral setae; distiphallus dentate along antero-ventral
margin (Fig. 61); Florida and southern Georgia (Fig. 84)
S. floridensis n. sp. (Figs. 19, 43), p. 288
10' Three postsutural dorsocentral setae; distiphallus with several distinct, small or
large hooks along antero-ventral margin (Figs. 60, 62) 11
11 (100 Male 12
1 V Female S. maculata Staeg. (Fig. 42), p. 286
and S. intrudens (Cn.) (Fig. 44), p. 288
[separation difficult without male associates; see discussion under S. maculata,
“Recognition”]
12 (11) Distiphallus U-shaped in anterior view, with 3-4 large hooks along
antero-ventral margin (Fig. 62); preabdomen black in ground colour overall, or
North American species of Siphona Meigen
283
vittate and yellow in ground colour laterally on T1+2 and T3; transcontinental
(Fig. 85) S. intrudens (Cn.) (in part) (Fig. 20), p. 288
12' Distiphallus V-shaped in anterior view, with several small hooks (Fig. 60);
preabdomen vittate, yellow in ground colour laterally on T1+2 and T3;
transcontinental (Fig. 82) S. maculata Staeg. (Fig. 18), p. 286
13 (8') Distiphallus U-shaped in anterior view, with 3-4 large hooks along
antero-ventral margin (Fig. 62); 3 postsutural dorsocentral setae; T1+2 with
weak lateral marginal setae in most specimens; proboscis shorter than 2. OX
head height; transcontinental (Fig. 85)
S. intrudens (Cn.) (in part) (Figs. 20, 44), p. 288
13' Distiphallus V-shaped in anterior view, anteroventral margin dentate or with
small hooks; 3 or 4 postsutural dorsocentral setae; T1+2 with weak or strong
lateral marginal setae, if weak (some specimens of S. pacifica ), then 4
postsutural dorsocentral setae; proboscis longer than 1.85X head height 14
14 (130 Male 15
14' Female 15
15 (14) Surstylus with apex extended beyond apex of cerci by more than preapical width
of surstylus (Fig. 71); short, thick aristomere 3 (0.35-0.45 head height) in most
specimens; most specimens from eastern North America with broad, truncate
first flagellomere (Fig. 31), specimens from elsewhere with first flagellomere
varied (Figs. 30,32); transcontinental (Fig. 81)
S. hokkaidensis Mesn. (in part) (Figs. 30-32), p. 301
15' Surstylus with apex even with or extended slightly beyond apex of cerci;
aristomere 3 of average length (0.50-0.65 head height), tapered to fine tip; first
flagellomere with shape varied 16
16 (15') Length of first flagellomere less than 0.50 head height; western USA 17
16' Length of first flagellomere greater than 0.50 head height; widely distributed . . 18
17 (16) Three postsutural dorsocentral setae; tarsal claws large (as in Fig. 5); vibrissal
angle with 3-6 setae and several setulae in addition to vibrissa and hairs;
distiphallus sharply bent downward from basiphallus, with small hooks on
antero-ventral margin (Fig. 75); head with characteristic habitus in profile;
recorded from Washington and western Idaho (Fig. 88)
S. macronyx n. sp. (Fig. 33), p. 313
17' Four postsutural dorsocentral setae; tarsal claws of average size (as in Figs. 4,6);
vibrissal angle with 2-3 setae in addition to vibrissa and hairs; distiphallus bent
slightly downward from angle of basiphallus, dentate on antero-ventral margin
(Fig. 64); recorded from California and Washington (Fig. 87)
S. pacifica n. sp. (Fig. 22), p. 291
18 (17') Proboscis shorter than 2.3X head height; length of first flagellomere greater
than 0.76 eye height; first flagellomere fuscous to black; southwestern USA and
Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 23), p. 298
18' Proboscis longer than 2.25X head height, or from central or eastern North
America ( S . illinoiensis ); length of first flagellomere less than 0.85 eye height;
first flagellomere varied in colour, testaceous to black 19
19 (18') Distiphallus narrow in profile, apex sloped sharply postero-ventrally in most
specimens (Fig. 9); slope of distiphallus even with that of basiphallus in most
Quaest. Ent., 1982, 18 (1-4)
284
O’Hara
specimens, slightly bent in few; proboscis shorter than 2.5X head height; length
of first flagellomere less than 0.69 eye height; central and eastern North
America (Fig. 92) S. illinoiensis (Tnsd.) (Fig. 1), p. 307
19' Distiphallus broad in profile, apex not sloped sharply postero-ventrally in most
specimens (Figs. 66-67); distiphallus bent sharply downward from angle of
basiphallus; proboscis longer than 2.25X head height; length of first
flagellomere greater than 0.65 eye height; transcontinental (Figs. 93, 95)
S. cristata (Fabr.) (Fig. 24), p. 292
and S. multifaria n. sp. (Fig. 25), p. 293
[see “Recognition” under S. cristata for discussion concerning separation of
these species]
20 (14r) First flagellomere truncate apicaily; aristomere 3 short (0.35-0.45 head height)
in most specimens; transcontinental (Fig. 81)
S. hokkaidensis Mesn. (in part) (Fig. 51), p. 301
20' First flagellomere rounded distally along ventral margin; aristomere 3 average
length (0.50-0.65 head height) 21
21 (20') Vibrissal angle with 3-6 setae and several setulae in addition to vibrissa and
hairs; 3 postsutural dorsocentral setae; recorded from Washington and western
Idaho (Fig. 88) S. macronyx n. sp. (Fig. 56), p. 313
2 1' Vibrissal angle with 2-3 setae in addition to vibrissa and hairs; 3-4 postsutural
dorsocentral setae 22
22 (2F) Proboscis shorter than 2.3X head height; western USA 23
22 ' Proboscis longer than 2.25X head height, or from central or eastern North
America (S. illinoiensis) 24
[for specimens from western USA within or near overlap in proboscis length,
check distribution maps and see “Recognition” under appropriate species]
23 (22) Four postsutural dorsocentral setae; recorded from California and Washington
(Fig. 87) S. pacifica n. sp. (Fig. 45), p. 291
23' Three postsutural dorsocentral setae in most specimens; recorded from
southwestern USA and Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 49), p. 298
24 (220 Eye large (Fig. 54); proboscis shorter than 2.5X head height; central and
eastern North America (Fig. 92) S. illinoiensis (Tnsd.) (Fig. 54), p. 307
24' Eye of average size (Figs. 46-47); proboscis longer than 2.25X head height;
transcontinental (Figs. 93, 95) S. cristata (Fabr.) (Fig. 46), p. 292
and S. multifaria n. sp. (Fig. 47), p. 293
[see “Recognition” under S. cristata for discussion concerning separation of
these species]
Key to adults of Siphona species of Middle America
1 Male 2
V Female 9
[female of S. longissima unknown]
2 (1) Length of first flagellomere less than 0.52 head height 3
2' Length of first flagellomere greater than 0.58 head height, or not south of
Tropic of Cancer (5. multifaria) 4
North American species of Siphona Meigen
285
3 (2) Preabdomen light coloured with narrow vitta; 3 postsutural dorsocentral setae in
all but very few specimens; palpus yellow in most specimens, slightly infuscate
apically in few; distiphallus triangular and pointed in profile (Fig. 73); widely
distributed, from central Mexico to Costa Rica (Fig. 96)
S.futilis Wulp (Fig. 34), p. 305
3' Preabdomen dark brown, not vittate, with narrow light coloured bands across
anterior of T3, T4 and T5; 4 postsutural dorsocentral setae; palpus fuscous or
black on apical half; distiphallus slender, apex truncate in profile (Fig. 74);
Chiapas (Fig. 97) S. brunnea n. sp. (Fig. 35) p. 308
4 (20 Preabdomen broadly vittate (vitta wider than distance between median
marginal setae on T3), dark coloured; palpus infuscate apically; distiphallus
short and broad in profile with small hooks along antero-ventral margin (Fig.
76); southern Mexico (Fig. 99) S. akidnomyia n. sp. (Fig. 36), p. 31 1
4' Preabdomen average vittate (vitta not wider than distance between median
marginal setae on T3), light coloured; palpus yellow; distiphallus not shaped as
above, with or without small hooks along antero-ventral margin 5
5 (4') Proboscis shorter than 2.3X head height; distiphallus slightly bent downward
from basiphallus, shaped as in Fig. 65; southwestern USA to Mexico state,
Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 23), p. 298
5' Proboscis longer than 2.6X head height, or if shorter ( S . multifaria 2. 25-3. OX)
then distiphallus sharply bent downward from basiphallus and not south of
Tropic of Cancer 6
6 (5") North of Tropic of Cancer; distiphallus as in Fig. 67; widely distributed in
Canada and USA (Fig. 93), presence in northern Mexico likely
S. multifaria n. sp. (Fig. 25), p. 293
6' South of Tropic of Cancer 7
7 (60 Length of first flagellomere less than 0.75 eye height; surstylus with apex
extended slightly to far beyond apex of cerci (Fig. 79); distiphallus with apex
rounded in profile; bend of R] bare; Veracruz (Fig. 98)
S. rizaba n. sp. (Fig. 39), p. 310
T Length of first flagellomere greater than 0.75 eye height; surstylus with apex
even with or extended slightly beyond apex of cerci (Figs. 77-78); distiphallus
with apex pointed in profile; bend of R, with single setula or bare 8
8 (70 Eye height greater than 0.86 head height; bend of R, with single setula on at
least one side in most specimens; slope of distiphallus even with that of
basiphallus; distiphallus dentate along antero-ventral margin (Fig. 78);
Veracruz to Costa Rica (Fig. 101) S. tropica (Tnsd.) (Fig. 38), p. 309
8' Eye height less than 0.85 head height; bend of R, bare; distiphallus bent sharply
downward from angle of basiphallus; distiphallus with small hooks along
antero-ventral margin (Fig. 77); Chiapas (Fig. 100)
S. longissima n. sp. (Fig. 37), p. 31 1
9 (10 Proboscis longer than 2.7X head height 10
9' Proboscis shorter than 2.7X head height 12
[N. multifaria keys through both halves of couplet]
10 (9) North of Tropic of Cancer; widely distributed in Canada and USA (Fig. 93),
presence in northern Mexico likely
Quaest. Ent., 1982, 18 (1-4)
286
O’Hara
S. multifaria n. sp. (in part) (Fig. 47), p. 293
10' South of Tropic of Cancer 11
11 (100 Bend of with a single setula on at least one side in most specimens; Veracruz
to Costa Rica (Fig. 101) S. tropica (Tnsd.) (Fig. 57), p. 309
1 V Bend of Rj bare; Veracruz (Fig. 98) S. rizaba n. sp. (Fig. 58), p. 310
[male associates required for positive identification]
12 (90 Proboscis longer than 2.5X head height; wing brownish hyaline; southern
Mexico (Fig. 99) S. akidnomyia n. sp. (Fig. 59), p. 31 1
12' Proboscis shorter than 2.5X head height; wing yellowish or brownish hyaline 13
13 (120 Palpus with apical half fuscous or black; 4 postsutural dorsocentral setae; body
dark brown; Chiapas (Fig. 97) S'. brunnea n. sp. (Fig. 55), p. 308
13' Palpus yellow; 3 postsutural dorsocentral setae in all but very few specimens, or
not south of Tropic of Cancer; body blue-gray with brown areas, not extensively
dark brown 14
14 (130 Head profile as shown in Fig. 53; eye relatively large, first flagellomere
relatively short; widely distributed, from central Mexico to Costa Rica (Fig. 96)
S.futilis Wulp (Fig. 53), p. 305
14' Head profiles as shown in Figs. 47,49; eye relatively small; first flagellomere
relatively long 15
15 (140 Proboscis shorter than 2.3X head height; southwestern USA to Mexico state,
Mexico (Fig. 83) S. pisinnia n. sp. (Fig. 49), p. 298
15" Proboscis longer than 2.25X head height; widely distributed in Canada and
USA (Fig. 93), presence in northern Mexico likely
S. multifaria n. sp. (in part) (Fig. 47), p. 293
The S. maculata group
This group is characterized by a short to very short proboscis and a slender first flagellomere
broadly rounded apically along ventral margin. Weak lateral marginal setae on T1+2 are
presumed symplesiotypic for the group, but this state is not expressed in all specimens (and not
at all in Siphona lurida). There is a trend toward a smaller than average eye. Except for
Siphona pacifica, adults are seen in spring and early summer.
Siphona maculata Staeger
Siphona maculata Staeger in Zetterstedt, 1849: 3212- Mesnil, 1965: 871 (redescription).
Type material examined. - Lectotype (by designation of Andersen, in press “a”), male,
labelled: “<$/ Chariot-/ tenlund [Denmark] [handwritten]”; “Coll./ Staeger [yellow label]”;
“TYPE [red label]”; “Macula/ ta Staeg. [handwritten]”; “TYPE [red label]”; “Lectotype 6
[on left side of label]/ Siphona maculata/ Staeger in Zett. [handwritten]/ det. Stig. Andersen”
(ZMUC).
Recognition. - Average-sized eye and first flagellomere, very short proboscis. Most
distinctive features are weak lateral marginal setae on T1+2 and small but distinct hooks on
antero-ventral margin of distiphallus. Colouration of preabdomen average, vittate. Three
postsutural dorsocentral setae.
Similar to S. floridensis , from which it can be separated by number of postsutural
dorsocentrals, structure of distiphallus and geographical distribution.
North American species of Siphona Meigen
287
It is most easily confused with S. intrudens. Males are separated by structure of distiphallus.
There are several trends which are useful for distinguishing between S. intrudens and S.
maculata, these apply to both male and female specimens, though more conspicuous in former:
S. maculata has a shorter proboscis, smaller tarsal claw, larger eye and longer first
flagellomere than S. intrudens. With respect to females, preabdomen of most specimens of S.
intrudens is black in ground colour overall, while preabdomen of S. maculata in most
specimens has a short, broad vitta, with yellow ground colour laterally on T1+2 and
antero-laterally on T3.
Description. — Specimens examined: 263 males, 117 females from North America; ca. 20 European specimens.
Length: 3. 5-5. 0mm.
Male. Head (Fig. 18). Colouration: frontal vitta yellow to testaceous; gena and parafacial white to light yellow; frontal
plate light yellow to light brown; scape and pedicel yellow to fuscous, first flagellomere in most specimens fuscous or black,
testaceous in few light coloured specimens; palpus yellow; proboscis reddish-brown to fuscous. Eye: size average for genus,
0.755-0.825 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: average
for genus. Antenna: first flagellomere 0.564-0.640 head height (Fig. 14); length of aristomere 3 average for genus.
Mouthparts: proboscis short, 1 .30-1 .74 head height (Fig. 1 5).
Thorax. Colouration: dorsum average, 3 brown vittae distinct in light coloured specimens, dorsum predominantly
brown to dark brown in dark coloured specimens; tegula testaceous, light brown to brown; wing yellowish hyaline; legs
average. Macrotrichia: postsutural dorsocentrals, 44 specimens with three, 1 with four [males and females], Acropod:
claws and pulvilli average-sized.
Preabdomen. Colouration: faintly vittate; most specimens black in ground colour on T4 and T5 and medially on Tl+2
and T3, yellow in ground colour laterally on T1+2 and T3; areas black in ground colour with surface blue-gray or gray, dark
brown or black around macrotrichial insertions, especially around median marginals on T3 and T4; areas yellow in ground
colour with surface yellow to light brown; extent of black ground colour varied among specimens, in few greater than
average (posterior portion of T3 black in ground colour) or less than average (anterior portion of T4 yellow in ground
colour). Macrotrichia: median marginals absent from T, +2; lateral marginals on T]+2 weak, in most specimens scarcely or
not at all differentiated from marginal hairs.
Genitalia (Fig. 60). Thirty-eight examined, including 2 from European specimens. Apex of surstylus even with or
extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus;
antero-ventral margin V-shaped in anterior view, armed with several well-developed hooks (though smaller than hooks on
distiphallus of S. intrudens ); in profile tapered to narrow apex.
Female. As described for male except as follows.
Head (Fig. 42). Palpus longer and thicker than in males in few specimens.
Preabdomen. Colouration: highly varied, most specimens darker than average male; average specimen yellow in
ground colour laterally on T|+2 and antero-laterally on T3, rest black in ground colour, dorsal vitta short and broad; range
from entirely black in ground colour (vitta absent), to average condition described for male (vitta present); dark specimens
with large dark brown or black areas around macrotrichial insertions.
Geographical distribution (Fig. 82). - Europe and North America. Transcontinental in
North America; most records within a rather narrow belt along southern limit of boreal forest.
Habitat. - I have collected specimens of this species in mixed forest dominated by maple
(Gatineau Co., Que.) or aspen (Edmonton, Alta.). Individuals flew close to the ground in a zig
zag pattern, apparently searching for mates or ground-dwelling hosts.
Chorological affinities. - S. maculata and S. intrudens are widely sympatric over much of
their apparent ranges. A few examined series of specimens contain both S. intrudens and S.
maculata , suggesting their ecological requirements are similar.
A closely related species, S. floridensis, is allopatric in distribution.
Phylogenetic relationships (Fig. 103). - The phenetic similarity between S. maculata and
S. floridensis suggests they might be sister species, though a synapotypy has not been found to
substantiate this.
Of the North American taxa of the S. maculata group, only S. maculata and S. intrudens
share the derived condition of hooks on the antero-ventral margin of the distiphallus (Figs. 60,
62). However, on the basis of other characters, this condition is interpreted as derived at the
base of the lineage terminating in S. maculata , S. intrudens and S. lurida (Fig. 103),
Quaest. Ent., 1982, 18 (1-4)
288
O’Hara
precluding a sister group relationship between S. maculata and S. intrudens.
Siphona floridensis new species
Holotype. - Male, labelled: “Orlando,/ Fla. [Florida, USA] III.3 . [ 1 9] 46 / F. Snyder”
(AMNH). Genitalia in microvial on pin below specimen.
Allotype. - Female, same data as holotype (AMNH).
Derivation of specific epithet. - With only one record outside Florida, this species is named
for its restricted known distribution.
Recognition. - Average-sized eye and first flagellomere, very short proboscis. Lateral
marginal setae on T1+2 weak or strong. Distiphallus average in structure. Preabdomen vittate
or black in ground colour overall. Four postsutural dorsocentral setae.
Externally very similar to the allopatric S. maculata. The proboscis of both species is very
short. S. floridensis has a slightly larger eye and slightly shorter first flagellomere than S.
maculata. Males are separated by lack of hooks on antero-ventral margin of distiphallus (small
hooks present in S. maculata) and number of postsutural dorsocentrals (three in S. maculata ,
four in S. floridensis).
Description. - Length: 3. 5-5. 5mm.
Male. Head (Fig. 19). Colouration: as described for S. maculata. Eye: size average for genus, 0.783-0.839 head height
(Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first
flagellomere slightly shorter and thinner than in S. maculata , 0.548-0.574 head height (Fig. 14); length of aristomere 3
average for genus. Mouthparts: proboscis short, within range for S. maculata , 1.37-1.59 head height (Fig. 15).
Thorax. Colouration: dorsum average, 3 light brown vittae visible in most specimens, scutum and scutellum
predominantly brown in few specimens (generally somewhat lighter than in S. maculata ); tegula testaceous; wing
yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, all specimens with four [males and females].
Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: specimens in “Orlando” series vittate, as described for average S', maculata ; other males
black in ground colour, surface light blue-gray or gray, dark brown to black around macrotrichial insertions. Macrotrichia:
median marginals absent from T1 + 2; lateral marginals on T]+2 weak in 2 males and 4 females from “Orlando” series,
strong in others.
Genitalia (Fig. 61). Four examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus
bent very slightly downward from angle of basiphallus in 3 specimens, bent more sharply (as in S. maculata) in one
specimen; antero-ventral margin dentate (without hooks); apex broad in profile.
Female. As described for male except as follows.
Head (Fig. 43). Palpus longer and thicker than in males.
Preabdomen. Specimen from Lake Placid mottled black and yellow in ground colour, weakly vittate; other specimens
black in ground colour, surface light blue-gray or gray.
Geographical distribution (Fig. 84). - Known only from Florida and southern Georgia.
Chorological affinities. - This is the only Siphona species restricted to extreme southeastern
United States, and the only member of the S. maculata group recorded from that area. S.
multifaria is the only Siphona species known to be sympatric with this species.
Phylogenetic relationships (Fig. 103). - This species is plesiotypic in almost all states, so its
placement beyond species group cannot be clarified. It might be the sister species to S.
maculata , as inferred from phenetic similarity and zoogeographic analysis.
Paratypes. — Four males, 7 females. USA: Florida: Orlando, 3,15.111.46, F. Snyder, 2M, 4F; Wakulla Co.,
Ochlockonee R. St. Pk„ 29. IV. 70, W.W. Wirth, 1M; Highlands Co., Archbold Biological Station, 14.111.62, S.W. Frost,
1 M; Archbold Biological Sta., L. Placid, 10. IV. 66, R.W. Dawson, IF; Biscayne Bay, IF. Georgia: Liberty Co., St.
Catherines Island, 24-28. IV. 72, Thompson and Picchi, 1 F. (Deposited in AMNH, CNC, PSU, USNM and WSUP.)
Siphona intrudens (Curran)
Bucentes intrudens Curran, 1932: 14.
North American species of Siphona Meigen
289
Siphona intrudens : Sabrosky and Arnaud, 1965: 1064.
Type material examined. - Holotype, male, labelled: “Bucentes/ intrudens/ Curran <3 [red
label]”, “Castle Rock/ IV- 1 7- [ 1 9]08 Pa [Pennsylvania, USA]” “Accession]. 5642 A.J. Weidt
coll.”; “Bucentes/ intrudens/ Curran” (AMNH).
Allotype , female, same data as holotype except without fourth label (AMNH).
Recognition. - Small to average-sized eye, average-sized first flagellomere and short
proboscis. Most specimens with weak lateral marginal setae on T1+2, large tarsal claws and a
dark blue-gray preabdomen with dorsal vitta. A few specimens very similar to S. maculata , but
the U-shaped distiphallus (in anterior view), with 3-4 large hooks along antero-ventral margin,
is unique to S. intrudens. Three postsutural dorsocentral setae.
S. intrudens can be confused with either S. maculata or S. lurida. For discussion concerning
former, see “Recognition” under that species.
Specimens of S. lurida with a first flagellomere of average Siphona length are similar in
general appearance to S. intrudens. The dark colouration of these species, with blue-gray,
non-vittate preabdomens, contributes to their similarity and with other characteristics attests to
their close phylogenetic relationship.
S. intrudens specimens with strong lateral marginals on T1+2 can be separated from S.
lurida by the larger eye, presence of only 2-3 setae on vibrissal angle (besides vibrissa) and by
the unique structure of the distiphallus.
Description. — Specimens examined: 408 males, 338 females. Length: 3. 5-5. 5mm.
Male. Head (Fig. 20). Colouration: as described for S. maculata. Eye: small to average for genus, 0.703-0.817 head
height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle with 2 or 3
setae and several hairs, setae in most specimens 2 in number, 1 laterad or above vibrissa and 1 below. Antenna: first
flagellomere 0.474-0.593 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short,
1 .45-2.00 head height (Fig. 1 5).
Thorax. Colouration: dorsum average, blue-gray to blue with varied amounts of brown; 3 brown vittae not clearly
differentiated in most specimens ( S . intrudens generally with more blue on dorsum than S. maculata , and darker than S.
floridensis)', tegula testaceous to fuscous; wing yellowish to cinereous hyaline; legs average to darker than average, femur
of hindleg predominantly reddish-brown in few specimens. Macrotrichia: postsutural dorsocentrals, 68 specimens
examined, all with three [males and females]. Acropod: claws and pulvilli large in most specimens (Fig. 5), average in few.
Preabdomen (Fig. 7). Colouration: highly varied, from black in ground colour, surface predominantly blue-gray or
gray, to vittate condition described for S. maculata\ most specimens from western North America dark form, both dark
and light (vittate) forms well-represented in examined material from prairies to eastern North America. Macrotrichia:
median marginals absent from T]+2; lateral marginals on T,+2 in most specimens weak, slightly stronger than
surrounding hairs, in few specimens not differentiated, in few strong.
Genitalia (Fig. 62). Thirty-seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci.
Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin U-shaped in anterior view,
with several large, well developed hooks (larger than in S. maculata)\ in profile tapered to narrow apex.
Female. As described for male except as follows.
Head (Fig. 44). Palpus longer and thicker than in males in most specimens.
Preabdomen. Colouration: most specimens black in ground colour, surface predominantly blue-gray or gray; few
specimens vittate, as described for average specimen of female S', maculata.
Geographical distribution (Fig. 85). - Transcontinental species; records mostly south of
boreal forest, and particularly well recorded from northeastern United States.
Chorological affinities. - Among S. maculata group members, this species is evidently
parapatric or narrowly sympatric with S. lurida and S. pacifica. Also see notes under S.
maculata.
Phylogenetic relationships (Fig. 103). - This is a highly derived species, closely related to S.
lurida. A decision on whether these species are sister species awaits a cladistic analysis of
related (undescribed) Oriental species of Siphona.
See also notes under S. maculata.
Quaest. Ent., 1982, 18 (1-4)
290
O’Hara
Siphona lurida Reinhard
Siphona lurida Reinhard, 1943: 20-- Sabrosky and Arnaud, 1965: 1064.
Type material examined. - Holotype , male, labelled [lines not recorded]: “Rainier, Ore.
[Oregon, USA] III.3. [ 1 9] 30”; “R.E. Dimick Coll.”; “HOLOTYPE Siphona lurida Reinhard
[red label]”; “Siphona lurida Rnh R’43 [Reinhard det. label]” (CNC).
Allotype , female [abdomen missing], labelled: “Union Hills/ 11-20-1940/ R.E. Rieder,
col.”; “ALLOTYPE/ Siphona/ lurida/ Reinhard [red label]”; “Siphona/ lurida/ Rnh. R.
[Reinhard det. label]”; “PARATYPE/ Siphona/ lurida/ Reinhard [yellow label]” (FSCA).
Paratypes. — , 4 males, top two labels: “Union Hills/ Marion Co[unty], Ore. [Oregon, USA]/ 11-20-1940/ R.E.
Rieder, col.”; “Paratype/ Siphona/ lurida/ Reinhard [red label]” (2 in CNC, 1 each in FSCA, OSU).
Recognition. - Eye smaller than any other species, first flagellomere long in most specimens.
Proboscis short. Cluster of 5-10 setae on vibrissal angle besides vibrissa and hairs, and above
average number of hairs on parafacial and frontal plate (up to ca. 15). Dark coloured species,
preabdomen black in ground colour overall, apex of palpus infuscate in most specimens. Tarsal
claws large in most specimens. Wing cinereous hyaline. Lateral marginal setae on T1+2 strong.
Three postsutural dorsocentral setae. Adults are seen in very early spring to mid-summer; one
southern record from December.
A few specimens are very similar to S. intrudens, as discussed under “Recognition” of that
species.
S. lurida specimens with a very small eye and large first flagellomere are easily identified.
The holotype and paratypes are of this form, as are examined specimens of S. lurida from
Japan. It is uncertain how prevalent specimens with a short first flagellomere are, for the
species has been inadequately collected.
Description. — Specimens examined: 1 1 males, 9 females from North America; 1 male, 2 females from Japan.
Length: 4. 5-5. 5mm.
Male. Head (Fig. 21). Colouration: frontal vitta bright orange to reddish-brown; gena and lower parafacial orange or
yellow to gray, upper parafacial and frontal plate yellow or blue-gray to brown; orbital plate light to dark brown; scape,
pedicel and a portion of first flagellomere above aristal insertion orange to fuscous, rest of first flagellomere fuscous to
black; palpus yellow, apex infuscate in most specimens; proboscis reddish-brown to fuscous. Eye: very small, 0.627-0.677
head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia: number and length
of hairs on upper parafacial and fronto-orbital plate above average for genus in most specimens, up to 15 present; vibrissal
angle with tight cluster of 5-10 setae and several hairs in addition to vibrissa. Antenna: first flagellomere highly varied in
length, 0.508-0.677 head height (Fig. 14); length of aristomere 3 average for genus. Mouthparts: proboscis short,
1.45-2.00 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or blue with 3 clearly differentiated brown vittae in most specimens,
few specimens with dorsum predominantly brown (very close to S. intrudens)-, tegula fuscous; wing cinereous hyaline; legs
average to darker than average, femur of foreleg predominantly reddish-brown in few specimens. Macrotrichia:
postsutural dorsocentrals, 20 specimens examined, all with three [males and females]. Acropod: claws and pulvilli large in
most specimens, average in few.
Preabdomen. Colouration: not vittate; black in ground colour, surface varied, from predominantly blue-gray to
predominantly dark brown mixed with black with blue-gray restricted to bands on anterior third of T3, T4, and T5; dark
brown or black around macrotrichial insertions. Macrotrichia: median marginals absent from T1+2; lateral marginals on
T1 + 2 strong.
Genitalia (Fig. 63). Six examined, including one Japanese specimen. Apex of surstylus even with or extended slightly
beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin
V-shaped in anterior view, dentate; in profile tapered to rounded apex.
Female. As described for male except as follows.
Head (Fig. 41 ). Palpus longer and thicker than in males in most specimens.
Preabdomen. Colouration: black in ground colour; surface varied, from predominantly blue-gray or gray to
predominantly dark brown with or without narrow bands of blue-gray on anterior margins of T3, T4, and Ts.
Geographical distribution (Fig. 86). - Japan (new record) and western United States.
North American species of Siphona Meigen
291
Chorological affinities. - In North America, this species is apparently parapatric or
narrowly sympatric with S. maculata and the closely related S. intrudens. The range of S.
pacifica coincides with that of another S. maculata group member, S. lurida.
Phylogenetic relationships (Fig. 103). - Like S. intrudens , this species is highly derived. It
might be sister species to S. intrudens , as mentioned under that species.
Siphona pacifica new species
Holotype. - Male, labelled: “Dartford, WASH. [Washington, USA]/ Malaise trap”; “R.D.
Gray/ VI. 10-11 1969” (WSUP).
Allotype. - Female, labelled: “Dartford, Wash/ VII-5- 1 970”; “Malaise trap/ R.D. Gray”
(WSUP).
Derivation of specific epithet. - This species is named in recognition of its western
distribution.
Recognition. - Proboscis length and eye height average, first flagellomere short.
Preabdomen light blue-gray or gray, with or without dorsal vitta. Lateral marginal setae on
T , _|_ 2 weak or strong. Distiphallus bent slightly downward from angle of basiphallus, apex
broad in profile. Four postsutural dorsocentral setae.
This species has a unique set of characteristics that distinguish it from other species.
However, taken singly these features are unremarkable, and S. pacifica is therefore difficult to
recognize. It is placed in the S. maculata group because the first flagellomere is slender and
broadly rounded apically along ventral margin, lateral marginal setae on T)+2 are weak in
some specimens, and general colouration is typical of the group.
Description. - Length: 3.5-5.0mm.
Male. Head (Fig. 22). Colouration: frontal vitta orange, testaceous to reddish-brown; gena and parafacial white, light
yellow to light brown; fronto-orbital plate light brown to brown; scape and pedicel and in few specimens portion of first
flagellomere above aristal insertion yellow; first flagellomere fuscous to black; palpus yellow; proboscis reddish-brown.
Eye: size average to large for genus, 0.796-0.850 head height (Fig. 13), broad, widest at center, evenly rounded along
anterior margin. Macrotrichia: vibrissal angle with 1 seta above vibrissa (or laterad) and in most specimens 2 setae below,
and several short hairs. Antenna: first flagellomere short, 0.431-0.472 head height (Fig. 14); length of aristomere 3 average
for genus. Mouthparts: proboscis length average for genus, 1.87-2.15 head height ( Fig. 1 5).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in most specimens; tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals,
four in all specimens. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: varied, vittate in few specimens; most specimens predominantly or entirely black in ground
colour, few specimens with yellow ground colour laterally on T)+2 and T3 or less; surface light coloured, areas of black
ground colour light blue-gray or gray in surface colour, brown to black around macrotrichia! insertions; areas yellow in
ground colour with surface yellow or light brown. Macrotrichia: median marginals absent from T]+2; lateral marginals on
T|+2 differentiated from marginal hairs, weak or strong.
Genitalia (Fig. 64). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus bent slightly downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge
straight, apex broad.
Female. As described for male except as follows.
Head (Fig. 45). Length of palpus slightly longer than in males.
Preabdomen. Colouration: not vittate; black in ground colour, surface blue-gray or gray.
Geographical distribution (Fig. 87). - Recorded from several localities in California and
one in Washington.
Chorological affinities. - The western distribution of this species is very similar to that of S.
lurida. Two other species of the S. maculata group, S. maculata and S', intrudens , are
evidently parapatric or narrowly sympatric with this species.
Phylogenetic relationships (Fig. 103). - S. pacifica is interpreted as distantly related to
most other North American members of the S. maculata group. This species possesses the
Quaest. Ent., 1982, 18 (1-4)
292
O’Hara
synapotypic feature of the group - a slender, pointed first flagellomere - as well as the
symplesiotypies, weak lateral marginal setae on T1+2 and a relatively short proboscis.
Paratypes. — Seven males, 4 females. USA: California: Sugar Loaf Mt., 28.V.46, J. Sperry, 1M; Marin Co., Mill
Valley, IX. 51, E.S. Ross, 1M; Placer Co., Brockway, VII. 41, G.E. Bohart, 1M. Washington: Spokane Co., Dartford, 6-9,
20-31. VI, 2-3. VII, 2-3.VIII.69, 5.VII.70, R.D. Gray, 4M, 3F; ibid., 23.VI.70, IF. (Deposited in CAS, USNM and
WSUP.)
The S. cristata group
Characteristics varied; most species with an average to long proboscis, average-sized eye,
broad first flagellomere rounded or slightly truncate apically along ventral margin (not
distinctly pointed), and an anterior indentation on the distiphallus. The three predominantly to
entirely yellow North American Siphona species belong to this group.
Siphona cristata (Fabricius)
Stomoxys cristata Fabricius, 1805: 281.
Siphona cristata: Mesnil, 1964: 861 (redescription).— Curran, 1932: 13 (in key; probable misidentification).
Siphona chaetolyga Rondani, 1865: 194.- Herting, 1969: 192.
Siphona palpina Zetterstedt, 1859: 6064.- Mesnil, 1964: 861.
Mesnil (1964) and Crosskey (1976b) followed earlier authors and listed Siphona oculata
Pandelle as a synonym of S. cristata. Herting recently examined the type of S. oculata and
found it conspecific with Siphona pauciseta Rondani (Herting, 1978: 6).
Type material examined. - Stomoxys cristata Fabricius, HOLOTYPE, female, labelled:
“TYPE [red label]”; “S. cristata/ [locality indecipherable; Denmark] [handwritten]”;
“Holotype 2/ Siphona/ cristata (F.) [handwritten]”/ det. “Stig Andersen” (ZMUC).
I cannot with certainty confirm that the type of S. cristata is conspecific with the North
American species I identify as such. The type is a female with features characteristic of (my) S.
cristata and S. multifaria. Therefore my interpretation of S. cristata is based primarily on
Mesnil’s description (1964) and male and female specimens determined by Drs. Mesnil and
Herting.
Recognition. - Eye height and first flagellomere length average, proboscis longer than
average. Colouration highly varied; in few specimens thoracic pleura yellow in ground colour,
dorsum dark in ground colour, rest of specimens with thorax predominantly dark in ground
colour. Preabdomen of both sexes varied, from entirely yellow, to yellow with narrow median
vitta, to average vittate with predominantly tawny surface colour. Very few specimens with a
single or 1 pair median marginal setae on T1+2 (3% each). Distiphallus bent sharply downward
from angle of basiphallus, in most specimens with anterior edge curved inward ca. 0.33 of its
length from ventral edge. Postsutural dorsocentral setae three (35%) or four (65%).
Due to varied nature of this species, identification can be difficult. A few specimens
resemble S. lutea (as discussed under “Recognition” of that species), but do not pose a serious
identification problem.
Separation of S. cristata and S. pisinnia is discussed under the latter species.
Specimens of S. cristata can be very difficult to separate from those of S. multifaria , for
they are similar in most characteristics. Colouration is useful, but not always trustworthy. The
yellow form of S. cristata is very distinct from S. multifaria , but specimens with vittate
preabdomen are not always readily separable from S. multifaria specimens.
In Edmonton, Alberta, both species occur at the same time in the same place. S. cristata
specimens are slightly larger and have a tawny, vittate preabdomen. The preabdomen of S.
North American species of Siphona Meigen
293
multifaria specimens is yellow laterally (not tawny as in S. cristata ) and has a narrow brown
pruinose vitta medially, overlying a indistinct light blue-gray vitta. Differences in the
distiphallus are recognizable, but do not hold for all specimens.
Since both S. cristata and S. multifaria are quite varied, the differences seen in Edmonton
populations are not suitable for species discrimination in all areas. For this reason I have not
separated these species in the key. Most male specimens should be identifiable on the basis of
the foregoing remarks, though female specimens will not meet with equal success.
Description. — Specimens examined: 78 males, 112 females from North America, ca. 10 European specimens.
Length: 4. 5-5. 0mm.
Male. Head (Fig. 24). Colouration: as described for S', lutea except for antenna and proboscis; scape, pedicel and
portion of first flagellomere above aristal insertion yellow in most specimens, fuscous in few; rest of first flagellomere
testaceous to fuscous, in few specimens black; proboscis brown to reddish-brown. Eye: size average to above average for
genus, highly varied, 0.754-0.855 head height (Fig. 13), slender to broad, widest at, or slightly above, center, narrowed or
rounded below. Macrotrichia: average for genus. Antenna: first flagellomere 0.547-0.672 head height (Fig. 14), resembling
first flagellomere of S. lutea but more varied in length and shape; first flagellomere broad, varied from truncate to broadly
rounded along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length above average for
genus, highly varied, 2.25-3.00 head height (Fig. 1 5).
Thorax. Colouration: black in ground colour in most specimens, light brown in ground colour on pleura in few
specimens; dorsum from yellow or light brown finely and evenly mixed with blue-gray or gray to entirely yellow, golden or
brown; 3 brown or golden vittae dorsally in very few specimens; tegula testaceous; wing yellowish hyaline; legs average.
Macrotrichia: postsutural dorsocentrals, 20 specimens with three, 37 with four [males and females]. Acropod: claws and
pulvilli average in size.
Preabdomen. Colouration: one form predominantly yellow in ground colour, with or without narrow dark vitta, surface
yellow to bright tawny; pattern varied from yellow form to average Siphona vittate form, with T5, most of T4, triangular
area on T3, and T,+2 medially black in ground colour, surface gray to brown; dark brown to black around macrotichial
insertions; rest of preabdomen yellow in ground colour, surface yellow to tawny. Macrotrichia: most specimens without
median marginals on T1+2, few with 1 seta on one side (6 specimens) or 1 pair (5 specimens); lateral marginals on T1+2
strong.
Genitalia (Fig. 66). Thirty-two examined, including 2 from European specimens. Apex of surstylus even with or
extended slightly beyond apex of cerci. Aedeagus: distiphallus bent sharply downward from angle of basiphallus;
antero-ventral margin dentate; in profile anterior edge varied, in most specimens indented ca. 0.33 distiphallus length from
ventral edge, straight in few specimens; apex broad in profile.
Female. As described for male except as follows.
Head (Fig. 46): Palpus longer and thicker than in males in most specimens.
Geographical distribution (Fig. 95). - Wide ranging in the Palearctic and Nearctic
Regions, though sparsely recorded from latter over much of its apparent range.
Habitat. - I have collected specimens of S. cristata , S. lutea and S. multifaria on flowers of
the composite, Erigeron philadelphicus L., in river valley of the North Saskatchewan R.,
Edmonton, along shaded bank of a small stream in vicinity of poplar and scattered spruce.
Chorological affinities. - In North America this wide ranging species is sympatric with the
closely related species S. multifaria and S. lutea , in addition to most other Siphona species of
America north of Mexico. As noted above, specimens of S. multifaria and S. lutea have been
caught in the same locality as those of S. cristata. In southwestern New Mexico, specimens of
S. pisinnia were collected with specimens of this species.
Phylogenetic relationships (Fig. 104). - Close to S. multifaria among North American
species, though the sister species might be a Palearctic species.
Siphona multifaria new species
Holotype. - Male, labelled: “MI. 87 [also known as km. 140.5], Y.T. [Yukon Territory,
Canada]/ Dempster Hwy ./ 8-12.VII.1973/ G.&D.M. Wood” (CNC).
Allotype. - Female, same data as holotype except date 16-17.VII.1973 (CNC).
Quaest. Ent., 1982, 18 (1-4)
294
O’Hara
Derivation of specific epithet. - From the Latin multifarius, meaning “having great
variety”. S. multifaria is so named because it is a widely distributed species of varied
appearance, being particularly difficult to identify because of its resemblance to S. cristata.
Recognition. - Eye height and length of first flagellomere average, proboscis longer than
average. Wing yellowish hyaline in most specimens, cinereous in few. Male preabdomen varied,
in most specimens light coloured, vittate, with a very narrow brown pruinose vitta (unassociated
with vitta produced by ground colour), few specimens predominantly black in ground colour.
Female preabdomen darker than in male, not vittate. Very few specimens with 1 median
marginal seta on one side on T1+2 (2%). Distiphallus bent sharply downward from basiphallus,
anterior edge slightly or not at all curved inward ca. 0.33 of its length from ventral edge. Most
specimens (65%) with three postsutural dorsocentral setae.
Most difficult to separate from S. cristata , as discussed under “Recognition” of that species.
S. multifaria and S. pisinnia are narrowly sympatric in southwestern United States.
Specimens from that region should be checked carefully against data in the Hubbs-Hubbs
diagrams, for colouration is essentially the same in both species, and differences between
distiphalli are not diagnostic. The data base for S. multifaria and S. pisinnia is large enough
that measurements cited in key, especially with reference to male specimens, should be
representative of the two species.
Description. - Length: 3. 5-5. 0mm.
Male. Head (Fig. 25). Colouration: frontal vitta orange or yellow to testaceous; gena and lower parafacial white to
light yellow; upper parafacial and fronto-orbital plate light yellow to brown, darkest on orbital plate, in few specimens
fronto-orbital plate bright yellow (these specimens resembling S. cristata ); scape, pedicel, and in few specimens portion of
first flagellomere above aristal insertion yellow to testaceous, in few specimens reddish-brown or fuscous; first flagellomere
reddish-brown, fuscous or black; palpus yellow; proboscis light brown to reddish-brown or fuscous. Eye: size average to
above average for genus, highly varied, 0.755-0.854 head height (Fig. 13), slender, widest at center, evenly rounded along
anterior margin. Macrotrichia: vibrissal angle with 2 or 3 setae and several hairs, setae in most specimens 2 in number
(average for genus), 1 laterad or above vibrissa and 1 below. Antenna: first flagellomere highly varied in length,
0.531-0.675 head height (Fig. 14); shape of first flagellomere varied, as described for S. cristata ; length of aristomere 3
average for genus. Mouthparts: proboscis length above average for genus, highly varied, 2.31-3.02 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula testaceous; wing yellowish hyaline in most specimens, cinereous hyaline in few; legs
average. Macrotrichia: postsutural dorsocentrals, 68 specimens with three, 37 with four [males and females]. Acropod:
claws and pulvilli average-sized.
Preabdomen. Colouration: faintly or clearly vittate; most specimens black in ground colour on T5, most or all of T4,
and medially on T]+2 and T3, surface blue-gray, gray or light brownish-gray; dark brown to black around macrotrichial
insertions; surface of most specimens with a faint or distinct narrow median brown pruinose vitta, narrower than vitta
formed by black ground colour; areas yellow in ground colour with surface yellow or light brown; in few specimens black
ground colour more extensive, covering most of T3, all of T4 and T5, with wide vitta T1+2. Macrotrichia: most specimens
without median marginals on T1+2, 7 with 1 on one side (1 specimen from Labrador, 6 from western North America);
lateral marginals on T, +2 strong.
Genitalia (Fig. 67). Fifty-six examined. As described for S. cristata , except anterior edge of distiphallus in profile
straight or slightly indented in most specimens, very few specimens with indentation as strong as in average S. cristata.
Female. As described for male except as follows.
Head (Fig. 47). Palpus longer and thicker than in males in most specimens.
Preabdomen. Colouration: darker than male; most specimens not vittate, black in ground colour except laterally on
T, +2; surface blue-gray, mixed with brown, in few specimens very light, in few very dark.
Geographical distribution (Fig. 93). - Very widely distributed throughout eastern and
western North America; records absent for central plains.
Habitat. - See S. cristata.
Chorological affinities. - Due to the ubiquitous nature of this species, it is sympatric with
all other Siphona species of America north of Mexico. Specimens of S. cristata and S. lutea
have been caught together with those of this species.
North American species of Siphona Meigen
295
Phylogenetic relationships (Fig. 104). - Because this species is wide ranging in North
America and recorded from northern localities, I consider its presence in the Palearctic Region
likely. Therefore its sister species is not necessarily a species of Nearctic distribution. S.
cristata is evidently the closest Siphona species to S. multifaria in North America.
Paratypes. — One hundred and eigthy-nine males, 128 females. CANADA. Alberta: George L., 53°57'N
114°06'W, 13-17.V1II.66, P. Graham, 1 M; Edmonton, 19.V.47, E.H. Strickland, IF; ibid., 1932, O. Bryant, 1M; ibid.,
29.V1.80, J.F. Landry, 2M; ibid., 7,8.V11.80, 30.VI, 3,6.VII.81, J.E. O'Hara, 13M; Mildred L„ 57°03'N 111°35'W,
11-15.V1I.79, J. Ryan and G. Hilchie, 1M; Banff, 3.VI1.49, E.H. Strickland, IF; Banff N.P., 8. VII. 55, R. Coyles, 1 M;
Frank, 15.VI.62, W.R.M. Mason, 1 M; Medicine Hat, 22. VI. 74, J.F. McAlpine, 1 M; Cypress Hills, 10.VIII.49, E.H.
Strickland, 3F. British Columbia: Summit L., mi. 392, Alaska Hwy., 4500', 29-30. VI, 2-4,1 1-14. VII. 59, R.E. Leech, 2M,
IF; ibid., 4200', 21, 31. VII, 19-21 VIII.59, R.E. Leech, 6M, IF; ibid., 19-21. VIII. 59, E.E. MacDougall, 2F; Lakelse L.
bog, s. ofTerrace, 1 1,27.VI.60, B. Heming, 2M; ibid., 11.VI1.60, C.H. Mann, 1M; Lac La Hache, 15.VI1.73, H.J.Teskey,
1M; Keremeos, 18.VI.23, C.B. Garrett, 1M; Atlin, 2200', 3.VII.55, H.J. Huckel, 1M; Horsefly, 14.VII.73, H.J. Teskey,
1M; Ketchum L„ 58°22'N 131°45'W, 3600', 23.VIII.60, W.W. Moss, IF; ibid., R. Pilfrey, 2F; McQueen L„ 10 mi. n.
Kamloops, 2. VII. 73, H.J. Teskey, IF. New Brunswick: Kouchibouquac N.P., 10, 24. VI. 77, J.R. Vockeroth, 3M, 6F; ibid.,
7. VII. 77, J.F. McAlpine, 1M; St. Andrews, 3. VIII. 57, G.E. Shewed, 1M. Newfoundland: St. John's, 21.VII.67, J.F.
McAlpine, IF; Cartwright, Labrador, 19.VII.55, E.E. Sterns, IF. Northwest Territories: Norman Wells, 8, 16. VIII. 69,
G.E. Shewed, 2F. Ontario: Maynooth, 9. VIII. 74, D.M. Wood, 1M; Mer Bleu, 5 mi. e. Ottawa, 3,11. VI. 66, D.D. Munroe,
2M; Mink L., nr. Maynooth, 27. VII. 55, J.F. McAlpine, 1M; Forestry Station, Petawawa, 28.V.59, J.R. Vockeroth, 1M;
Petawawa, 28.V.59, J.R. Vockeroth, 3F; Go Home Bay, 8 mi. w. Bala, 22.V.59, J.G. Chillcott, 1M; Foxboro, 10.V.63,
A.F. Johnson, 1M. Quebec: Gatineau Co., Masham Twp., 25-31. VII. 74, D.M. Wood, 2M; Lac Larouche, La Verendrye
Pk„ 30.VII.72, D.M. Wood, 2M, 2F; Mistassini Post, 1 8. VII. 56, J.R. Lonsway, 1M;4 mi. n. Eardley, 20,25. VI 1 1.7 1 , D.M.
Wood, 1M, 3F; Kam. Co., Parke Reserve, 28.VII.57, G.E. Shewed, 1M; Duncan L., nr. Rupert, 28.VI1, 9.VIII.70, 31. VII,
14. VIII. 71, J.F. McAlpine, 3M, 2F. Yukon Territory: 14 mi. e. Dawson, 1300', 4. VIII. 62, P.J. Skitsko, IF; 17 km. wnw.
Burwash Flats, 1250m, 10. VI 1.80, Wood and Lafontaine, IF; mi. 40, Dempster Hwy., 1-6. VI 1.73, G. and D.M. Wood,
1M; mi. 87, Dempster Hwy., 27-30.VI, 1-12, 18-27. VII, 4-8.V1II.73, G. and D.M. Wood, 37M, 23F; mi. 51, Dempster
Hwy., 18-27.VII.73, G. and D.M. Wood, 1 M; km. 140.5, Dempster Hwy., 900m, 27-29.VI1, 1.VIII.80, Wood and
Lafontaine, 1 M, 7F; km. 1 55, Dempster Hwy., 950m, 29.V1-3.VII.80, Wood and Lafontaine, 2M, 3F.
USA: Alabama: Eufaula, 18. VI. 54, R.L. Fischer, IF. Alaska: Richard Hwy., Donnelly Dome, 25. VI. 51, W.R.M.
Mason, 1 M; Cape Thompson, 1 .VI 1 1.61 , R. Madge, 1 F; Unalakleet, 1 7. VI 1, 1 1 .VI 11.61 , B.S. Heming, 2F; Kodiak, IX. 17,
J.S. Hine, IF; Douglas, 4. VIII. 01, E. Jenne, IF; Katmai, VIII. 17, J.S. Hine, 1M; King Salmon, Naknek R., 10. VII,
10.VIII.52, W.R. Mason, 1 M, IF; ibid., 2-3. VIII. 52, J.B. Hartley, 3F; Naknek, 18. VII, 8. VIII. 52, W.R. Mason, 3F.
Arizona: Sunnyside Canyon, Huachuca Mts., 9. VI 1.40, D.E. Hardy, 2F; Cochise Co., Southwestern Research Station, 8
km. w. Portal, 1650m, 21.VII1.62, 2M; ibid., 27.X.64, IF; ibid., 3.XI.64, V.D. Roth, IF; ibid., 14.VIII.65, 2M; ibid.,
9. VII, 27.VIII.65, V.D. Roth, 2M; ibid., 5-25.IX.65, C.W. Sabrosky, 3M, IF; ibid., 23.IX.66, P.H. Arnaud, Jr., 1M; ibid.,
26. IX. 66, V.D. Roth, IF. California: Mono Co., Leavitt Meadow, 7200', 12. VIII. 63, H.B. Leech, IF; Trinity Co., Butter
Creek, 3450', ca. 12 mi. se. Hyampom, 21, 22. VII. 68, H. Leech, 4M. Colorado: Mt. Evans, Doolittle Ranch, 9800',
22,23. VII, 3. VIII. 61, J.G. Chillcott, 3M; ibid., 3.VIII.61, W.R.M. Mason, 4M, IF; ibid., C.H. Mann, 1M; ibid., B.H.
Poole, 1M, IF; Mt. Evans, Echo L., 10,600', 12. VIII. 61, C.H. Mann, IF; Boulder, 1922, 1M; Pingree Pk., 17. VIII. 32, IF;
Mesa Co., Glade Pk., 8. VI 1.53, Gurney, IF. Florida: Inverness, Robertson, 1M; S. Florida, Robertson, 2M, 2F; Lake Co.,
Paisley, 18.11.66, C.L. Felshaw, 1M. Georgia: Dallas, 2. VI. 40, P.W. Fattig, IF; Lumpkin Co., 15mi. nw. Dahlonega,
25. VI. 69, F. Santana, 1M; Oconee Co., 4mi. s. Farmington, 20.111.75, B.F. Freeman, 1M. Idaho: Newman L., 16.V.25,
A.L. Melander, IF; Viola, 26. VI. 12, J.M. Aldrich, IF. Massachusetts: Chester, 3. VIII, C.W. Johnson, 1M; Essex,
20. VII. 11, IF; Cuttyhunk Is. and Elisabeth Is., 21. VIII. 71, C.T. Parsons, IF. Michigan: Thayers, 16.V.55, J.R.
Vockeroth, IF; Montcalm Co., Flat River Game Area, 14.V.55, R. L. Fischer, 1M; Emmet Co., 27.V.60, R. and K.
Dreisbach, 6M; Charlevoix Co., 31.V.60, R. and K. Dreisbach, 3M, IF; Iron Co., 26. V 1 1 1.59, R. and K. Dreisbach, 1M;
Hillsdale Co., 21 .V.60, R. and K. Dreisbach, 1 M, 2F; Saginaw Co., 24.V.60, R.R. Dreisbach, 1 M. Montana: Granite Co.,
Elkhorn Ranch, 9 mi. s. Clinton, 3800', 26.VI.71, J.R. Powers, 1M. New Hampshire: Flume, n. Woodstock, 19. VI 1.35,
Blanton and Borders, IF; Franconia, 1M; Halfway House, to Gorham, 20. VII. 35, Blanton and Borders, 1M; Gorham,
19. VIII. 56, W.A. Drew, IF; ibid., R.W. Hodges, 1M; ibid., 14.VI1I.58, J.R. Vockeroth, 1M. New Jersey: Avalon, 8.VI,
2M; ibid., 30.VI, IF. New York: Lake Placid, 2000', 19.VII.62, J.R. Vockeroth, 3M, IF; ibid., J.G. Chillcott. 3M, IF;
Keene Valley, 10.V111.16, E.L. Diven, 1M, IF; Dug Mt., 8.VII1.12, D.B. Young, 1M; Olean, 5. IX. 13, IF; Babylon, L.I.,
9. VI. 35, 28.VIII.36, Blanton and Borders, 8M, 12F; ibid., 5.VIII.37, F.S. Blanton, IF. North Carolina: Highlands,
3-5000', IV. 36, R.C. Shannon, 1M; ibid., 3800', 29.V.57, J.R. Vockeroth, IF; Cumberland Co., Fort Bragg, 16.V,
27.IX-3.X.67, J.D. Birchim, 1 M, IF. Oregon: Deschutes Co., Meadow Cpgd., Bend, 3840', 29.VII.70, P.H. Arnaud, Jr.,
1M. Pennyslvania: West Co., Pitcairn, 17.V.65, J.G. Chillcott, 1M. Tennessee: Gatlinburg, 22. VI. 40, R.C. Osburn, 1M.
Utah: Grand Co., Warner Ranger Station, 28 mi. ese. Moab, 9200', 2. V 1 1 1 .60, F., P. and B. Rindge, 1M. Vermont:
Chittenden, Rutland, 1-15. VIII. 16, J. Bequaert, 3M; Norwich, 7. VII, C.W. Johnson, IF. Washington: Glacier Pk.,
Avalanche L., 14. VII. 35, A.L. Melander, 1 M; Asotin Co., Fields' Spr. St. Pk., 15. VI. 72, W.J. Turner, 1M; Yakima Co., 8
mi. sw. Tieton R.S., Bear Creek, 16, 28. VII. 72, W.J. Turner and W.B. Garnett, 7M. Wisconsin: Polk Co., VII, Baker, IF.
Quaest. Ent., 1982, 18 (1-4)
296
O’Hara
Wyoming: Sweetwater Co., (Old) Hwy. 187, 11.5 mi. s. Eden, 23. VIII. 65, H.B. Leech, 1M. (Deposited in AMNH,
CAS, CNC, CUI, FSCA, INHS, JEOH, MCZ, MSU, OUCO, UASM, UGA, UGG, UKL, USNM, and WSUP.)
Siphona lutea (Townsend)
Crocuta lutea Townsend, 1919: 584.
Siphona lutea : Sabrosky and Arnaud, 1965: 1064.
Siphona tenuis Curran, 1933: 10.-- Sabrosky and Arnaud, 1965: 1064. New synonymy.
Notes about synonymy. - Curran did not include S. lutea (Townsend) in his 1933 key to
adults of North American Siphona species. This suggests he was unaware of Townsend’s
description of this species when he described S', tenuis in 1933. I have examined the types of
both nominal species and regard them conspecific.
Type material examined. - Crocuta lutea Townsend, 2 syntypes. One female specimen here
designated LECTOTYPE, labelled: “TD 4519 [handprinted]"; “Franconia/ NH [New
Hampshire, USA]’’; “CHT Townsend/ coll"; “21. VII, [19] 1 5“[handprinted]; “Type/ No.
U.S.N.M. [red label; no number]’’; “Crocuta/ lutea/ T. [handprinted]/ Det CHTT"
(USNM). My lectotype label “LECTOTYPE/ Crocuta/ lutea Tnsd./ O’Hara designation/
Selected 1982 [red bordered, handprinted label]” has been attached to this specimen. One
paralectotype, female, same locality data as lectotype except collected 20.VII.1915 (USNM).
Siphona tenuis Curran, HOLOTYPE, male [left wing, 4 legs missing], labelled: “Siphona/
tenuis/ Curran 6 [red type label]’’; “Timagami, Ont. [Ontario, Canada]/ 1. VIII. 1932/
A.W.A. Brown“; “Siphona/ tenuis/ Curran” (AMNH).
Recognition. - Particularly distinct because of yellow ground and surface colour overall.
Most specimens examined (78%) with R4+5 setulose beyond crossvein r-m on at least one side.
Similarly, Rj distally with 1-2 setulae on at least one side in half the specimens (53%). Most
specimens (94%) with four postsutural dorsocentral setae.
Shares with S. oligomyia the synapotypic character state of yellow ground and surface
colouration. Differs from S. oligomyia in possessing a shorter proboscis and four postsutural
dorsocentrals and lacking a modified male sternum 5.
The only other species resembling S. lutea is S. cristata. The preabdomen of most specimens
of S. cristata is yellow with a thin dark median vitta; in few specimens it is darker, or entirely
yellow as in S. lutea. The darker thorax of S. cristata serves to separate specimens of that
species from S. lutea where abdominal colouration is ambiguous. Also, most specimens of S.
lutea are setulose on R4+5 beyond crossvein r-m, unlike other North American species.
Description. — Specimens examined: 41 males, 39 females. Length: 4. 0-5. 5mm.
Male. Head (Fig. 26). Colouration: frontal vitta orange or yellow to testaceous; gena and parafacial white to yellow;
fronto-orbital plate yellow, bright yellow to gold; scape, pedicel, palpus and proboscis yellow; first flagellomere yellow to
testaceous. Eye: size average for genus, 0.760-0.821 head height (Fig. 13), slender, widest at center, in most specimens
evenly rounded along anterior margin, in few slightly narrowed below center. Macrotrichia: average for genus. Antenna:
first flagellomere 0.561-0.643 head height (Fig. 14), broad and slightly truncate; length of aristomere 3 average for genus.
Mouthparts: proboscis slightly shorter than average for genus, 1.83-2.14 head height (Fig. 15).
Thorax. Colouration: yellow in ground colour and pruinosity; tegula yellow; wing yellowish hyaline; femora and tibiae
yellow. Macrotrichia: postsutural dorsocentrals, 2 specimens with three, 31 specimens with four [males and females];
wing, 32 males and females examined, R4+5 setulose beyond crossvein r-m on at least one side in 25 specimens, R, with at
least 1 setula on apical third on at least one side in 17 specimens, CuA, with 1 to 3 setulae in 4 specimens. Acropod: claws
and pulvilli average-sized (Fig. 6).
Preabdomen. Colouration: yellow in ground and surface colour; few specimens with red or dark areas posteriorly,
where dark coloured internal structures are visible through translucent exoskeleton. Macrotrichia: median marginals
absent from T, +2; lateral marginals on T, +2 strong.
Genitalia (Fig. 68). Nine examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus
bent sharply downward from angle of basiphallus in most specimens; antero-ventral margin dentate; in profile anterior
edge slightly indented ca. 0.33 distiphallus length from ventral edge in most specimens, straight in few; apex broad or
North American species of Siphona Meigen
297
narrow in profile. Sternum 5 (Fig. 10): average for genus.
Female. As described for male except as follows.
Head (Fig. 48). Palpus longer and thicker than in males in most specimens.
Geographical distribution (Fig. 89). - Transcontinental, in narrow belt south of boreal
forest.
Habitat. - Label data indicates specimens of this species have been collected from
sphagnum bogs. Also see S. cristata.
Chorological affinities. - This species is evidently parapatric with sister species S.
oligomyia. The range of this species is broadly sympatric with ranges of closely related S.
cristata and S. multifaria , as well as with ranges of S. hokkaidensis, S. maculata, S. intrudens
and S. medialis.
Phylogenetic relationships (Fig. 104). - This species and S. oligomyia are interpreted as
sister species on the basis of their highly derived colouration and setula on bend of R,. Together
I interpret these species as the sister lineage to the S. cristata - S. multifaria lineage.
Siphona oligomyia new species
Holotype. - Male, labelled: “Keremeos, B.C. [British Columbia, Canada]/ 20. VI 1923/
C.B. Garrett” (CNC). Genitalia in microvial on pin below specimen.
Derivation of specific epithet. - Formed from oligos and myia , Greek for “few” and “fly”
respectively. This combination was chosen because the type-series is comprised of but two
specimens, collected more than 1000 kilometers apart.
Recognition. - Yellow ground and surface colour as in S. lutea. This is the only North
American species with antero-lateral arms of sternum 5 hooked inward. R, distally with one
setula on only one side in both specimens. Three postsutural dorsocentral setae. Female of
species unknown.
This species is compared with S. lutea under “Recognition” of that species.
The characterization of S. oligomyia remains incomplete until more specimens are collected
and examined and female of the species recognized. In particular, usefulness of proboscis
length as a means for distinguishing S. oligomyia from S. lutea needs testing, and wing setular
characteristics require additional data to establish trends.
Description. - Length: 4.0-4. 5mm.
Male. Head (Fig. 27). Colouration: as described for S. lutea. Eye: size average for genus though smaller than eye of S.
lutea , 0.741-0.750 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin. Macrotrichia:
average for genus. Antenna: first flagellomere 0.574-0.596 head height (Fig. 14), broadly rounded distally along ventral
margin; length of aristomere 3 average for genus. Mouthparts: proboscis longer than in S. lutea, average for genus,
2.30-2.35 head height (Fig. 15).
Thorax. Colouration: yellow in ground colour and pruinosity; tegula yellow; wing yellowish hyaline; femora and tibiae
yellow. Macrotrichia: postsutural dorsocentrals, both specimens with three; wing: R4 + 5 not setulose beyond crossvein r-m
(ie. average, cf. S. lutea); R, with single setula on apical third on one wing only in both specimens. Aeropod: claws and
pulvilli slightly larger than average.
Preabdomen. Colouration: yellow in ground and surface colour. Macrotrichia: median marginals absent from T]+2;
lateral marginals on T, +2 strong.
Genitalia (Fig. 69). Both holotype and paratype examined. Apex of surstylus ca. even with apex of cerci. Aedeagus:
distiphallus slightly bent downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior edge
convex, not indented (as it is in most S. lutea); apex broad in profile. Sternum 5 (Fig. 1 1): antero-lateral margins hooked
inward.
Female. Unknown.
Geographical distribution (Fig. 90). - Known only from Lake Tahoe, Calif., and Keremeos,
B.C.
Quaest. Ent., 1982, 18 (1-4)
298
O’Hara
Chorological affinities. - Though this species has been inadequately collected, it is
apparently parapatric with sister species S. lutea. Among other S', cristata group members, it is
sympatric with S. cristata and S. multifaria. Similar in distribution are S. lurida and S.
pacifica of the S. maculata group.
Phylogenetic relationships (Fig. 104). - This species is sister species to S. lutea, as
mentioned under that species.
S. cuthbertsoni Curran (described from Salisbury, Zimbabwe) and Siphonopsis species
have the antero-lateral arms of male sternum 5 hooked inward, as in S. oligomyia. S.
oligomyia is not closely related to S. cuthbertsoni , and neither is closely related to Siphonopsis
species.
Paratype. — One male. USA: California: Lake Tahoe, 20. Vi. 1 5, A.K. Fisher (USN M).
Siphona pisinnia new species
Holotype. - Male, labelled: “USA N.M. [New Mexico] Grant Co./ Gila Nat. For. Cherry/
Ck. Campgrd. 21km.n./ Silver City 2250m./ 3-4.VIII.1980/ J.E. & W.M. O’Hara” (CNC).
Allotype. - Female, same data as holotype (CNC).
Derivation of specific epithet. - To my knowledge “pisinnia” has no classical meaning, and
was chosen as an arbitrary combination of letters.
Recognition. - Eye height and proboscis length average, first flagellomere long and broad in
most specimens (the most distinctive external feature). Male preabdomen light coloured,
faintly vittate. Female preabdomen darker, not vittate. Distiphallus slightly bent downward
from angle of basiphallus, shape not very distinctive; anterior edge straight or slightly .rounded,
apex broad or narrow in profile. Three postsutural dorsocentral setae in most specimens (87%).
In Mexico male and female specimens are easily distinguished by head characteristics. Male
genitalia of S. pisinnia are quite different from those of Mexican species.
In southwestern United States S. cristita specimens have been caught with specimens of S.
pisinnia. Differences in colouration, eye size, first flagellomere and proboscis lengths will in
most instances separate specimens of these species. Most S. cristita specimens are slightly
larger than those of S. pisinnia.
Very similar to S. pisinnia is S. multifaria. Differentiation between the two is discussed
under “Recognition” of S. multifaria.
Description. - Length: 3. 0-5. 0mm.
Male. Head (Fig. 23). Colouration: frontal vitta yellow to brown or reddish-brown; gena and lower parafacial white to
light yellow; upper parafacial and fronto-orbital plate light yellow to brown, darkest on orbital plate; scape and pedicel
testaceous to fuscous; first flagellomere fuscous to black; palpus yellow; proboscis reddish-brown or fuscous. Eye: size
average for genus, 0.725-0.811 head height (Fig. 13), slender, widest at center, evenly rounded along anterior margin.
Macrotrichia: average for genus. Antenna: first flagellomere highly varied, in most specimens long and broad, 0.577-0.740
head height (Fig. 14), in most specimens rounded distally along ventral margin, in few specimens slightly truncate; length
of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 1.87-2.32 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray, finely mixed with brown in most specimens, 3 brown vittae
visible in few 'specimens; tegula testaceous to fuscous; wing yellowish hyaline; legs average. Macrotrichia: postsutural
dorsocentrals, 53 specimens with three, 8 with four [males and fema-les]. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: faintly vittate; most specimens black in ground colour on T5, T4 and medially on T)+2 and
T3, rest of preabdomen yelldw in ground colour; surface blue-gray, gray or light brownish-gray on areas of black ground
coloiir, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow or light
browp; in few specimens black ground colour extensive, over most of T3, all of T4 and T5, with broad vitta on T1+2.
Genitalia (Fig. 65). Thirteen examined. Apex of surstylus even with or extended slightly beyond apex of cerci.
Aedeagus: distiphallus slightly bent downward from angle of basiphallus; antero-ventral margin dentate; in profile anterior
edge straight or slightly rounded; apex broad or narrow in profile.
North American species of Siphona Meigen
299
Female. As described for male except as follows.
Head (Fig. 49). Palpus longer and thicker than in males in most specimens.
Preabdomen. Colouration: darker than male, not vittate; black in ground colour in most specimens, yellow laterally
on T, _j_2 *n few specimens; surface blue-gray or gray, mixed with light brown.
Geographical distribution (Fig. 83). - A western species, ranging from southwestern United
States to central Mexico.
Habitat. - I have collected specimens from one locality each in western New Mexico and
eastern Arizona. At the former, in the pine-juniper zone, a large series was collected from
flowers of a short geranium (unidentified), while a few specimens were caught on white sweet
clover ( Meli lotus alba). In eastern Arizona, in the pine-spruce zone, specimens were swept
from grass under cover of pine trees.
Chorological affinities. - Uniquely distributed throughout southwestern United States and
western Mexico, this species is narrowly sympatric with S. multifaria and S. cristita along the
northern limit of its range. It is allopatric to other S. cristita group members.
Phylogenetic relationships (Fig. 104). - Though not well characterized, the S. cristita
group is considered monophyletic, and S. pisinnia is tentatively regarded as sister species to the
other members of the group. Phenetically, S. pisinnia is close to S. multifaria.
ParatypeS. — Sixty-eight males, 73 females. USA: Arizona: Rustler Pk., Portal, 8200', 3. VIII. 55, R.R. Dreisbach,
1M; Santa Catalina Mts., 14. VII. 50, L.D. Beamer, 1M; Dripping Springs, Organ Pipe, 31.111.67, D.M. Wood, 1 M;
Sunnyside Canyon, 11. VII. 40, D.G. Hall, 1M; ibid., Huachuca Mts., 9. VI 1.40, E.E. Kenaga, IF; ibid., D.E. Hardy, 2F;
Chiricahua Mts., 6. VIII. 33, Bryant, 1M; ibid., 7. VIII. 41, R.H. Beamer, 1M; Cochise Co., Chiricahua Mts., Flys Peak,
85-9700', 5. VIII. 27, J.A. Kusche, 1 M; Cochise Co., Southwestern Research Station, 8 km. w. Portal, 1650m, 1 3. VII. 56, E.
Ordway, 1M; ibid., 3.IX.59, D.D. Linsdale, 1M; ibid., 9,22.X.64, P.H. Arnaud, Jr., 3F; ibid., 27.VII.65, 1M; ibid.,
2. VIII. 65, IF; ibid., 22.IX.65, IF; ibid., 2.X.65, IF; ibid., 5-25.IX.65, C.W. Sabrosky, 2M, 13F; ibid., 6.VIII.65,
8, 26. IX. 66, V.D. Roth, 1M, 9F; ibid., 1 1,19, 22, 24. IX. 66, P.H. Arnaud, Jr., 1 M, 1 1 F. California: Snow Creek, 1500',
7. III. 55, W.R. Richards, IF; Garnet, 4.IV.45, A.L. Melander, IF; Indio, 13.XII.44, A.L. Melander, 2F; nr. Indio,
23.11.49, A.L. Melander, 1 M; Riverside, 2.V.35, A.L. Melander, 1 M; Riverside Co., Deep Canyon, 23.11.64, M.E. Irwin,
IF; ibid., P.L. Boyd Desert Research Center, s. Palm Desert, 16-17. X, 4-6.XI.69, S. Frommer and R. Worley, 4F; ibid.,
3 1 .1- 10.11.70, S. Frommer, R. Worley and L. La Pre, 3M, IF; ibid., 4- 1 1 .11.73, A. Tabet, 7F; ibid., 1 2.IV.75, J.B. Tucker,
1 M; ibid., J.B. Turner, 1 F. Nevada: Austin, 12.VIII.40, D.E. Hardy, 1 M. New Mexico: Grant Co., Gila Nat. For., Cherry
Creek Cpgd., 21 km. n. Silver City, 2250m, 3-4.VIII.80, J.E. and W.M. O'Hara, 23M, 9F; Socorro Co., Magdalena Mts.,
19.V1II.51, E.L. Kessel, 2F.
MEXICO: Chihuahua: Mesa del Huracan, 108°15'N 30°04'W, 7400', 21-25. VII. 64, J.E.H. Martin, 2M. Durango:
Rio Chico, 20 mi. w. Durango, 7000', 10. VIII. 64, J.F. McAlpine, 1 M; 10 mi. w. El Salto, 9000', 21.VI.64, J.F. McAlpine,
1 M; 6500', VII 1.64, J.F. McAlpine, 1 M; 24 mi. w. La Ciudad, 7000', 28.VI.64, W.R.M. Mason, 2M. Mexico: Pedregal de
San Angel, 5-8.VII.78, G. and M. Wood, 16M, 2F. Sinaloa: El Palmito, 6400', 2.VII.64, J.F. McAlpine, 1 M. (Deposited
in AMNH, CAS, CNC, DMW, JEOH, MSU, UASM, UCR, UKL and USNM.)
The S. geniculata group
North American species of the S. geniculata group have a short aristomere 3 (0.35-0.45
head height), average length proboscis and at least one of the following two characteristics: one
pair of median marginal setae on T1+2 (Fig. 8), and/or a long surstylus which extends beyond
apex of cerci by more than preapical width of surstylus (Figs. 70-71).
Siphona geniculata (De Geer)
Musca geniculata De Geer, 1776: 38.
Crocuta geniculata'. Coquillett, 1910: 518.
Siphona geniculata-. Mesnil, 1965: 866 (redescription).- Following New World records are based on misidentifications.
Coquillett, 1897: 76.— Aldrich, 1905: 444.— Aldrich, 1934: 109.— Sabrosky and Arnaud, 1965: 1 064. —
Cortes and Hichins, 1969: 57. -Cortes and Campos, 1970: 98.- Arnaud, 1978: 458.
Siphona analis Meigen, 1824: 157.— Herting, 1972: 3.
Siphona cinerea Meigen, 1824: 156.-- Herting, 1972: 4.
Bucentes cinerea Latreille, 1809: 339.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Siphona meigenii (Lepeletier and Serville in Latreille, et al.), 1828: 501 ( Bucentes ).
Stomoxys minuta Fabricius, 1805: 282.
Siphona nigrovittata Meigen, 1824: 157.— Herting, 1972: 11.
Siphona tachinaria Meigen, 1824: 156.-- Herting, 1972: 13.
Musca urbanis Harris, 1780: 153.
The type of Musca geniculata De Geer is lost (Herting, per. comm.). Nevertheless the
species now recognized as S. geniculata (whether or not the original one) is readily
distinguishable so a neotype is not required.
Recognition. - Specimens are relatively large for Siphona. Eye height and proboscis length
average, first flagellomere and aristomere 3 short. Most specimens with above average number
of hairs (10-20) on upper parafacial and frontal plate. This is one of two species with both a
long surstylus and 1 pair of strong median marginal setae on T1+2 (see below). Abdominal vitta
present, broader than average, and obscured by dense pruinosity. Four postsutural dorsocentral
setae.
With median marginal setae on T1+2 and a long surstylus, S. geniculata is unlike any North
American species other than S. hokkaidensis. Generally, shape and length of male first
flagellomere permits separation of specimens of these species, since most S. hokkaidensis
specimens with median marginals on T1+2 have large first flagellomeres (as in Fig. 32).
Description. — Specimens examined: 6 males, 3 females from North America, 16 males and 11 females from
Europe. Length: 4. 5-6. 5mm.
Male. Head (Fig. 28). Colouration: frontal vitta testaceous, brown to reddish-brown; gena and lower parafacial white
to light yellow; fronto-orbital plate and upper parafacial yellow to dark brown, darkest on orbital plate; scape and pedicel
testaceous to fuscous; first flagellomere fuscous to black; palpus yellow, apex slightly infuscate in most specimens;
proboscis reddish-brown or fuscous. Eye: size average for genus, 0.786-0.833 head height (Fig. 13), broad, widest at
center, evenly rounded along anterior margin. Macrotrichia: cluster of 10-20 hairs on upper parafacial and frontal plate in
most specimens; vibrissal angle average in most specimens, more heavily setose than average in few specimens with 2 or 3
setae and several setulae in addition to vibrissa and several hairs. Antenna: first flagellomere short, 0.420-0.484 head
height (Fig. 14), rounded distally along ventral margin; aristomere 3 short for genus, in most specimens 0.35-0.45 head
height. Mouthparts: proboscis length average for genus, 2.02-2.50 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown, not vittate; tegula testaceous to
reddish-brown; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals, 25 specimens examined, all
with four [males and females]. Acropod: claws and pulvilli average to slightly larger than average.
Preabdomen. Colouration: dark, faintly vittate; T4, T5 and broad vitta on T, +2 and T3 black in ground colour, rest of
preabdomen yellow in ground colour; surface dark gray or brownish-gray on areas of black ground colour, dark brown to
black around macrotrichial insertions; areas of yellow ground colour with surface light brown or tawny. Macrotrichia: 1
pair median marginals on T1+2; 1 or 2 pair strong lateral marginals on T1+2 and T3.
Genitalia (Fig. 91). Four examined, including 3 from European specimens. Apex of surstylus extended beyond apex of
cerci by more than preapical width of surstylus. Aedeagus: slope of distiphallus even with angle of basiphallus (not bent);
antero-ventral margin dentate; in profile parallel-sided and elongate; apex truncate or nearly so.
Female. As described for male except as follows.
Head (Fig. 50). Palpus equal or slightly longer than in males.
Preabdomen. Colouration: dark, not vittate; black in ground colour, surface gray finely mixed with light brown.
Geographical distribution (Fig. 91). - A wide ranging species in the Palearctic Region;
introduced and established in the lower Fraser Valley of British Columbia. It has not, to date,
been recorded elsewhere in North America. It is unlikely S. geniculata will spread beyond its
restricted range unless its introduced host, Tipula paludosa , extends its range, or S. geniculata
adapts to a native Tipula species.
Chorological affinities. - At present S. geniculata occupies a narrow range in North
America, and it is uncertain whether this species interacts with other Siphona species (such as
closely related S. hokkaidensis ) in the area.
Phylogenetic relationships (Fig. 105). - The sister species to this species is probably a
Palearctic species. In North America, S. hokkaidensis is most closely related to it.
North American species of Siphona Meigen
301
Siphona hokkaidensis Mesnil
Siphona hokkaidensis Mesnil, 1957: 36.— Mesnil, 1965: 869.- Herting, 1967: 9 (comparison of S. hokkaidensis Mesnil
and S'. silvarum Herting).
Siphona silvarum Herting, 1967: 9. New synonymy.
Type material examined. - Siphona hokkaidensis Mesnil, HOLOTYPE, female, labelled:
“Obihiro [Hokkaido, Japan]/ S. Takano [handwritten label]”; “295 [green label]”; “Siphona/
hokkaidensis Mesnil/ L.P. Mesnil det., 1970”; “Type [red label]”; “EX/ L.-P. MESNIL/
COLLECTION 1970” (CNC).
Siphona silvarum Herting, HOLOTYPE, male, labelled: “ex larvae/ Tipula/ irrorata Mg.
[handwritten label]”; “Mooswald/./ Freiburg [West Germany] i.B./ IV. 1958 [handwritten
label]”; “Siphona/ silvarum Hert./ L.P. Mesnil det., 1970”; “TYPE [red label]”; “EX/ L.-P.
MESNIL/ COLLECTION 1 970” [slash immediately following Mooswald is on label] (CNC).
Two puparia accompany type on same pin.
Recognition. - Eye height, first flagellomere and proboscis lengths varied. Aristomere 3
short and thick in most specimens. Most eastern specimens without marginal setae on T]+2 and
with first flagellomere subtriangular (Fig. 31). Specimens from Alberta, Saskatchewan and
Manitoba with short slender first flagellomere (Fig. 30). Northwestern specimens varied, few
with 1 or pair of median marginal setae on T1+2 and very large first flagellomere (Fig. 32).
Few specimens with above average number of setae on vibrissal angle, up to 4 present. Apex of
surstylus extended far beyond apex of cerci, distiphallus truncate or nearly so. Most specimens
with four postsutural dorsocentral setae.
Specimens with median marginal setae on T1+2 are similar to S. geniculata, and are
discussed under “Recognition” of that species.
Eastern specimens of S. hokkaidensis form a homogeneous group, readily distinguished in
most instances by the subtriangular (male), apically truncate (both sexes) first flagellomere
and short (and in most males, thick) aristomere 3. S. hokkaidensis specimens from other
regions varied, as discussed below. In all regions male genitalia permit separation of this species
from others, because of the very long surstylus and characteristic distiphallus (Fig. 71).
Description. — Specimens examined: 271 males, 144 females. Length: 3. 5-5. 5mm.
Male. Head (Figs. 30-32). Colouration: frontal vitta testaceous, brown to reddish-brown; gena and parafacial light
yellow to light brown; fronto-orbital plate brown to dark brown; scape, pedicel, and in few specimens portion of first
flagellomere above aristal insertion yellow, testaceous, or (in few specimens) fuscous; first flagellomere reddish-brown,
fuscous to black; palpus yellow, in few specimens apex infuscate; proboscis testaceous to fuscous. Eye: size highly varied,
0.71 1-0.820 head height (Fig. 13), slender and narrowed below center to broad and evenly rounded along anterior margin.
Maerotrichia: vibrissal angle with 2-4 setae and several hairs, in addition to vibrissa. Antenna: length and shape of first
flagellomere extremely varied, 0.426-0.667 head height (Fig. 14), treated below under geographic variation; aristomere 3
short for genus, in most specimens 0.35-0.45 head height and thickened almost to tip. Mouthparts: length of proboscis
highly varied, 1 .96-2.53 head height (Fig. 1 5).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula testaceous to fuscous; wing yellowish hyaline; legs average. Maerotrichia: postsutural
dorsocentrals, 7 specimens with three, 91 with four. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: extremely varied, vittate in all but very few specimens; eastern specimens, black ground
colour on T5, vittate or entirely black on T4, vittate on T1+2 and T3, rest of preabdomen yellow in ground colour; areas
black in ground colour with surface blue-gray, dark brown to black around macrotrichial insertions; areas yellow in ground
colour with yellow or light brown surface; specimens from Alberta, Saskatchewan and Manitoba with ground colour as for
eastern specimens, surface predominantly light coloured, brownish-gray on areas of black ground colour, yellow or light
brown on areas of yellow ground colour; vitta visible though faint through light coloured pruinosity; dark areas around
macrotrichial insertions very reduced; pattern in western specimens highly varied, most specimens similar to eastern
specimens, few similar to specimens from Alberta, Saskatchewan and Manitoba, and few specimens unique - more
extensively black in ground colour, yellow ground colour restricted to extreme lateral edge of T1 + 2 and antero-lateral edge
of T3, not vittate; surface predominantly blue-gray, wide area around macrotrichial insertions dark brown to black.
Maerotrichia: median marginals on T1+2 present or absent, data recorded in Fig. 81; 1 or 2 pair strong lateral marginals
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
on T1 + 2 and T3, data recorded for T3 in Fig. 81.
Genitalia (Fig. 71). Twenty-seven examined, including 1 from a European specimen. Apex of surstylus extended
beyond apex of cerci by more than preapical width of surstylus. Aedeagus: slope of distiphallus even with that of
basiphallus, or distiphallus very slightly bent downward; antero-ventral margin dentate; in profile nearly parallel-sided;
apex truncate or nearly so.
Female. As described for male except as follows.
Head (Fig. 51). Length of palpus subequal or slightly longer than in males.
Preabdomen. Colouration: in all geographic regions ground colour pattern very similar, not vittate, yellow ground
colour restricted to extreme lateral edge of T1+2 and antero-lateral edge of T3; eastern specimens very dark, surface
blue-gray to gray, wide areas around macrotrichial insertions dark brown to black, black ground colour very evident
through light coloured pruinosity; specimens from Alberta, Saskatchewan and Manitoba with surface brownish-gray,
dark areas around macrotrichial insertions reduced, ground colour obscured by light coloured pruinosity; specimens
from Washington, British Colombia, Northwest Territories, Yukon and Alaska with surface highly varied, from light
gray or brownish-gray with light appearance as in Alberta, Saskatchewan and Manitoba specimens, to very dark like
most eastern specimens.
Variation, with notes about synonymy. - Polymorphism in S. hokkaidensis was investigated
in some detail to provide the basis for a taxonomic decision regarding morphs. Two character
systems in particular were amenable to analysis. These were shape of first flagellomere and
number and position of abdominal setae. These characteristics were at least in part
geographically distributed, as shown in Fig. 81.
Three basic shapes of first flagellomere were recognized in males of S. hokkaidensis ,
permitting specimens to be sorted into three groups on that basis. The three antennal types,
designated 1, 2 and 3, are illustrated in Figs. 30-32. Intermediate forms were few and limited to
specimens from western North America in regions where more than one antennal type were
present. Specimens with intermediate first flagellomeres were assigned the antennal type they
most closely resembled.
Females of S. hokkaidensis have smaller first flagellomeres than males, so the three
antennal types recognized in males could not be applied to females. Females were treated in a
fourth catagory (“f” in Fig. 81).
Except for one sample from northern Quebec (Indian House Lake), all male specimens
examined from Ontario eastward had a type 2 antenna. In other characteristics this group
varied little.
Males from Alberta, Saskatchewan and Manitoba also formed a homogeneous group,
sharing type 1 antenna, a slightly larger eye and shorter proboscis than average for the species,
and similar colouration. This group was called S. hokkaidensis “a” and treated separately from
other specimens of the species ( S . hokkaidensis^ b”) in the Hubbs-Hubbs diagrams. Locality
records for specimens of this group are represented by squares on the distribution map (Fig.
81).
Throughout most of the Northwest Territories type 3 antenna predominated among
specimens examined. In Alaska and the Yukon all three antennal types were represented in
localities in relatively close proximity to one another, with mixed series recorded in two
samples.
Abdominal setae reflected a similar geographic pattern to the one exhibited by antennal
variation (Fig. 81). Median marginal setae on T1+2 were present in a greater percentage of
specimens with type 3 antenna than any other. However, in northwestern North America
specimens with antennal types 1 and 2 had a higher occurrence of median marginals than they
did anywhere else. The number of lateral marginals on T3 also conformed to this pattern (Fig.
81).
My interpretation of these data is as follows. Whatever events produced polymorphism in S.
hokkaidensis , they apparently did not result in permanent interruption of gene flow between
North American species of Siphona Meigen
303
morphs, as exemplified by the heterogeneous populations in northwestern North America.
Therefore I regard all morphs as members of a single species, S. hokkaidensis.
Most distinctive of all were specimens with type 1 antenna from Alberta, Saskatchewan and
Manitoba, and for this reason they were treated separately in the Hubbs-Hubbs diagrams.
Unfortunately, there were too few specimens of S. hokkaidensis from the western provinces to
permit detailed interpretation of the relationship between S. hokkaidensis “a” and “b”.
Primarily because specimens with a type 1 antenna were found in northwestern North America
with other characters varied, I concluded that S. hokkaidensis “a” does not represent a new
species or subspecies. Furthermore, two autapomorphies support the present interpretation of
S. hokkaidensis as one species: male genitalia are unique and almost unvaried over the North
American range of the species, and the arista is thickened in most specimens irrespective of
antennal type.
Differences between specimens with antennal types 2 and 3 are slight, so there is little
reason to suspect they represent more than morphs of a single species. Dr. Herting noted (per.
comm.) that both antennal types are represented in European specimens of “ S . silvarum", so
the situation is geographically widespread.
Mesnil first described S. hokkaidensis from a female specimen from Hokkaido, Japan
(1957), then redescribed the species from European specimens in his treatment of Siphona in
Lindner’s “Die Fliegen der palaearktischen Region” (1965). Herting (1967) compared
European and Japanese specimens and noted the former lacked median marginals on T1+2 and
differed in colouration from the latter. At the time neither form was known from North
America, and their distributions appeared disjunct in the Palearctic Region. There was
sufficient reason for believing the forms represented two species, so Herting (1967) named S.
hokkaidensis of Mesnil, 1965 (not 1957) Siphona silvarum.
From data presented herein the varied nature of abdominal setae in S. hokkaidensis is
documented. The type of S. silvarum conforms in all respects to the present interpretation of S.
hokkaidensis - bearing a type 2 antenna and lacking median marginals on T]+2 - so it has been
synonymized with S. hokkaidensis.
Geographical distribution (Fig. 81). - Holarctic, in the Palearctic Region ranging over
Europe, USSR and Japan; in the Nearctic Region widely distributed in Canada, Alaska and
extreme northeastern United States. This species has a more northern distribution in Canada
than any other Siphona species.
Chorological affinities. - This species is sympatric with another S. geniculata group
member, S. medialis, along the southern edge of its range. It is widely sympatric with S.
maculata, S. intrudens, S. lutea, S. multifaria and S. cristata.
Phylogenetic relationships (Fig. 105). - Among North American Siphona species, this
species is most closely related to S. geniculata. However, several European species resemble S.
hokkaidensis , and one of these is probably its sister species.
Siphona medialis new species
Holotype. - Male, labelled: “Cranberry I. [Island]/ Lockeport, N.S. [Nova Scotia,
Canada]/ 25.VII.1958/ J.R. Vockeroth” (CNC).
Allotype. - Female, same data as holotype (CNC).
Derivation of specific epithet. - S. medialis is the only wide ranging North America species
that is characterized by strong median marginal setae on T1+2. S', medialis is named in
Quaest. Ent., 1982, 18 (1-4)
304
O’Hara
recognition of this attribute.
Recognition. - A dark coloured species with a very large eye, short first flagellomere and \
average-sized proboscis. Aristomere 3 short in most specimens. Legs of western specimens very
dark, specimens from other areas with legs average to darker than average. Preabdomen black |
in ground colour (not vittate), with 1 pair median marginal setae on T1+2. Three or four
postsutural dorsocentral setae.
This is one of the more distinctive species, with its dark colouration, large eye, median !
marginals on T1+2 and (in most specimens) dark legs. It can only be confused with the two
other species with median marginals on T1+2.
Male specimens of S’, hokkaidensis have a smaller eye, longer surstylus and are lighter j
coloured than S. medialis. In addition, most specimens of S. hokkaidensis with median
marginals on T1+2 are from northwestern North America and have large first flagellomeres (as
in Fig. 32).
S. medialis is most easily distinguished from S. geniculata by colouration, relative position
of the apices of surstylus and cerci, and in almost all instances by range differences.
Description. - Length: 4. 0-5. 5mm.
Male. Head (Fig. 29). Colouration: frontal vitta testaceous to reddish-brown; gena and parafacial white to light
brown; fronto-orbital plate light to dark brown, darkest on orbital plate; scape and pedicel black in most specimens, fuscous
in few; first flagellomere black; palpus yellow, in few specimens apex infuscate; proboscis fuscous to black. Eye: very large
for genus, 0.786-0.833 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin. Macrotrichia:
vibrissal angle average in most specimens, few specimens more heavily setose than average with 2-4 setae in addition to
vibrissa and several hairs. Antenna: first flagellomere on average slightly longer than, but similar in shape to, that of S.
geniculata , 0.439-0.533 head height (Fig. 14); aristomere 3 short for genus, in most specimens 0.35-0.45 head height.
Mouthparts: proboscis length average for genus, 2.00-2.51 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula reddish-brown to black; wing yellowish to cinereous hyaline; leg colouration geographically
varied; specimens examined from Washington, Idaho, Oregon and California with femora and tibiae fuscous in most
specimens, reddish-brown in few; specimens from other areas average to darker than average, reddish-brown or fuscous
portion covering dorsal and posterior surfaces of fore femur in few specimens, reddish-brown portions on mid and hind
femora not covering more than apical two-thirds, most extensive dorsally, tibiae predominantly yellow. Macrotrichia:
postsutural dorsocentrals, 1 1 specimens with three, 14 with four. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: dark; most specimens not vittate, black in ground colour, surface blue-gray, gray or
brownish-gray, dark brown to black around macrotrichial insertions; in very few specimens T]+2 and T3 laterally yellow in
ground colour, surface light brown. Macrotrichia: one pair median marginals on T1+2; lateral marginals on T1+2 strong.
Genitalia (Fig. 72). Seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus bent slightly or sharply downward from angle of basiphallus; antero-ventral margin dentate; apex broad in
profile.
Female. As described for male except as follows.
Head (Fig. 52). Palpus equal or slightly longer than in males.
Preabdomen. Colouration: not vittate; as described for average male.
Geographical distribution (Fig. 94). - Transcontinental; widely distributed in western
United States, but restricted to region along Canada-United States border in central and
eastern North America.
Habitat. - I have collected specimens of this species only once, in southwestern Utah. Adults
were swept from a damp meadow of sedge and grass. Specimens collected by J.F. McAlpine
and R.L. Hurley in Ninette, Manitoba, were caught in Poa- sedge meadows (label data).
Chorological affinities. - This species is sympatric with S. hokkaidensis along the northern
edge of the formers’ range. In addition to being transcontinental, S. medialis is widely
distributed in western United States, and is sympatric with all Siphona species of America
north of Mexico except S. pisinnia, S. illinoiensis and S. floridensis.
Phylogenetic relationships (Fig. 105). - This species is related to the S. geniculata - S.
hokkaidensis lineage of the S. geniculata group.
North American species of Siphona Meigen
305
Paratypes. - Twenty-six males, 40 females. CANADA: Alberta: 15mi. e. Morley, 23. VI. 62, W.R.M. Mason, 1M;
13mi. n. Banff, Banff-Jasper Hwy., 4500', 26. VII. 55, R. Coyles, IF. Manitoba: Ninette, 14. VI. 58, J.F. McAlpine, 1M;
ibid., R.L. Flurley, IF; 2mi. e. Douglas, 27. VI 1.58, N.B. Chillcott, 1 F. New Brunswick: Kouchibouguac N.P., 7,1 3. VI 1.77,
J.F. McAlpine, 2M, IF; Birch Cove, nr. Chamcook, 4. VII. 65, G.E. Shewell, 4M. Nova Scotia: Cranberry I., Lockeport,
18,20,24-25, 27-28, 30.VII.58, J.R. Vockeroth, 2M, 6F; Halifax Co., Lawrencetown, 19-20.VII.67, D M. Wood, 2M, IF.
Prince Edward Island: Green Gables, Cavendish Beach, 22. VII. 67, D M. Wood, 2F. Quebec: mi. 139, rte. 58, La
Verendrye Prov. Pk„ 29. VI, 1 VI 1.65, D.M. Wood, 2M, 3F.
USA: California: Nevada Co., Sagehen, 5mi. nw. Hobart Mills, 20. VI. 54, M.T. James, 2F; ibid., 25. VI. 54, R.M.
Bohart, IF; ibid., 2.VII.54, E.I.Schlinger, IF; ibid., 22. VI. 72, R.A. Belmont, IF; ibid., 15.VII.64, M.E. Irwin, 1 M, IF;
Merced Co., Dos Palos, 7.V.49, IF; Sierra Co., 4.8mi. se. Sierraville, 14. VI. 59, G.W. Byers, 1M. Colorado: Estes Pk.,
7500', 2. VII. 61, C.H. Mann, 2M, IF; Hartsel, 20.VI.40, A. L. Melander, 2M, 2F; Tenn. Pass, 26.VII.17, J.M. Aldrich, IF;
2061, IF. Idaho: Moscow, 30.V.1 1, J.M. Aldrich, 1M. Maine: Sagahadoc Co., Popham Beach, 15. VII. 71, P. Ward, IF.
Michigan: Iosco Co., 30. VII. 48, R.R. Dreisbach, IF. Minnesota: Eaglesnest, 30. VI. 59, W V. Balduf, IF. Montana:
Bozeman, 20. VI. 06, 1M; Powell Co., Nigger Hill, VII, W.M. Mann, IF. Oregon: Klamath Co., Mare's Egg Spring,
12. VI. 64, J. Schuh, 2M; ibid., Fort Klamath, 26. VI. 52, E.I. Schlinger, IF. Utah: Grizzly Ridge Camp, 30mi. n. Vernal,
8400', 8. VII. 61, J.G. Chillcott, IF; Kane Co., Dixie Nat. For., Duck Ck. Cpgd., 50km. se. Cedar City, rte. 14, 2620m,
16-17.VIII.80, J.E. and W.M. O'Hara, 1 M, 5F. Washington: Pullman, 23.V.18, 4.VI.22, A.L. Melander, 1 M, IF.
(Deposited in CNC, INHS, JEOH, MSUB, UASM, UCD, UCR, UKL, USNM and WSUP.)
The S.futilis group
Members have an average length proboscis, large eye and narrow, short to average length
first flagellomere. S. brunnea is a provisional member.
Siphona futilis Wulp
Siphona futilis Wulp, 1890: 1 25.— Aldrich, 1905: 444.— Guimaraes, 1971: 170.
Bucentes futilis : Curran, 1932: 13 (in key).
Bucentes ceres Curran, 1932: 14. New synonymy.
Siphona ceres’. Guimaraes, 1971: 170.— Arnaud, 1978:458.
Notes about synonymy. - Curran included S. futilis in his 1932 key to adults of North
American Siphona species. However, Curran’s key appears to have been written primarily from
published species descriptions. There is no indication that Curran examined specimens of S’.
futilis prior to his description of S. ceres. I regard the type of -S', ceres as conspecific with the
lectotype of S.futilis.
Type material examined. - Siphona futilis , 14 syntypes from BMNH, 2 syntypes from
USNM. This series includes specimens of 2 species. One species is Siphonopsis plusiae
(Coquillett), and I have chosen to retain this name as valid because this species is the
type-species of Siphonopsis , and its life history has been investigated under that specific epithet
(Bloeser, 1914). The other species has been described by Curran under the name Bucentes
ceres , subsequent to the description of S. futilis. The lectotype here designated for Siphona
futilis is conspecific with B. ceres , and replaces that name.
LECTOTYPE, male, labelled: “Co-/ type [round yellow-ringed label]”; “Omilteme,/
Guerrero, [Mexico]/ 8000 ft./ July. H.H. Smith.”; “B.C.A. Dipt. II./ Siphona/ futilis,/
v.d.W.“; “(5”; “Central America./ Pres, by/ F.D. Godman./ O. Salvin./ 1903-172“ (BMNH).
My lectotype label ’’LECTOTYPE/ Siphona/ futilis/ van der Wulp/ O’Hara designation/
Selected 1981 [red bordered, handprinted label]” has been attached to this specimen.
PARALECTOTYPES, 7 specimens of Siphonopsis plusiae (Coq.) (6 BMNH, 1 USNM).
Eight specimens (7 BMNH, 1 USNM) conspecific with lectotype: MEXICO, DISTRITO
FEDERAL: Mexico City, V.1888, H.H.S., 1M. GUERRERO: same data as lectotype, 2M,
2F; Xucumanatlan, 7000ft, VII, H.H. Smith, 2M. VERACRUZ: Orizaba, XII. 1887, H,H.S.
& F.D.G., IF. [BMNH specimens damaged in transit, most in very poor condition.] To these
specimens both paralectotype and determination labels have been attached.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Bucentes ceres , HOLOTYPE, female, labelled: “Antigua, Guatemala]./ 5-V-1931/ D.M.
Bates 506”; “Bucentes/ TYPE/ ceres 9/ Curran/ No. [red label]”; “Bucentes/ ceres / Curran
[handwritten det. label]” (AMNH). My determination label “Siphona/ futilis / van der Wulp/
det./ J.E. O’Hara 1981” has been attached to this specimen.
Recognition. - Eye large, first flagellomere short and proboscis length average. Scape and
pedicel fuscous to black. Palpus yellow in most specimens, slightly infuscate apically in few
specimens. Male preabdomen light coloured, vittate. Female preabdomen darker than in male;
without vitta. Distiphallus bent sharply downward from angle of basiphallus, narrow in profile,
apex sloped sharply postero-ventrally in most specimens. Three postsutural dorsocentral setae
in most specimens (95%).
Eye and proboscis sizes are very similar between S. futilis and S. illinoiensis. S. futilis has a
shorter first flagellomere. There are also slight colour differences: scape and pedicel are dark
coloured in S. futilis and light coloured in S. illinoiensis ; female preabdomen broadly vittate in
S. illinoiensis , entirely dark in ground colour in S. futilis. There is a marked similarity between
distiphalli of these species, attesting to their close relationship. The main difference is that the
distiphallus is bent sharply downward from angle of basiphallus in S. futilis and is even with
basiphallus in S. illinoiensis.
Since S. futilis and S. illinoiensis are allopatric, the only species likely to be confused with
the former is S. brunnea. Head characteristics are similar, but dark colouration of S. brunnea
in combination with other features readily separate these species (see S. brunnea ,
“Recognition”).
Description. — Specimens examined: 66 males, 53 females. Length: 4. 0-5. 0mm.
Male. Head (Fig. 34). Colouration: frontal vitta testaceous to reddish-brown; gena and parafacial white, light yellow
to light brown; fronto-orbital plate light to dark brown, darkest on orbital plate; scape and pedicel fuscous to black; first
flagellomere black; palpus yellow, slightly infuscate at apex in few specimens; proboscis reddish-brown or fuscous. Eye:
large for genus, 0.847-0.911 head height (Fig. 13), broad, widest at center, evenly rounded along anterior margin.
Macrotrichia: average for genus. Antenna: first flagellomere 0.450-0.517 head height (Fig. 14), slender, rounded distally
along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 2.10-2.41
head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula testaceous to reddish-brown; wing yellowish hyaline; legs average. Macrotrichia:
postsutural dorsocentrals, 39 specimens with three, 2 with four. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: light coloured, vittate; yellow ground colour laterally on T1+2, T3 and anterolaterally on T4,
rest black in ground colour; vitta long and narrow; areas black in ground colour with surface blue-gray, gray or
brownish-gray, dark brown to black around macrotrichial insertions; areas yellow in ground colour with surface yellow to
light brown; pattern varied in few specimens, slightly greater or lesser than average amounts of black ground colour.
Macrotrichia: median marginals absent from T1+2; lateral marginals on T]+2 strong.
Genitalia (Fig. 73). Seven examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin dentate; in profile narrow, apex
sloped sharply postero-ventrally in most specimens.
Female. As described for male except as follows.
Head (Fig. 53). Palpus longer than in males in few specimens.
Preabdomen. Colouration: darker than male, without vitta; most specimens black in ground colour, very few laterally
yellow in ground colour on T, +2; surface as described for male.
Geographical distribution (Fig. 96). - Recorded from central Mexico to Costa Rica.
Chorological affinities. - The range of this species encompasses most of the range of S.
tropica , and the entire known ranges of S. brunnea, S. rizaba, S. akidnomyia and S.
longissima. The ranges of S. futilis and S. pisinnia overlap in central Mexico. S. futilis is
allopatric to sister species S. illinoiensis.
Phylogenetic relationships (Fig. 107). - This species is sister species to 5. illinoiensis , from
which it differs only slightly.
North American species of Siphona Meigen
307
Siphona illinoiensis Townsend
Siphona illinoiensis Townsend, 1891: 368.- Coquillett, 1 897: 76 (as syn. of S. geniculata).- Aldrich, 1 905: 444 (as syn. of
S', geniculata).- Bezzi and Stein, 1907: 382 (as syn. of S. geniculata).- Sabrosky and Arnaud, 1965: 1064.
Type material examined. - One syntype, here designated LECTOTYPE, male [not
female], labelled: “9”; “Robertson/ S. Illinois”; “Siphona/ illinoiensis/ Tws”; “Type”;
“COTYPE/ Siphona/ illinoiensis/ Tns. [red label]” (UKL). My lectotype label
“LECTOTYPE/ Siphona/ illinoiensis Tnsd./ O’Hara designation/ Selected 1981 [red
bordered label]” has been attached to this specimen.
Townsend described S. illinoiensis from 2 specimens. The UKL collection has only one
specimen labelled “COTYPE”; the one herein designated lectotype. However, the UKL
collection has a male specimen of S. illinoiensis with same locality label as the type. It is
probably the missing syntype.
Recognition. - Eye large, first flagellomere and proboscis lengths average. Scape and
pedicel yellow to testaceous. Male preabdomen light coloured, vittate. Female preabdomen
slightly darker than male, with short broad vitta. Slope of distiphallus even with that of
basiphallus. Distiphallus narrow in profile, apex sloped sharply postero-ventrally in most
specimens. Three postsutural dorsocentral setae in most specimens (86%).
Similarities and differences between S. illinoiensis and S. futilis are discussed under
“Recognition” of the latter species.
Within central and eastern North America, specimens of S. cristita and S. multifaria may
be mistaken for S. illinoiensis. Most male specimens of S. illinoiensis can be distinguished by
characteristics given in key and by comparing specimen measurements with those presented in
the Hubbs-Hubbs diagrams. Male genitalia of S. illinoiensis are relatively unvaried, and
sufficiently different from S. cristita and S. multifaria to aid in identifications.
Not all female specimens of S. illinoiensis can be distinguished from those of S. cristita and
S. multifaria. Eye and proboscis sizes can yield ambiguous results. In most specimens the light
brownish-gray, non-vittate female preabdomen of S. illinoiensis can be distinguished from the
abdominal colouration of S. cristita and S. multifaria. However, decisions regarding subtle
colour differences are untrustworthy if specimens of only one species are at hand.
Description. — Specimens examined: 217 males, 163 females. Length: 3. 0-5. 0mm.
Male. Head (Fig. 1 ). Colouration: frontal vitta yellow, testaceous to reddish-brown; gena and parafacial white to light
yellow; fronto-orbital plate light yellow to light brown or brown, darkest on orbital plate; scape, pedicel, and in few
specimens portion of first flagellomere above aristal insertion yellow to testaceous; first flagellomere fuscous; palpus
yellow; proboscis testaceous to reddish-brown. Eye: large for genus, 0.807-0.909 head height (Fig. 13), broad, widest at
center, evenly rounded along anterior margin. Macrotrichia: average for genus. Antenna: first flagellomere 0.500-0.596
head height (Fig. 14), slender, rounded distally along ventral margin; length of aristomere 3 average for genus.
Mouthparts: proboscis length average for genus, 1.93-2.49 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula testaceous; wing yellowish hyaline; legs average. Macrotrichia: postsutural dorsocentrals,
91 specimens with three, 15 with four. Acropod: claws and pulvilli average-sized (Fig. 4).
Preabdomen. Colouration: light coloured, vittate; most specimens black in ground colour on T4, T5 and medially on
Ti+2 and T3, rest of preabdomen yellow in ground colour; areas black in ground colour with surface brownish-gray, brown
to black around macrotrichial insertions; areas yellow in ground colour with surface light yellow or light brown to tawny;
pattern varied in few specimens, areas black in ground colour slightly more, or less, extensive. Macrotrichia: median
marginals absent from T, +2; lateral marginals on T, +2 strong.
Genitalia (Fig. 9). Twenty examined. As described for S. futilis, except slope of distiphallus even with that of
basiphallus in most specimens, slightly bent downward in few.
Female. As described for male except as follows.
Head (Fig. 54). Palpus longer and thicker than in males in most specimens.
Preabdomen. Colouration: average specimens slightly darker than male, with yellow ground colour restricted to lateral
edge of T1+2 and antero-lateral edge of T3, dark vitta short and broad; surface colour and variation as decribed for male.
Quaest. Ent., 1982, 18 (1-4)
308
O’Hara
Geographical distribution (Fig. 92). - Widely distributed in eastern United States.
Chorological affinities. - This uniquely distributed species is parapatric or narrowly
sympatric with S. hokkaidensis, S. maculata, S. lutea, S. medialis and S', cristata along the
northern edge of its range. Widely sympatric with S. illinoiensis are S. intrudens and S.
multifaria. S. illinoiensis is allopatric to sister species S.futilis.
Phylogenetic relationships (Fig. 107). - This species is sister species to the Mexican species
S. futilis.
Siphona brunnea new species
Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] 9600ft./ Zontehuitz, nr. S.
Crist. [San Cristobal de las Casas]/ 17 May 1969/ W.R.M. Mason” (CNC).
Allotype. - Female, same data as holotype (CNC).
Derivation of specific epithet. - Brunneus is Latin for brown. This species is named for its
very dark brown colouration.
Recognition. - Eye large, first flagellomere short and proboscis length average. A dark
brown species, with apical half of palpus fuscous or black, wing brownish hyaline and legs in
some specimens darker than average. Male tarsal claws slightly longer than average. Male
preabdomen not vittate, surface dark brown with a light coloured band across anterior portion
of each tergite. Female abdominal colouration varied. Surstylus slightly longer than cerci.
Distiphallus bent sharply downward from basiphallus, narrow and truncate in profile. Four
postsutural dorsocentral setae.
An easily recognized species, quite different from members of the Mexican S. tropica group.
Similar in head characteristics to S.futilis , but distinguishable on basis of colouration, number
of postsutural dorsocentrals and in particular by structure of distiphallus.
Description. - Length: 4.5-5.0mm.
Male. Head (Fig. 35). Colouration: frontal vitta dark brown to reddish-brown; gena and parafacial light brown;
fronto-orbital plate dark brown; antenna black; palpus yellow basally, apical half fuscous or black; proboscis fuscous or
black. Eye: large for genus, 0.833-0.843 head height (Fig. 13), broad, widest above center, slightly narrowed below center.
Macrotrichia: average for genus. Antenna: first flagellomere 0.463-0.510 head height (Fig. 14), short and distally rounded;
length of aristomere 3 average for genus. Mouthparts: proboscis length average for genus, 2.35-2.44 head height (Fig. 15).
Thorax. Colouration: dorsum dark brown; tegula black; wing brownish hyaline; legs average to darker than average,
reddish-brown regions covering dorsal and posterior surfaces of fore femur and distal half of mid and hind femora in two
specimens. Macrotrichia: postsutural dorsocentrals, four in all specimens. Acropod: claws and pulvilli slightly larger than
average.
Preabdomen. Colouration: dark, not vittate; black in ground colour except for extreme lateral edge of T1+2 and T3;
surface dark brown with light bands across anterior 0.25 of T3, T4 and T5. Macrotrichia: median marginals absent from
T, + 2; lateral marginals on T, +2 strong.
Genitalia (Fig. 74). One examined. Apex of surstylus extended slightly beyond apex of cerci. Aedeagus: distiphallus
bent sharply downward from angle of basiphallus; antero-ventral margin dentate; in profile narrow, apex truncate.
Female. As described for male except as follows.
Head (Fig. 55). Length of palpus subequal in males and females.
Preabdomen. Colouration: light bands slightly wider than in male; one female with same ground colour pattern as
male, other female vittate, with yellow ground colour dorso-laterally on T, +2.
Geographical distribution (Fig. 97). - Known only from Chiapas, Mexico.
Chorological affinities. - From label data this species has evidently been collected from the
same locality as S. akidnomyia, though at a different time of year.
Phylogenetic relationships (Fig. 107). - Due to the highly derived nature of this species, it is
difficult to place with certainty. It is here interpreted as a member of the S.futilis group on the
basis of its large eye and general habitus.
North American species of Siphona Meigen
309
Paratypes. — One male, 1 female. MEXICO: Chiapas: Zontehuitz, nr. San Cristobal de las Casas, 9600', 17.V.69,
W.R.M. Mason, 1M, IF. (Deposited in CNC.)
The * S', tropica group
Group characteristics are: long to very long proboscis and a very long first flagellomere, the
latter in profile apically wide and broadly rounded along ventral margin. Most species have a
large eye.
Siphona tropica (Townsend)
Phantasiosiphona tropica Townsend, 1915: 93.— Townsend, 1936: 150 (in key to genera of the Siphonini), Townsend,
1940: 286 (redescription of genus).- Guimaraes, 1971: 169.
Bucentes tropica-. Curran, 1932: 13 (in key).
Type material examined. - PARATYPE, male [not female], labelled: “Granada /
Nicaragua/ Coll. Baker”; “V/ 77 [tiny handwritten label]”; “Paratype No./ 19954/
U.S.N.M. [red label]”; “Phantasiosiphona/ tropica/ Tns. det/ Sabrosky” (USNM).
Recognition. - Eye large, first flagellomere and proboscis long. Wing brownish hyaline, with
single setula on bend of Rj in most specimens (89%). Preabdomen light coloured, vittate in both
sexes. Distiphallus distinctive, slope even with that of basiphallus, apex narrow and pointed in
profile. Three postsutural dorsocentral setae.
Similar to S. longissima and S. rizaba in proboscis length, and to latter in eye height. Most
specimens of S. tropica are readily identified by presence of a setula distally on R,. No other
Middle American species is known to exhibit this character state. Structure of the male
distiphallus is particularly diagnostic for this species.
Description. — Specimens examined: 14 males, 6 females. Length: 4. 0-5. 0mm.
Male. Head (Fig. 38). Colouration: frontal vitta testaceous to dark brown; gena and parafacial white, light yellow to
light brown; fronto-orbital plate light to dark brown, darkest on orbital plate; antenna fuscous; palpus yellow; proboscis
testaceous, reddish-brown to fuscous. Eye: large for genus, 0.860-0.891 head height (Fig. 13), slender, widest above center,
narrowed below center. Macrotrichia: average for genus. Antenna: first flagellomere very long and broad for genus,
0.654-0.745 head height (Fig. 14), rounded distally along ventral margin and in most specimens produced slightly forward
distally along dorsal margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for genus, 2.76-3.02
head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula fuscous to black; wing brownish hyaline; legs average. Macrotrichia: postsutural
dorsocentrals, three in all 19 specimens examined; wing, 19 specimens examined: one seta on both wings in 14 specimens; 1
seta on one wing only in 3 specimens; setae absent, 2 specimens. Acropod: claws and pulvilli average-sized.
Preabdomen. Colouration: vittate; varied, yellow ground colour ranging from laterally on T1+2 and antero-laterally on
T3 to laterally on T]+2, T3 and antero-laterally on T4, surface yellow, light brown to tawny; areas black in ground colour
with surface blue-gray, gray to brown, darker around macrotrichial insertions. Macrotrichia: median marginals absent
from T, +2; lateral marginals on T, +2 strong.
Genitalia (Fig. 78). Four examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
slope of distiphallus even with that of basiphallus (not bent); antero-ventral margin dentate; apex narrow and pointed in
profile.
Female. As described for male except as follows.
Head (Fig. 57). Length of palpus subequal in males and females.
Preabdomen. Colouration: vittate; yellow ground colour laterally on T1 + 2 and antero-laterally on T3, otherwise as
described for male.
Geographical distribution (Fig. 101). - Recorded from southern Mexico to western
Colombia.
Habitat. - Recorded from cloud forest at 5100' in Puebla, 5 mi. ne. of Teziutlan. In
Veracruz (Fortin de las Flores) specimens of this species were collected between 2500-3000'
(from label data).
Quaest. Ent., 1982, 18 (1-4)
310
O’Hara
Chorological affinities. - Though widely distributed throughout Central America and
southeastern Mexico, records do not show specimens of this species as having been collected
with those of other Middle American Siphona species.
Phylogenetic relationships (Fig. 106). - Relationships among S. tropica group members are
unclear, for reliable synapotypies have not been discovered. S. tropica and S. rizaba are
tentatively grouped as sister species on the basis of a very large eye and the loss of hooks on the
distiphallus.
Siphona rizaba new species
Holotype. - Male, labelled: “Orizaba, Ver[acruz].,/ Mex[ico]. 8/12-22/61/ R&K
Dreisbach” (MSU). Genitalia in microvial on pin below specimen.
Allotype. - Female, same data as holotype, with Reinhard determination label: “ Bucentes /
tropica/ Tm. R. [Reinhard initial]” (MSU).
Derivation of specific epithet. - This species is named after the type-locality, Orizaba, from
which the first letter has been removed.
Recognition. - Eye large, first flagellomere and proboscis long. Wing yellowish to brownish
hyaline. Preabdomen light coloured, vittate in both sexes. Externally without good diagnostic
characteristics. Apex of surstylus extended slightly to far beyond apex of cerci. Distiphallus
slightly bent downward from basiphallus, apex rounded in profile. Three postsutural
dorsocentral setae.
This species is the least distinctive of the S. tropica group. It shares the general
characteristics of the other members of the group, but lacks good diagnostic features. S. tropica
specimens without a single setula on R] can be confused with this species. Dissection of male
genitalia is advisable where ratios of first flagellomere to eye and head heights are near upper
values for S. rizaba and lower values of S. tropica and S. longissima (Figs. 14,16).
Description. - Length: 4. 0-5. 0mm.
Male. Head (Fig. 38). Colouration: frontal vitta testaceous to brown; gena and lower parafacial white, light yellow to
light brown; upper parafacial and frontal plate light brown to dark brown; orbital plate brown to dark brown; scape and
pedicel testaceous to reddish-brown or fuscous; first flagellomere fuscous; palpus yellow; proboscis testaceous to
reddish-brown. Eye: large for genus, 0.846-0.870 head height (Fig. 13), slender, widest at center, slightly narrowed below
center. Macrotrichia: average for genus. Antenna: first flagellomere long and broad, 0.593-0.648 head height (Fig. 14),
broadly rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for
genus, 2.74-3.03 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula reddish-brown to fuscous; wing yellowish to brownish hyaline; legs average. Macrotrichia:
postsutural dorsocentrals, three in all specimens. Acropod: claws and puivilli average-sized.
Preabdomen. Colouration: vittate; T4, T5 and vitta on T1+2 and T3 black in ground colour, surface gray,
brownish-gray or brown, darker around macrotrichial insertions; T1+2 and T3 laterally yellow in ground colour, surface
yellow to light brown. Macrotichia: median marginals absent from T1+2; lateral marginals on T]+2 strong.
Genitalia (Fig. 79). Four examined. Apex of surstylus extended slightly beyond to more than preapical width of
surstylus beyond apex of cerci. Aedeagus: distiphallus bent slightly downward from angle of basiphallus; antero-ventral
margin dentate; apex rounded in profile.
Female. As described for male except as follows.
Head (Fig. 58). Palpus subequal in length in males and females.
Preabdomen. Colouration: short vittate; yellow ground colour restricted to T1+2, otherwise as described for male.
Geographical distribution (Fig. 98). - Recorded from two localities in Veracruz, Mexico.
Chorological affinities. - This species is evidently sympatric with S. futilis and S. tropica ,
but specimens of it have not been caught together with those of any other species.
Phylogenetic relationships (fig. 106). - See this heading under S. tropica.
Paratypes. - Three males, 2 females. MEXICO: Veracruz: Orizaba, 12-22. VIII. 61, R. and K. Dreisbach, 1M, 2F;
Cordoba, 13,20.VII.66, J.S. Buckett, M R. and R.C. Gardner, 2M; (Deposited in CNC, MSU and UCD.)
North American species of Siphona Meigen
311
Siphona longissima new species
Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] lOmi NE./ San Cristobal [de
las Casas]/ 13.V.69 7500'/ H.J. Teskey” (CNC). Genitalia in microvial on pin below
specimen.
Derivation of specific epithet. - From longissimus, Latin for “longest”. S. longissima has
the longest proboscis of the species described in this paper.
Recognition. - Eye large and first flagellomere long. Proboscis very long, one specimen with
longest proboscis of any recorded North American species (3.2X head height). Wing brownish
hyaline. Tarsal claws intermediate in size between average and large. Preabdomen vittate in
male. Distiphallus distinctive, bent sharply downward from basiphallus, antero-ventral margin
with small hooks, apex narrow and pointed in profile. Three postsutural dorsocentral setae.
Female of species unknown.
Most similar to S. tropica and S. rizaba. Other than characteristic sizes of eye, first
flagellomere and proboscis, S. longissima can be distinguished by shape of distiphallus and the
small hooks (as in S. macronyx and S. akidnomyia) along its antero-ventral margin.
Description. - Length: 4.5-5.0mm.
Male. Head (Fig. 37). Colouration: frontal vitta brown; gena and parafacial light brown; fronto-orbital plate brown to
dark brown, darkest on orbital plate; antenna fuscous to black; palpus yellow; proboscis fuscous. Eye: slightly larger than
average for genus, 0.796-0.850 head height (Fig. 13), slender, widest above center, narrowed below center. Macrotrichia:
average for genus. Antenna: first flagellomere very long and broad for genus, 0.650-0.667 head height (Fig. 14), rounded
distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis long for genus, 2.74-3.20
head height (Fig. 15).
Thorax. Colouration: dorsum darker than average, dark brown; tegula fuscous to black; wing brownish hyaline; legs
average. Macrotrichia: postsutural dorsocentrals, three in all specimens. Acropod: claws and pulvilli intermediate in size
between average and large.
Preabdomen. Colouration: vittate; yellow ground colour laterally on T1+2 and T3, surface yellow; areas black in
ground colour with surface gray to brown, darker around macrotrichial insertions. Macrotrichia: median marginals absent
from T| +2; lateral marginals on T, +2 strong.
Genitalia (Fig. 77). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus sharply bent downward from angle of basiphallus; antero-ventral margin with small hooks (similar to S.
macronyx and S. akidnomyia)-, apex narrow and pointed in profile.
Female. Unknown.
Geographical distribution (Fig. 100). - Recorded from southern Mexico.
Chorological affinities. - Specimens of this species have been caught in the same area as
those of S.futilis and S. akidnomyia , but on different dates (from label data).
Phylogenetic relationships (Fig. 106). - The position of this species within the S. tropica
group is tentatively as sister species to the highly derived S. akidnomyia.
Paratypes. - Two males. MEXICO: Chiapas: 3mi. n. San Cristobal de las Casas, 7000', 29.V.69, H.J. Teskey, 1M.
Distrito Federal: Desierto de Los Leones, X.65, N.L.H. Krauss, 1 M. (Deposited in CNC and USNM.)
Siphona akidnomyia new species
Holotype. - Male, labelled: “MEX. [Mexico] Chis. [Chiapas] 9600ft./ Zontehuitz, nr. S.
Crist. [San Cristobal de las Casas]/ 25 June 1969/ W.R.M. Mason” (CNC). Genitalia in
microvial on pin below specimen.
Allotype. - Female, labelled: “MEXICO, Chiapas/ Mt. Tzontehuitz/ 9400' 12mi. NE
San/ Cristobal [de las Casas] 27. V./ 1969 B.V. Peterson” (CNC).
Derivation of specific epithet. - Derived from the Greek for feeble ( akidnos ) fly ( myia ).
This is in reference to Siphona in general, which lack the size and strength of their larger
relatives.
Quaest. Ent., 1982, 18 (1-4)
312
O’Hara
Recognition. - Eye size average, first flagellomere and proboscis long. Palpus long, apex
infuscate in most specimens. Wing brownish hyaline. Male tarsal claws large. Male
preabdomen dark, broadly vittate, female preabdomen average vittate. Distiphallus unique,
short and broad in profile with small hooks on antero-ventral margin. Three postsutural
dorsocentral setae.
Closest in appearance to, yet readily distinguished from, S. tropica , S. rizaba and S.
longissima. S. akidnomyia is the darkest of these species (though lighter than S. brunnea ), and
has a shorter proboscis, smaller eye and longer tarsal claws than the others. The infuscate
palpus is very large for Siphona , especially in the female. The distiphallus has small hooks
along its antero-ventral margin (as in S. macronyx and S. longissima ), and a very distinctive
shape.
Description. - Length: 4. 0-5. Omm.
Male. Head (Fig. 36). Colouration: frontal vitta brown; gena and parafacial white to light brown; fronto-orbital plate
brown to dark brown, darkest on orbital plate; antenna fuscous to black; palpus yellow, apex infuscate; proboscis fuscous.
Eye: size average for genus, 0.781-0.804 head height (Fig. 13), slender, widest at center, slightly narrowed below center.
Macrotrichia: average for genus. Antenna: first flagellomere long and broad for genus, 0.585-0.660 head height (Fig. 14),
broadly rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: palpus longer than
average for genus; proboscis long for genus, 2.49-2.66 head height (Fig. 15).
Thorax. Colouration: dorsum average to dark brown; tegula fuscous; wing brownish hyaline; legs average.
Macrotrichia: postsutural dorsocentrals, three in all specimens. Acropod: claws and pulvilli large.
Preabdomen. Colouration: broadly vittate; laterally on T|+2 and laterally or antero-laterally on T3 yellow in ground
colour, surface yellow; areas black in ground colour with surface blue-gray, gray to brown, darker around macrotrichial
insertions. Macrotrichia: median marginals absent from T1+2; lateral marginals on Tl+2 strong.
Genitalia (Fig. 76). Five examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus slightly or sharply bent downward from angle of basiphallus; narrow section between basiphallus and
distiphallus longer than average; antero-ventral margin with small hooks (similar to 5. macronyx and S. longissima ); in
profile distiphallus short and broad, apex pointed.
Female. As described for male except as follows.
Head (Fig. 59). Palpus larger than in males, very long for genus, apex infuscate in one female.
Preabdomen. Colouration: vitta average; surface colour as described for male, except slightly lighter.
Geographical distribution (Fig. 99). - Recorded from southern Mexico.
Chorological affinities. - Evidently sympatric with S. longissima, S. brunnea and S.futilis.
Specimens of this species have been recorded from the same localities as the former two species,
but were not collected on the same dates.
Phylogenetic relationships (Fig. 106). - This is the most highly derived species of the S.
tropica group. Intragroup relationships are difficult to interpret without additional characters.
S. longissima is tentatively placed as sister species to S. akidnomyia on the basis of larger than
average tarsal claws.
Paratypes. - Four males, 1 female. MEXICO; Chiapas: Mt. Tzontehuitz, 9400', 12mi. e. San Cristobal de las Casas,
27.V.69, B.V. Peterson, 1M; 8mi. ne. San Cristobal de las Casas, 7500", 15.V.69, H.J. Teskey, 1M; San Cristobal de las
Casas, 7087', 13, 28. VI. 69, B.V. Peterson, 1M,1F. Distrito Federal: Desierto Leones, 111, IV, V. 65, N.L.H. Krauss, 1M.
(Deposited in CNC and USNM.)
The S. macronyx group
This group contains one highly autapotypic species, S. macronyx. The characteristics of this
species (see “Recognition” below) are such that it cannot, at this time, be placed in one of the
other North American Siphona species groups. Further study and additional characters might
indicate that S. macronyx is a derived member of one of the other recognized groups. If so, then
it should be placed in that group.
North American species of Siphona Meigen
313
Siphona macronyx new species
Holotype. - Male, labelled: “Pullman, Wash. [Washington, USA]/ Malaise Trap.”; “R.W.
Dawson/ May 6 1969” (WSUP). Genitalia in microvial on pin below specimen.
Allotype. - Female, same data as holotype except date May 13, 1969 (WSUP).
Derivation of specific epithet. - From the Greek words for large claw, makros and onyx,
after this attribute of the species.
Recognition. - Head with characteristic habitus due to broad eye, above average number of
setae and setulae on vibrissal angle, short first flagellomere and dark colouration. Proboscis
average length. Male tarsal claws large. Preabdomen dark, in male broadly vittate, in female
entirely black in ground colour. Distiphallus bent sharply downward from angle of basiphallus
with several small hooks along antero-ventral margin. Three postsutural dorsocentral setae.
The derived features of this species are such that specimens are not closely similar to those
of any other species.
Description. - Length: 4.0-5. 5mm.
Male. Head (Fig. 33). Colouration: frontal vitta orange, testaceous to reddish-brown; gena and parafacial light brown
in most specimens, light yellow in few; fronto-orbital plate brown to dark brown; scape, pedicel, and in few specimens
portion of first flagellomere above aristal insertion yellow, testaceous to reddish-brown; first flagellomere fuscous to black;
palpus yellow; proboscis reddish-brown to fuscous. Eye: size average for genus, 0.742-0.804 head height (Fig. 13), broad,
widest at center, evenly rounded along anterior margin. Macrotrichia: vibrissal angle in most specimens with cluster of 3-6
setae and several setulae in addition to vibrissa. Antenna: first flagellomere short, 0.403-0.500 head height (Fig. 14),
rounded distally along ventral margin; length of aristomere 3 average for genus. Mouthparts: proboscis length average for
genus, 2.00-2.34 head height (Fig. 15).
Thorax. Colouration: dorsum average, blue-gray or gray finely mixed with brown in most specimens, 3 brown vittae
visible in few specimens; tegula testaceous to reddish-brown; wing brownish hyaline; legs average. Macrotrichia:
postsutural dorsocentrals, 60 specimens with three, 1 with four [males and females]. Acropod: claws and pulvilli large.
Preabdomen. Colouration: dark, broadly vittate; T4, T5 and vitta on T1+2 and T3 black in ground colour, surface gray
to brown, black around macrotrichial insertions; yellow ground colour laterally on Tl+2 and T3 in most specimens, surface
yellow to light brown; in 2 specimens yellow ground colour very reduced. Macrotrichia: median marginals absent from
T1+2; lateral marginals strong.
Genitalia (Fig. 75). Two examined. Apex of surstylus even with or extended slightly beyond apex of cerci. Aedeagus:
distiphallus bent sharply downward from angle of basiphallus; antero-ventral margin with small hooks (similar to S.
longissima and 5. akidnomyia)', apex broad in profile.
Female. As described for male except as follows.
Head (Fig. 56). In most specimens gena yellow, parafacial and frontal plate bright yellow tinged with brown (near
golden); length of palpus slightly longer than in male in few specimens.
Preabdomen. Colouration: not vittate; black in ground colour except for faint traces of yellow ground colour laterally
on T1+2 in few specimens; surface light blue-gray, gray or brownish-gray, mixed with brown medially.
Geographical distribution (Fig. 88). - Known only from Washington and western Idaho.
Chorological affinities. - The small known range of this species is sympatric with the
northern limits of the ranges of western species S. lurida , S. oligomyia and N. pacifica. S.
macronyx is parapatric or sympatric with S. hokkaidensis, S. maculata, S. intrudens, S. lutea,
S. multifaria, S. medialis and S. cristita, though specimens of it have not been collected with
specimens of any of these species (from label data).
Phylogenetic relationships (Fig. 103). - See discussion under species group heading above.
Paratypes. - Eleven males, 49 females. USA: Idaho: Moscow Mt., 5. VII. 19, A.L. Melander, 1M. Washington:
Whitman Co., Almota, 27. IV. 72, W.J. Turner, 1 M, 2F; Jefferson Co., falls outside Olympic Nat. Pk. entrance, 15km. ne.
Quinault, P.H. Arnaud, Jr., IF; Asotin Co., Fields’ Spr. St. Pk., 15. VI. 72, W.J. Turner, 3F; Dewatto, 7. VI. 06, J.M.
Aldrich, IF; Chelan Co., Squillchuck St. Pk., 9mi. se. Wenatchee, 17, VI. 73, M. Jackson, IF; Lyle Grove, Pullman,
3. VI. 69, H.S. Telford, IF; ibid., 10. VI. 11, A.L. Melander, IF; Pullman, WSU Campus-bee area, 3.V.71, W.J. Turner,
1M; Pullman, nr. SCS Pond, WSU, 29.V.71, W.J. Turner, IF; Pullman, 9,1 1-12, 14, 19,28, 30.V, 1,4,6.VI.65, R.D. Akre,
4M, 9F; ibid., 7,18. V.66, 5-6,8, 15-17, 22,25-29.V, 1.VI.69, R.W. Dawson, 4M, 29F. (Deposited in CAS, USNM and
WSUP.)
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
PHYLOGENETIC INTERPRETATION
Introduction
In the ensuing pages phylogenetic aspects of Siphona species are considered. The approach I
have taken is cladistic, wherein character states are evaluated and polarized according to their
inferred plesiotypic (ancestral) or apotypic (derived) nature. Assuming only apotypic states to
be analyzable, these states are arranged in a nested pattern of hierarchies to produce a
cladogram (a diagram depicting a succession of apotypic character states representing
hypothetical phylogenetic relationships) most parsimonious with the data available.
Since publication of Hennig’s English version of his book “Phylogenetic Systematics”
(1966a), numerous practical accounts of cladism have been published (eg. Griffiths, 1972;
Ross, 1974; Eldredge, 1979; Wiley, 1981). The reader is referred to these sources for a detailed
exposition of the cladistic method of phylogenetic analysis used here.
The term monophyly is subject to different interpretations. As used here it excludes
paraphyly, being monophyly in Hennig’s sense (equivalent to holophyly of Ashlock, 1971).
Within the Siphonini a monophyletic group of genera is distinguished on the basis of two shared
derived character states (cf. Andersen, in press “b”):
1) anal vein extended to wing margin at least as fold (ground plan for the Tachinidae is anal
vein terminating in wing membrane)5, and
2) three katepisternal setae, of which postero-dorsal is strongest and antero-ventral is longer
than antero-dorsal (groundplan for at least the Siphonini is antero-dorsal seta longer than
antero-ventral).
This group is here referred to as the Siphona group, including the Old World genera recognized
by Andersen (in press “b”) and 3 genera endemic to the New World. Genera included are:
Aphantorhapha Townsend, Asiphona Mesnil, Ceranthia Robineau-Desvoidy, Pseudosiphona
Townsend, Siphona Meigen and Siphonopsis Townsend. Peribaea Robineau-Desvoidy is
excluded from this group, though it possesses state (1) above. If Peribaea is related to this
group, it is evidently sister genus to all the others. Andersen (in press “b”) hypothesizes that
Peribaea is the most primitive Old World siphonine genus, and this possibility is not excluded
here.
With the exception of Siphona, the genera of the Siphona group form an assemblage of
species of unclear relationships. Of these genera I have examined many undescribed species
from Middle and South America and suspect other inadequately collected areas also harbour
new species. Recognized genera do not bear well such scrutiny, and I predict a generic revision
will significantly alter their limits. Due to their similarities I refer to them collectively as
Ceranthia s. lat. to distinguish them from Siphona and to focus attention on their unsettled
status.
Siphona is probably monophyletic, as evidenced by proboscis structure. The proboscis is
long and geniculate, with the labella at least as long as prementum and at least (and in most
species considerably more than) eye height in length (Fig. 1,18-59).
Considered by Andersen (in press “b”) to be the ground plan state for the Siphonini, with
subsequent loss in the Actia group ( Goniocera B. & B., Ceromya R.D. and Actia R.D.).
North American species of Siphona Meigen
315
Most species of Ceranthia s. lat. have a labellar disc similar to that of a housefly. In North
American species in which it is elongate ( Siphonopsis and Pseudosiphona species in
particular), it can be distinguished by characteristics given in the “Reclassification” section.
Similarity between the proboscis of several Oriental and eastern Palearctic Actia species
and Siphona is also discussed in the “Reclassification” section.
A detailed comparison of male genitalia of Siphona species with those of species of
Ceranthia s. lat. will probably corroborate the monophyletic origin of Siphona species.
Currently, states of the major characters of male genitalia within taxa of the Siphona group
have not been polarized, so a cladistic analysis cannot be conducted. However, preliminary
studies suggest most genitalic features are plesiotypic in Siphona, even though the combination
of features that contribute to the general habitus of the male genitalia in that genus is unique.
A detailed study of the male genitalia of Ceranthia s. lat. taxa (including many undescribed
species) is needed to determine the synapotypic features of this character complex in Siphona
species. Such a study might also provide the necessary framework upon which to build a
classification for taxa of Ceranthia s. lat.
Character analysis
General notes. - I have recognized 15 characters as suitable for phylogenetic analysis of
North American Siphona species. Each is discussed below in relation to delimitation and
polarization of states. Difficulties peculiar to delimitation of mensural characters are discussed
under Character 1, eye height.
Out-group and in-group comparisons have been conducted to polarize character states. The
out-group is Ceranthia s. lat. (defined in the previous section). The in-group is world Siphona
species, as represented by specimens made available to me during the course of this study (see
world list for described species examined).
A character weighting system has not been used, as explained in the “Reconstructed
phylogeny” section.
One character useful for identifications has been excluded from the analysis. This character,
number of postsutural dorsocentral setae, is either 3 or 4, and in many species the number is of
diagnostic value. Both states are widely distributed in the out-group and in-group, so
consequently the apotypic state could not be determined; if it had been, then the number of
reversals necessary to account for its present distribution would probably have negated its
usefulness.
In the following section and in the character matrix (Fig. 102) an a represents the inferred
plesiotypic state. The first apotypic state is given an A. Other apotypic states not assignable to a
transformation series with the first are denoted by successive capitals letters; B, C, D, etc. The
order in which these are named refers only to their position in the matrix, starting from the top.
Transformation series are denoted by superscripts, negative numbers being used only with
mensural characters in which the plesiotypic state is inferred to be an average value (ie. eye
height).
Characters and states
Character 1 . Male eye height
This character and proboscis length are difficult to analyse because of their complex pattern
of inter- and intraspecific distribution. As such they represent transformation series of
phylogenetic importance, albeit without the relatively unambiguous delimitation of states
Quaest. Ent., 1982, 18 (1-4)
316
O'Hara
common to meristic characters. I have employed a practical solution to this problem:
delimitation of states based on apparent groupings about mean values (refer to Figs. 13,15),
with consideration of concordance with other characters for species of ambiguous placement.
The theoretical danger here is introduction of circularity into the analysis by allowing
perceived relationships among certain characters to affect delimitation of others. To offset this
departure from standard phylogenetic practice, I discuss doubtful species placements under
each species group. Despite this drawback I regard the characters eye height and proboscis
length as important to the elucidation of Siphona phylogenetics, and their treatment in the
manner indicated necessary for their interpretation.
The ranges given below for each eye height state are based only on consideration of species
under revision, in accordance with criteria discussed above. The plesiotypic state is inferred to
be an average-sized eye, as this state is the most widespread throughout the genus and across
species groups. Both small and large eyes are considered apotypic.
a - average-sized, mean between 0.76-0.82 head height.
A*2 - very small eye of S. lurida.
A-1 - small, though larger than eye of S. lurida , and less than 0.76 head height.
A1 - larger than average, between 0.82-0.85 head height.
A2 - very large, greater than 0.85 head height.
Character 2. Male first flagellomere
Six independent states are recognized in size and shape of male first flagellomere. The states
of this character are too varied in taxa of Ceranthia s. lat. to permit their polarization by
out-group comparisons. Since no state is distributed among examined species of world Siphona
in a manner suggestive of plesiotypy, all states are presumed apotypic. Relationships between
states are unknown (other than divisions of A, B and C), so they cannot be treated as a
transformation series.
A1 - narrow, average length to long, broadly rounded apically along ventral margin such that
dorsal edge is apically pointed.
A2 - shaped as in A1 though shorter.
B1 - relatively broad in profile, average length to long, dorsal and ventral surfaces parallel,
rounded or slightly truncate apically along ventral margin, not pointed as in A.
B2 - shaped as in B1 but very long.
C1 - narrow, short, rounded along ventral margin, not pointed.
C2 - shaped as in C1 but of average length.
D - triangular, apically truncate. Here S. hokkaidensis is treated separately according to the
shape of first flagellomere of eastern North American specimens (Fig. 31), though other
morphs resemble closely states C and F.
E - short and apically rounded.
F - very long, wide apically in profile and broadly rounded along ventral margin.
Character 3. Proboscis length
The plesiotypic state for Siphona at one time must have been a short proboscis, perhaps of
the sort exhibited by Siphonopsis species. To my knowledge no extant Siphona species has a
proboscis shorter than S. floridensis or S. maculata , so the truly ancestral state of the genus is
North American species of Siphona Meigen
317
no longer represented.
I have treated proboscis length as a transformation series from very short (plesiotypic) to
very long (apotypic), as polarized by out-group comparisons. A regression from a derived to less
derived state (ie. a reduction in proboscis length) is only inferred to have occurred in S', lutea
and S. akidnomyia.
An alternative exists to this polarization scheme. The S. maculata group is monophyletic,
based on shape of first flagellomere, and contains two character states interpreted as
symplesiotypies: a short proboscis and weak lateral marginal setae on Tl+2. Since both states
are virtually limited to that group, either of two explanations account for their restricted
distribution: (1) the S. maculata group branched from the main lineage of Siphona early in the
history of the genus, and retained to the present ancestral states which became modified in the
main Siphona lineage, or (2) the S. maculata group is relatively young, and states limited to it
are apotypic. Due to lack of corroborative evidence for the second hypothesis I consider the first
more parsimonious with available data.
Criteria for delimitation of character states is discussed under eye height.
a - very short, less than 1.55X head height.
A1 - short, mean between 1.55-1.8X head height.
A2 - slightly shorter than average, 1.8-2. IX head height.
A3 - average-sized, 2.1-2.5X head height.
A4 - long, 2.5-2.7X head height.
A5 - very long, greater than 2.7X head height.
Character 4. Aristomere 3
This character is represented by a transformation series which is easily polarized by
in-group comparisons. Species with the derived states are apparently monophyletic in Siphona,
for the states are in accord with other data. However, a few species in the ex-group (eg.
Ceranthia flavipes (Coquillett) and C. abdominalis (Robineau-Desvoidy)) also possess state
A1.
a - average length for Siphona , 0.50-0.65 head height.
A1 - shorter than average, 0.35-0.45 head height.
A2 - short as in A1 and thickened to near tip.
Character 5. Vibrissal angle
States were determined by in-group and out-group comparisons. Although the states seem to
represent a transformation series, no other evidence suggests the bearers of A1 and A2 ( S .
macronyx and S. lurida respectively) are closely related.
a - average condition, 2-3 setae and several hairs in addition to vibrissa.
A1 - 3-6 setae and several setulae and hairs in addition to vibrissa.
A2 - heavily setose, up to 10 setae and several setulae and hairs in addition to vibrissa.
Character 6. Male colouration
Colouration within the out-group is too varied to be useful in polarization of states in
Siphona.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
The most prevalent male colour pattern among Siphona species, and that least in
accordance with other characters, is predominantly blue-gray over most of thorax (where it is
mixed with light brown) and abdomen, the latter medially vittate and laterally yellow in ground
colour on T1+2 and T3. This state is regarded as plesiotypic, as are slight deviations from it.
a - given above.
A - abdomen black in ground colour, not vittate, deep blue-gray as basic pattern.
B1 - thorax dark in ground colour at least dorsally, abdomen partially to entirely yellow.
B2 - body yellow in ground and surface colour.
C - abdomen black in ground colour, not vittate, surface predominantly brownish-gray.
D1 - brown pruinosity over most of dorsal surfaces.
D2 - as in D1 only darker brown.
Character 7. Wing setulae on R
With varied frequency, R, wing setulae are present among species of a number of siphonine
genera. Except in a few genera their presence is evidently autapotypic. This is as true of the
genus Siphona as any other. There appears to be a predisposition for expression of wing setulae
on R, among these genera, so species sharing this trait are not necessarily closely related.
Among siphonines setulae on R] are expressed in one of three ways: (1) vein setulose along
its length, (2) limited to a row along the bend, or (3) represented by one or two setulae near the
bend. Most non -Siphona species with R, setulose belong to the first two categories. Siphona
bevisi Curran and S. setinerva Mesnil (both African) are in the first category, other examined
Siphona species belong to the third category - S. lutea, S. oligomyia, S. tropica and S.
grandistyla Pandelle (the last from the Pyrenees),
a - setulae absent.
A - one or two setulae near bend of R1.
Character 8. Wing setulae on R4+5
Wing setulae on R4+5 between bifurcation of R2+3 and R4+5 and crossvein r-m is
characteristic of the Siphonini, and represents the plesiotypic state in Siphona. As with R,
however, the apotypic state in Siphona has been independently derived in other genera and
presumably within Siphona as well. Of the species under revision only S. lutea has R4+5
setulose beyond r-m. Other Siphona species with this condition (of species examined) are S.
bevisi Curran, S. setinerva (Mesnil) and S. unispina (Mesnil) (all from Africa). The African
species resemble one another and further analysis may indicate they belong in the same group.
They do not share characteristics other than wing setulae with S. lutea , so are probably not
closely related to it.
a - vein R4+5 setulose between bifurcation of R2+3 and R4+5 and crossvein r-m.
A - vein R4+5 setulose beyond r-m.
Character 9. Male tarsal claws
The groundplan for Siphona is small tarsal claws. Large tarsal claws of males are present in
very few species of other Siphona group genera (eg. state A1 in Asiphona brunnescens (Vill.)).
These observations suggest the apotypic states, especially A1, are easily acquired and seldom
indicative of a monophyletic origin.
North American species of Siphona Meigen
319
a - average-sized, length of claw subequal to width of tarsomere 5 (Figs. 4,6).
A1 - intermediate in size between a and A2.
A2 - large claw, length subequal to length of tarsomere 5 (Fig. 5).
Character 10. Median marginal setae on T1+2
I am unaware of any Siphona group species not belonging to Siphona that possess a pair of
median marginals on T1+2. Nevertheless, consideration of more distantly related tachinids
reveals that median marginals on T1+2 are widespread, indicating their absence from most
members of the Siphona group almost certainly results from suppression of this attribute. This
may explain why a very few specimens of S. cristata and S. multifaria have median marginals
on T1+2, and why a few specimens of other species have slight development of a median
marginal hair on T1+2 such that it is distinctly stronger than others to either side.
a - absent.
A1 - a significant proportion of specimens with median marginals on T]+2 (here interpreted as a
regression from A2).
A2 - virtually all specimens with one pair median marginal setae on T, +2.
Character 1 1. Lateral marginal setae on T1+2
Most species of the Siphona group other than those belonging to Siphona have weak lateral
marginal setae on T1+2, although more distantly related tachinids in most species have strong
setae in this position. As with median marginals on T1+2, weak lateral marginals probably
result from suppression.
In Siphona weak lateral marginals on T1+2 are almost exclusive to species of the S.
maculata group. Since this state is widespread in the out-group, it is interpreted as
symplesiotypic in the S. maculata group (also see discussion under character 3).
a - setae absent or one pair weakly developed.
A - one pair strong setae.
B - two pair strong lateral marginal setae on T1+2 in significant number of specimens.
Character 12. Male sternum 5
Of male specimens examined, only two unrelated Siphona species, S. oligomyia and S.
cuthbertsoni Curran, possess a sternum 5 in which the antero-lateral arms are hooked inward.
This condition is characteristic of Siphonopsis species, which form either the sister group to
Siphona or branch from the Ceranthia lineage. Be that as it may, there is no other indication
that the hooked state of sternum 5 is plesiotypic in Siphona. In fact, the most parsimonious
explanation is independent evolution of the hooked state among Siphona species rather than its
loss in all but very few.
a - antero-lateral margins smoothly rounded (Fig. 10).
A - antero-lateral margins hooked inward (Fig. 1 1).
Character 13. Antero-ventral margin of distiphallus
By far the most prevalent condition of the distiphallus by out-group and in-group
comparisons is a dentate antero-ventral margin. This is presumed plesiotypic. The difference
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
between the dentate state and one in which small hooks are present (A1) is slight, hence is more
likely to occur in unrelated taxa than are states A2 and A3.
a - dentate.
A1 - small hooks (Figs. 75-77).
A2 - hooks medium in size (Fig. 60).
A3 - large hooks, margin U-shaped in anterior view (Fig. 62).
Character 14. Shape of distiphallus
The plesiotypic shape of the distiphallus in Siphona is unknown. I have chosen to regard all
shapes without apparent apotypic features as plesiotypic. This conservative approach should
ensure that species with the same apotypic shape have a high probability of sharing a uniquely
derived state.
A transformation series could not be discerned from distiphallus shapes exhibited, so each
apotypic state is treated independently.
a - without recognizable apotypic shape (ie. not necessarily plesiotypic, but treated as such).
A - subtriangular, apex sharply pointed in profile (Figs. 60,62).
B - anterior margin of distiphallus slightly indented in profile about one third of its length from
apex (Figs. 66-68).
C - anterior and posterior surfaces of distiphallus nearly parallel in profile, apex truncate (Figs.
70-71).
D - narrow in profile, sloped postero-ventrally (Figs. 9, 73).
E - narrow in profile, apically truncate (Fig. 74).
F - broad and short in profile (Fig. 76).
G - narrow in profile, subtriangular, apex pointed (Figs. 77-78, 80).
Character 15. Surstylus length
The most prevalent condition of the surstylus and one least in agreement with other
character states is presumed plesiotypic.
a - apex of surstylus even with or extended slightly beyond apex of cerci.
A - apex of surstylus extended beyond apex of cerci by more than preapical width of surstylus.
Reconstructed phylogeny
Introduction. - In reconstructing the phytogenies depicted in Figs. 103-107, I have not
employed a character weighting system. I recognize its advantages when applied to certain
types of data, as lucidly demonstrated by Hecht and Edwards (1976, 1977) for example, but
such a system is not appropriate to all situations. With respect to Siphona, the characters
analyzed are inadequate to permit elucidation of relationships among most species groups, so
where character weighting would have been most useful it cannot be applied. Furthermore,
with relatively few characters of broad import, the most parsimonious reconstructed phytogeny
for each species group has few alternatives. I have depicted only one phylogenetic scheme for
each species group, but I discuss problem species placements below, and consider the strengths
and weaknesses of the character states involved. Under these circumstances I consider the
utility of a character weighting system negated.
North American species of Siphona Meigen
321
Figs. 103-107 illustrate inferred relationships among North American Siphona species.
Exclusion of Palearctic species from these diagrams is not indicative of more distant
relationships between regions, for indeed the Siphona faunas of the Nearctic and Palearctic
Regions are intimately associated. My analysis of Palearctic Siphona species has not been
sufficiently detailed to warrant species placements beyond group. In general, Siphona species
from other regions have been considered to an even lesser extent.
The approach I have taken is thus:
1) inferred relationships among North American species are discussed under each species
group;
2) alternative phylogenies are considered;
3) the 1 1 described Palearctic Siphona species6 (excluding Holarctic species and S. seyrigi
Mesnil of the Canary Islands) are used to test my hypotheses concerning delimitation of
species groups; and
4) Siphona species from the Neotropical and Afrotropical Regions are considered where
their character states are relevant to the discussion.
The S. maculata group. - The ancestor to extant members of the S. maculata group is
inferred to have had a narrow, pointed first flagellomere, short proboscis and weak lateral
marginal setae on T,+2 (Fig. 103). The latter two states are interpreted as symplesiotypies (as
discussed above under polarization of states of proboscis length (character 3)), and identify the
S. maculata group as sister group to the other species groups.
Except for the apotypic shape of first flagellomere upon which the S. maculata group is
based (Figs. 18-22), there are only two synapotypies below the level of species pair. Therefore
relationships within this group are not well understood.
The only resolution is among S. maculata, S. intrudens and S. lurida. The latter two species
share four apotypies, clearly attesting to their close phylogenetic relationship. There are a
number of differences between S. intrudens and S. lurida , mostly because of the highly derived
condition of the latter.
S. maculata is interpreted as closely related to the S. intrudens - S. lurida lineage, because
of complex modifications of the distiphallus shared between S. maculata and S. intrudens
(characters 13 and 14 (Fig. 102), Figs. 60,62). Since S. lurida lacks these modifications (Fig.
63), the plesiotypic states of its distiphallus evidently represent losses (thus the coding of 13-a
and 14-a as derived in S. lurida in Fig. 103).
Distribution of character states are inadequate to resolve the trichotomy depicted for
lineages of the S. maculata group (Fig. 103). The autapotypic states of S. pacifica result in its
general similarity to plesiotypic members of other Siphona groups. However, there is no
evidence to question placement of S. pacifica in the S. maculata group.
The phylogenetic position of S. floridensis within the S. maculata group is also enigmatic.
This species has the fewest recognized apotypic states of species analyzed. Therefore, the
overall similarity between S. maculata and S. floridensis in external characteristics is the
result of symplesiotypies, which are unanalyzable cladistically. Also, to consider 5. maculata
and S. floridensis as sister species, one must regard the plesiotypic states of the distiphallus in
the latter (Fig. 61, cf. Fig. 60) as losses (as inferred for S. lurida ). Nonetheless, a close
relationship between these species is tenable on zoogeographic grounds. A zoogeographic
The new, as yet unpublished, Siphona species of Andersen (in press “a”) were not examined.
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scenerio whereby S. floridensis might have evolved from a S. maculata population during the
Quaternary is outlined in the “Zoogeographic” section. Pending further morphological
evidence, I leave the trichotomy in Fig. 103 unresolved.
Two Palearctic species have been identified as members of the S', maculata group: S. collini
Mesnil and S. efflatouni Mesnil. They are the only Palearctic Siphona species with weak
lateral marginal setae on T1+2. They possess an A1 type first flagellomere and are the only
Palearctic species with a short proboscis (not longer than A2). The distiphallus of S. collini has
small hooks along the antero-ventral margin (state A1). (Only the female holotype of S.
efflatouni was examined, so the distiphallus of this species was not studied.) In general habitus
S. collini and S. efflatouni also appear to belong to the S. maculata group.
Despite the plesiotypic nature of weak lateral marginal setae on T1+2 and a short proboscis,
both states are well correlated with the apotypic shape of first flagellomere in the S. maculata
group. As such these characteristics are useful indicators of S. maculata group members, even
though they are interpreted as symplesiotypies.
A test I have not had an opportunity to perform to date involves use of undescribed Oriental
Siphona species H. Shima notes (per. comm.) several species similar in appearance to S. lurida
from the Orient, and study of these would undoubtedly cast light on the phylogeny presented
here. It will be particularly interesting to note whether these species are more closely related to
S. lurida or S. intrudens, since the implications critically affect the zoogeographic
interpretation as well as the reconstructed phylogeny of the S. maculata group.
The S. cristata group. - The S. cristata group is based on one apotypic state, shape of first
flagellomere (Fig. 104).
S. oligomyia is included in this group, though its first flagellomere is not characteristic of
the group (possessing state 2- A instead of 2-B). Justification for inclusion of S. oligomyia in the
S. cristata group is based on two synapotypies between S’, lutea and S. oligomyia , as discussed
below.
At the next higher level on the cladogram (Fig. 104) is shape of distiphallus (state 14-B),
uniting the S. cristata - S. multifaria and S. lutea - S. oligomyia lineages. Again, S. oligomyia
does not share the derived state of the other members.
S. pisinnia is treated as sister species to the other species of the S. cristata group, though it
is actually not as distantly removed phenetically from the rest of the group as it appears in the
cladogram. Only three states separate it from S. multifaria , and one of these, distiphallus shape
(Fig. 65; cf. Fig. 67), is a consequence of the system I employed to delimit states. I have
recognized the indentation of the anterior margin of the distiphallus as an apotypic state, while
necessarily relegating forms near this shape without an indentation to the plesiotypic state, for
lack of distinguishing features. Because of the highly derived appearance of S’, lutea and S.
oligomyia , S. pisinnia is phenetically closer to S. multifaria and some specimens of S. cristata
than are the others.
Within the S. cristata group I have depicted S. multifaria and S. cristata as sister species
(Fig. 104). Under this scheme I have not followed the apparent transformation series of
character 6 (colouration), which suggests S. cristata is closer to S. lutea than to S. multifaria.
The distiphalli of S. cristata and S. lutea are also very close (Figs. 66, 68). Nevertheless,
characters 3 and 7 argue in favour of the depicted arrangement.
Additional support for the reconstruction comes from the Palearctic species S. flavifrons
Staeger. It is similar in colouration to the yellow form of S. cristata , yet probably belongs to the
S. geniculata group. This is evidence that the B1 colouration of S. cristata need not have shared
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323
an ancestry with B2 colouration of S. lutea and S. oligomyia.
Character state 6-B2 is highly apotypic. Through concordance with state 7-A, S. lutea and
S. oligomyia are well established as sister species. S. oligomyia is distinctive because of many
autapotypic features; had it been of average colouration its species group placement would be
uncertain.
A Palearctic species of dubious placement is S', setosa Mesnil. It has median marginal setae
on T1+2> but none of the other S. geniculata group apotypic characteristics. With its 2-B first
flagellomere, it might belong to the S. cristata group. Other characteristics are inconclusive:
distiphallus type 14-a, proboscis and eye average-sized, colouration average, etc. I leave S.
setosa unplaced.
Two Palearctic species are easily recognized as members of the S. cristata group. One, S.
confusa Mesnil, has an average-sized eye and proboscis and a first flagellomere varied between
states A1 and B1. The one distiphallus examined is close to state 14-B, but without an
indentation anteriorly. A few specimens have a tawny abdomen as in S. cristata , while others
are of average colouration.
The other Palearctic species, S. pauciseta Rondani, has an average-sized to A1 eye and a
proboscis of average length. First flagellomere and distiphallus possess the characteristics of the
group, with states 2-B1 and 14-B.
S. confusa and S. pauciseta do not share the apotypic states of S. lutea and S. oligomyia.
Their phylogenetic position relative to S. pisinnia , S. multifaria and S. cristata is unclear. As
members of the S. cristata group, S. confusa and S. oligomyia illustrate a deficiency in the
distinction between states A and B of first flagellomere. These states are not markedly
different, and both species indicate that 2-A is derivable from 2-B. Unfortunately, without
additional characters there is no alternative to 2-B1 as a synapotypy for the group.
S. confusa also illustrates a problem in delimitation of character 14; distiphallus shape. S.
pauciseta supports recognition of 14-B as a good synapotypic state, yet S. confusa , like S.
pisinnia , lacks this state. To incorporate S. confusa and S. pisinnia , S. medialis, S. floridensis
and S. pacifica would also have to be included, thereby considerably reducing the usefulness of
this uniquely derived state. A subjective element thus enters the analysis, for resemblance of a
type 14-a to a 14-B distiphallus is not without significance.
Regardless of the difficulties inherent in recognition of state 14-B, it is possible that
relegation of S. pisinnia to its depicted position is phylogenetically inaccurate. An alternative is
to disregard the slight distiphallus differences beween S. pisinnia and its relatives, and examine
the consequences to the cladogram. A trichotomy results, with the S. pisinnia lineage on the
same node as the S. cristata - S. multifaria and S. lutea - S. oligomyia lineages. Given this
possibility, similarity between S. pisinnia and S. multifaria suggests 51. pisinnia might be more
closely related to the S. cristata - S. multifaria lineage than to the other. To support this
alternative hypothesis (and refute the relationship shown in Fig. 104) a synapotypy is required,
and one has not yet been found.
The S. cristata group would benefit more than others from analysis of additional characters.
Possibilities include larvae, ethological and host data, differences in the reproductive system of
females, electrophoresis and cytotaxonomic data. As illustrated above, external features and
male genitalia are not particularly amenable to phylogenetic analysis.
The S. geniculata group. - In North America the S. geniculata group includes three species
(Fig. 105), one of which ( S . geniculata ), has been introduced by man to this continent. These
species share the derived states of one pair median marginal setae on T1 + 2 (albeit not all S.
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hokkaidensis specimens) and a short aristomere 3 (Figs. 28-32).
The apotypic states shared between S. geniculata and S. hokkaidensis attest to their close
phylogenetic relationship, even though most specimens of the latter are without either median
marginals on T1+2 or a type 2-A1 first flagellomere (= antennal type 1, Fig. 30).
Five Palearctic species are assignable to the S. geniculata group, though their
characteristics are varied. Only two species have median marginal setae on T1+2, S. flavifrons
Staeger and S. rossica Mesnil. S. rossica is evidently close to S. hokkaidensis , with its
medium-sized eye and proboscis, first flagellomere near type 2-D, two pair lateral marginal
setae on T3 and a short aristomere 3. I did not have an opportunity to dissect a male specimen,
but I predict the male genitalia of this species has a long surstylus.
I recognize S. flavifrons as a member of the S. geniculata group by its joint possession of
median marginals on T,+2 (most specimens) and a long surstylus. Its distiphallus is apically
truncate, but slightly too wide basally to be classed type 14-C.
S. flavifrons does not have a shortened aristomere 3 and its first flagellomere and
colouration are as in S. cristata. In light of other character state differences I view these
similarities as convergences. The characteristic shape of first flagellomere for the S. cristata
group has already been shown to be weakly based. I regard the average-sized aristomere 3 as a
significant departure from the condition characteristic of the S. geniculata group, for to my
knowledge S. flavifrons is the only species with median marginals on T1+2 and a long surstylus
for which the aristomere 3 is not shortened.
S. paludosa Mesnil lacks median marginals on T1+2, but has a short aristomere 3, type 14-B
distiphallus and a 15-A surstylus, strongly supporting its placement in the S. geniculata group.
I also include two other Palearctic species without median marginals on T1+2, S.
grandistyla Pandelle and S. nigricans (Villeneuve). Both have a first flagellomere resembling
the 2-D state of S. hokkaidensis , and a short aristomere 3. Unfortunately I did not have an
opportunity to dissect male specimens of these, which would have provided a good test of their
placement.
The only Palearctic species with median marginal setae on T1+2 that I exclude from the S.
geniculata group is S. setosa Mesnil. It is discussed under the S. cristata group.
I have examined specimens from Ecuador of three undescribed Siphona species with median
marginal setae on T1+2. In other respects these species are quite unlike members of the S.
geniculata group (in particular, they lack states 4-A and 15-A), so I exclude them. Discussion
of these species and their affinities more appropriately awaits a revision of South American
Siphona species. I mention them here to demonstrate that not all species with median
marginals on T1+2 belong to the S. geniculata group.
Examined Afrotropical and Oriental Siphona species do not have median marginals on T1+2
(see world list for species examined).
Despite lack of unequivocal character states for the S. geniculata group, concordance
among states functions well for distinguishing between included and excluded species.
Consideration of Palearctic species illustrates that the S. geniculata group is a complex
assemblage, yet evidently monophyletic.
The S. futilis group. - The apotypic character state for this group is a large eye, best
exemplified by the 1-A2 state in S. futilis and S. illinoiensis (Figs. 1,34). Similarity between S.
futilis and S. illinoiensis is such that I can polarize but one difference, length of first
flagellomere.
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325
I cannot with certainty place S. brunnea in the S'. futilis group. In general habitus it
resembles S. futilis and S. illinoiensis despite its autapotypies, so I include S. brunnea in this
group until such time as its relationship is clarified (Fig. 107).
With S. brunnea a provisional member, and only one apotypic (and not unique) state for the
group, consideration must be given to states of characters 1, 2 and 14 in placement of
non-North American species (Fig. 102).
The Palearctic species S. boreata Mesnil might belong to the S. futilis group. It has a large
eye, average length proboscis (characteristic of the group, but evidently plesiotypic), and
average colouration. However, first flagellomere is near type 2-F. I did not have an opportunity
to study male genitalia of this species, thereby limiting my analysis.
Examined specimens of at least two (possibly undescribed) South American species
probably belong to the S. futilis group, study of which might help define the group and clarify
the position of S', brunnea.
The S. tropica group. - This group is particularly noteworthy because it is well represented
in the Afrotropics. Of Afrotropical species examined, S. unispina (Mesnil) and S. gracilis
(Mesnil) are the most easily recognized as members. I hesitate to assign to this group other
Afrotropical species, since the characteristics that readily distinguish members in Middle
America are widespread among Afrotropical Siphona species and not restricted to the S.
tropica group.
I have used character states of S. unispina to help polarize states within the S. tropica
group. I have assumed the Middle American species to be more closely related among
themselves than to S. unispina. As a result, states of characters 13 and 14 have been polarized
differently than they otherwise would have been (Fig. 106).
In conflict with my understanding of the S. tropica group is character state 8- A: setulae on
R,. A single setula is present near bend of R, in S. tropica and S. unispina. Because it is absent
from examined specimens of S. rizaba, S. longissima and S. akidnomyia , I suggest it has been
independently derived as in other Siphona species ( S . lutea + S. oligomyia , S. grandistyla). S.
tropica could be postulated as closer to S. unispina than to the others, but this would not
explain the presence of hooks on the distiphallus of S. longissima and S. akidnomyia (Figs.
76-77), nor their other S. tropica group characteristics. Two separate ancestors could be
invoked to explain the anomaly, but are not justified on the basis of a single low weight
apotypy.
The shared characteristic between S. longissima and S. akidnomyia is a weak synapotypy.
Additional data are needed to elucidate relationships among S. tropica group members.
The S. tropica group is apparently absent from the Nearctic and Palearctic Regions. In the
Neotropics it is most diverse in Middle America. Other than S. tropica , I have examined one
species (probably undescribed) of this group from South America (Chile) and one from the
Philippines.
The Siphona fauna of Middle America is inadequately known. I have examined a single
specimen of a new species belonging to the S. tropica group that I have not described due to its
poor condition. I suspect several other Siphona species are undiscovered or undescribed from
this region as well.
The S. macronyx group. - S. macronyx is placed in a monobasic species group because its
phylogenetic position relative to other North American Siphona species has not been clarified
(Fig. 103).
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The large tarsal claws and above average number of setae and setulae on vibrissal angle of
S. macronyx are derived states shared with S. lurida of the S. maculata group. This species
also has hooks on the distiphallus, a derived state for the S. maculata, S. floridensis, S.
intrudens and S', lurida lineage. Nevertheless, a sister species relationship between S.
macronyx and S. lurida is untenable in light of their overall dissimilarity (Figs. 21,33) and the
synapotypies between S. intrudens and S. lurida (notably a small eye, short proboscis and
similar colouration). Furthermore, S. macronyx lacks the slender, pointed first flagellomere,
which is apotypic for the S. maculata group.
There is a slight similarity between S. macronyx and the S. geniculata group (characters
2,3 and 9 (Fig. 102)), but S. macronyx lacks two of three apotypic states upon which the group
is based.
Evolutionary trends
The most satisfactory method for obtaining information about most characters of varied
structure in Siphona adults is to observe living specimens under natural conditions. Though I
did this intermittently, I did not pursue it intensely. It has been my experience that adults of
Siphona searching for a mate or host are usually seen fleetingly, and in many instances rarely.
On the other hand, adults seeking nourishment can be observed for some time as they probe
deeply into flowers with their long proboscises, in quest of nectar. As a result, the functional
significance of most derived characteristics can only be gleaned indirectly through study of
dead specimens and by analogy with other taxa.
It is a basic tenet in evolutionary studies that most modifications in structure result from
selective pressures and have functional significance. If so, the most derived and varied feature
of Siphona adults, the proboscis, must have attained its present variety because of advantages
conferred on its bearers.
The only known function of the proboscis in Siphona adults is for procurement of food;
chiefly nectar from flowers. Unlike large tachinines, adults of most siphonines are too small to
force apart flower components to reach deep nectaries, so the benefits of a long proboscis are
obvious. Not only did adults of Siphona apparently evolve an elongate prementum and labella
to aid feeding, but so too did others, such as adults of certain species of Ceranthia s. lat. and
Actia. Adults of Clausicella achieved essentially the same end through elongation of the
prementum, albeit with less maneuverability distally than that provided by a geniculate
proboscis. In fact, an elongate prementum is found in various groups throughout the
Tachinidae, though an elongate labella is apparently unique to the Siphonini.
It is not possible to correlate development of a geniculate proboscis in adult Siphona with
success of the taxon. For one reason, certain genera like Ceromya, Peribaea and most species of
Actia have a short labellar disc, yet approach the diversity of Siphona. For another, too little is
known about parasite-host interactions to dismiss as minor, developments in that area.
However, it is possible that individuals with a long proboscis have a selective advantage over
others in varied environments, for they are physically less dependent upon certain kinds of
flowers than are their relatives with shorter proboscises. Few Siphona species other than
members of the primitive S. maculata group have a short proboscis, so once attained a long
proboscis seems seldom to have shortened (I have inferred just two such reductions; in S.
oligomyia and S. akidnomyia). Therefore a definite trend in adults of Siphona species from a
short to long proboscis is evident (cf. Fig. 15).
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327
There are no other trends in Siphona as unidirectional as that of proboscis length. Greatly
varied is eye size, but from the inferred plesiotypic state of an average-sized eye, both small and
large eyes are necessarily derived. Shape and size of first flagellomere are also considerably
varied (even within species), and a trend is not apparent. The significance of varied eye and
first flagellomere sizes among species is unknown, and speculation is best deferred until more
information on behaviour and environment of individual species is available. Within species,
males have the larger eye and first flagellomere, implying both are used in the search for a
mate. Females are probably mostly host oriented, and the slightly longer palpus of females in
most species might be involved in determining host suitability.
Notable in the evolution of Siphona species is the similarity among species groups in mean
number of apotypic states (Fig. 102). Though the S. maculata group is interpreted as the most
primitive, two included species ( S . intrudens and S. lurida) are very derived, so even this group
is not markedly removed from the others in degree of apotypy. This pattern implies that
evolution within Siphona has progressed at a relatively constant rate, without invasion of, and
diversification in, new adaptive zones.
ZOOGEOGRAPHIC CONSIDERATIONS
Diversity and general paterns
I prefer not to use the terms Nearctic and Neotropical in discussing New World Siphona
species distributions. Most Middle American Siphona species are not derived from extant
Nearctic or South American species groups, so to emphasize this point I. discuss distribution
patterns in terms of America north of Mexico, Middle America (Mexico to Panama) and South
America. Only two species are not confined within boundaries of one of these areas. S. pisinnia
(Fig. 83) ranges widely in America north of Mexico and Middle America. The distribution of
S. tropica (Fig. 101) extends into extreme northwestern South America, but for present
purposes it will be regarded as a Middle American species.
My use of terms Palearctic, Oriental and Afrotropical were discussed in the preamble to
“World list”. I treat S. geniculata as a Palearctic species in the ensuing discussion because in
North America it is an introduced species.
The 79 described Siphona species7 are distributed as follows, by region in order of
decreasing diversity:
Afrotropical - 40 species.
Palearctic - 17 species.
America north of Mexico - 14 species.
Middle America - 7 species.
South America - 3 species.
Oriental - 3 species.
Madagascar is included in the Afrotropical Region. It has three endemic species and five
others that are shared with the mainland. The Canary Islands off the northwest coast of Africa,
with endemic species S. seyrigi , is considered Palearctic. So too is Africa north of the Sahara,
from whence S. nigricans was described (from Algeria).
Excluding S. maculipennis Meigen, an unrecognized Palearctic species that is probably
conspecific with a recognized species.
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Four species (excluding introduced species S. geniculata ) are known to be shared between
America north of Mexico and the Palearctic Region: S. cristata, S. hokkaidensis, S. lurida and
S. maculata. The Palearctic Region is not known to share species with any other region.
America north of Mexico has S. pisinnia in common with Middle America.
The Siphona fauna of South America is inadequately known. I estimate from the little
material I have examined from the region that at least ten species live there, possibly more.
I have examined adults of one undescribed species of Siphona from Australia. Siphona
species from the Oriental Region are not well known and are probably more diverse than
indicated by three described species.
In the previous section a cladistic analysis helped define six North American Siphona
species groups. Within each group there is a general pattern of geographic distribution
involving at least two regions. Three general interregional patterns are recognizable among the
groups:
1) America north of Mexico - Palearctic Region.
This pattern is evidenced by a close relationship between species of both regions, with a high
proportion of Holarctic distributions for species with northern records (three of six New World
species). Representing this pattern are the S. maculata, S. cristata and S. geniculata groups.
2) America north of Mexico - Middle America.
Here belongs the S', futilis group, wherein closely related S. illinoiensis and S. futilis are in
adjacent regions. There is evidently a closer affinity to the fauna of South America than the
Old World, but this has not been firmly established.
3) Middle America - Afrotropical Region.
In the New World the S. tropica group is most diverse in Middle America. No species has been
assigned to the group from America north of Mexico or the Palearctic Region. Although
represented by at least one species in South America and one in the Philippines, the
zoogeographically most significant relationship is between Middle American and African
members of the group.
Within North America another set of distribution patterns, those involving species rather
than species groups, is evident. To a large extent these reflect biotic changes that have taken
place in the region since the Middle Tertiary. These are:
1) Transcontinental (mostly cool to cold temperate) - S. cristata, S. hokkaidensis, S. intrudens,
S. lutea, S. maculata, S. medialis and S. multifaria.
2) Western - S. geniculata (introduced), S. lurida, S. macronyx, S. oligomyia, S. pacifica and
S. pisinnia.
3) Eastern - S. floridensis and S. illinoiensis.
4) Mexican - S. akidnomyia, S. brunnea, S. futilis, S. longissima, S. rizaba and S. tropica.
Historical zoogeography
It is premature to speculate in detail about historical events responsible for present day
Siphona species distributions in North America. The three main distribution patterns identified
in the previous section make obvious the fact that the North American assemblage of Siphona
species is polyphyletic. Indeed, interregional relationships are so complex that the monobasic S.
macronyx group is the only one that is exclusively North American.
The best zoogeographic hypothesis is one that stems from a thorough cladistic analysis of a
group. Only then are all requirements of a scientific hypothesis met: explanatory power,
internal consistency, predictive power and testability (falsification) (Ball, 1976; Noonan,
North American species of Siphona Meigen
329
1979). As a consequence the cladograms in Figs. 103-107 (representing only North American
species) are inadequate for detailed interpretation of the historical zoogeography of North
American Siphona species.
Siphona species groups, on the other hand, are suitable for zoogeographic analysis because
they are based on inclusive and not (as for North American species) exclusive data. Siphona
species groups are monophyletic units, members of which need not be cladistically arranged for
the groups to be zoogeographically analyzed, provided regional group relationships are known
(Hennig, 1966b). For instance, the very close phylogenetic relationships among taxa of the S.
maculata group between America north of Mexico and the Palearctic Region are indicative of
past faunal exchange between regions. This presumed interchange satisfies the four
requirements of a scientific hypothesis, so a zoogeographic analysis seeking a historical
explanation for the interchange is well founded.
Ideally all Siphona species groups and data about intergroup affinities should be included in
a zoogeographic treatment of the genus. It may be many years before sufficient information on
Siphona cladistics is gathered to realize this goal. Therefore I judge it timely to interpret
aspects of Siphona history from the data available now.
In the absence of a fossil record there is an alternative means for deriving age of a taxon.
The minimum age is that of the earliest demonstrable vicariant event within the taxon.
Maximum age is that already determined for a higher taxon to which the taxon in question
belongs, or as evidenced by the taxon’s global distribution pattern (provided the taxon is
widespread). For the Calyptratae in general the early fossil record is inadequate; they have
been assumed pre-Tertiary in age for many years (eg. Rohdendorf, 1974), but not until 1970
was the first calyptrate fly from the Mesozoic (Upper Cretaceous, 70 mybp') recorded
(McAlpine, 1970). To date, a fossil tachinid has not been discovered that has a significant
bearing on the age of origin of this family (van Emden, 1959), much less Siphona.
It has become possible in the past decade to outline in a broad sense major geologic and
climatic events of the earth since early Mesozoic time. Concomitant with this knowledge has
developed a general willingness on the part of biogeographers to interpret species distributions
according to established geologic and climatic theory. Fossils remain valuable means of
hypothesis testing, but are no longer central to most biogeographical analyses unless the fossil
record is unusually complete. Recent studies of Diptera reflect this trend. For example, Bowden
(1973) suggests a Late Cretaceous origin of Dischistus (Bombyliidae) based on distributional
data. Papavero (1977), places origin of the Oestridae between Late Jurassic and Early
Cretaceous on the basis of a Gondwanian distribution. Quite remarkable is McAlpine’s (1977)
suggestion of a Gondwanian distribution for an extant piophilid genus, Protopiophila (the first
Gondwanian distribution recognized for a genus of the Schizophora).
By analogy with studies on other Schizophora the Tachinidae are apparently pre-Tertiary in
origin. Provided distributional data suggest a more recent origin of Siphona than analogy has
for the Tachinidae, then there is no age conflict between them9.
Although Siphona species are cosmopolitan, they show no indication of a Late Cretaceous
or earlier origin. Had Siphona evolved in the Southern Hemisphere during the Late Cretaceous
8million years before present.
9as yet a time scale has not been proposed for origin of tachinid subfamilies and tribes, so these
taxa cannot be discussed here.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
a Gondwanaland distribution would be expected (assuming extinction was not a factor). By
Late Cretaceous Africa was drifting north, while South America, Antarctica and Australia
were still connected via at least an archipelagic route (Dietz and Holden, 1970; Smith, Briden
and Drewry, 1973; Howarth, 1981). In fact, the southern route through Antarctica may have
been in use by terrestrial animals as late as Early Oligocene (Raven and Axelrod, 1975).
An adequately supported argument against a Gondwanaland distribution for Siphona must
await a cladistic analysis of African, South American and Australian species. However, the few
Siphona species in South America and especially Australia, relative to Africa and the northern
continents, suggests dispersal of Siphona species was not taking place between southern
continents in the Late Cretaceous.
In the Northern Hemisphere during the Late Cretaceous, high sea levels had a greater
isolating effect on continents than did plate positions. Across the North Atlantic and through
Beringia, land (?filter) bridges were apparently in operation, though north-south seaways
hindered intracontinental faunal movements (Kurten, 1969; Hallam, 1973; Matthews, 1979;
Howarth, 1981).
If Siphona species were in the Northern Hemisphere during the Late Cretaceous I would
expect a more pronounced difference between Siphona faunas of America north of Mexico and
the Palearctic Region than evident in the S. maculata, S. cristata and S. geniculata groups.
Presumably Siphona species could have dispersed readily between northern regions had they
been there during the Late Cretaceous. Yet present faunal similarities are better explained by
Late Cenozoic exchange.
Based on available evidence and without comparable studies within the Tachinidae, I
hypothesize a maximum age of Siphona as Late Cretaceous. As mentioned earlier, minimum
age necessarily precedes the earliest demonstrable vicariant event. Before discussing how I
apply this principle to Siphona , it is appropriate to consider region of origin of the genus. This
simplifies the discussion further on.
I emphasize the term region of origin, not center of origin. Thus I am not concerned with the
controversy between centers of origin in the Darwinian sense and vicariance biogeography
(Croizat et al., 1974). In a region of continental size, taxa can evolve under either paradigm
and produce equal diversity, so the distinction is not important here.
Siphona probably arose in the Old World. The basis for this inference is twofold. First, Old
World Siphona species and species of siphonines related to Siphona ( Actia , Ceromya and
Peribaea spp.) considerably outnumber New World taxa. Secondly, an invasion of North
America from the Old World during the Tertiary by Siphona is compatible with New World
species distributions, while it is inconceivable without a number of assumptions that invasion of
Siphona into the Palearctic Region from North America would lead to the present patterns of
diversity and distribution in the Old World.
To account most parsimoniously for Old World Siphona patterns, Africa must be
interpreted as region of origin for the genus. Diversity favours this choice, but does not negate
the possibility of adaptive radiation in Africa with an origin elsewhere. More important than
diversity is the pattern of diversity, as determined through comparison of species numbers
among regions.
An African origin of Siphona is compatible with the observed decrease in number of species
from a high in the Afrotropical Region to a low in the Oriental and Australian Regions.
If the Palearctic Region were postulated as area of origin of Siphona , the observed pattern
of diversity would be unparsimonious. During most of the Tertiary, Africa was isolated from
North American species of Siphona Meigen
331
other regions by the Tethys Sea (Cooke, 1972; Hallam, 1973; Azzaroli and Guazzone, 1979;
Howarth, 1981). It is unlikely Palearctic Siphona individuals crossed this barrier enough times
to attain the present Afrotropical diversity in number of species and species groups without
similarly invading the more accessible Oriental Region.
Returning to the question of minimum age of Siphona , I limit my analysis to species groups
in which cladistic relationships between regions have been determined. These are the S.
maculata, S. cristata, S. geniculata and S. tropica groups. The first three do not reflect a
necessarily old distribution pattern. The S. tropica group is disjunct between Middle America
and Africa, and it is from this group I hypothesize minimum age of Siphona.
Concomitant with historical biogeographic hypotheses are assumptions. Assumptions should
be formalized, for they enhance the central hypothesis by clearly distinguishing the assumed
from the inferred (cf. Lewis, 1980: 554). Furthermore, while some assumptions are untestable
(and are, for that reason, assumptions) others result from insufficient data and become
falsifiable as new data are added. Assumptions of the latter type should be accompanied by
statements about their testability, to make the central hypothesis more scientific.
Assumptions upon which I base my analysis of the disjunction of the S. tropica group
between Middle America and Africa are:
1) The cladistic relationship between Middle American members of the S. tropica group and S.
unispina in Africa (Zaire) has been correctly interpreted. (Falsifiable by cladistic study of
additional taxa or reinterpretation of existing data.)
2) Members of the S. tropica group in South America are derived from Middle American
species; they are not Gondwanaland relicts. (Relationships testable by cladistic analysis.)
3) Africa was region of origin of Siphona and the S. tropica group. (Subject to corroboration or
refutation through study of world species groups of Siphona and closely related genera.)
4) Disjunction in distribution of the S. tropica group is not a result of long distance dispersal
between Middle America and Africa. The dispersal ability of Siphona adults is attested to by
their presence on islands: Madagascar, Canaries, Juan Fernandez, Tierra del Fuego,
Philippines and others. There are eight species recorded from Madagascar, of which five are
shared with Africa. Presumably the Mozambique Channel (500km wide) has not been an
efficient barrier to dispersal of Siphona individuals. There is no evidence, however, that
individuals cross wide barriers. For example, Siphona is not recorded from Hawaii. I suspect
water gaps are more easily crossed than other physical or climatic barriers.
5) Current hypotheses of paleoclimate and paleogeography relevant to my interpretation of past
events are accurate. (Testable by results of future research.)
6) Climate and habitat requirements of S. tropica group taxa have not changed significantly
throughout their existence. (Discovery of a S. tropica group species in a cold temperate
environment would cast doubt on this assumption.)
An explanation is sought that accounts for the geographically disjunct nature of the S.
tropica group. Assuming the genus originated in Africa subsequent to the Late Cretaceous,
direct dispersal across the Atlantic is considered improbable because South America and
Africa were widely separated by that time. The only available land route was northward to
Europe or Asia and into North America through Beringia or across a North Atlantic land or
filter bridge.
First consider the ecological aspects involved in a Northern Hemisphere route. The S.
tropica group is now restricted to the Tropics, so existence of northern land bridges is not
sufficient to account for Siphona migration into North America. Had a S. tropica group taxon
Quaest. Ent., 1982, 18 (1-4)
332
O’Hara
invaded the Northern Hemisphere from Africa during a relatively cool period, descendants
would presumably still be there. If instead there was a period of significant warming in the
north, a S. tropica group taxon could have entered Eurasia and spread to North America
without a major shift in way of life. If deterioration of the warm trend took place soon after the
ancestral New World Siphona taxon arrived in North America,' then retreat to Middle
America could have occurred without leaving northern, cool adapted descendants. These events
could not have occurred later than the Late Eocene (34 mybp) (Wolfe, 1978, 1980).
For many years the Geoflora10 concept dominated paleobotanical interpretations. Paleocene
and Eocene climates of northern North America were interpreted as subtropical to warm
temperate with gradual cooling from the Oligocene to Quaternary (Dorf, 1964). The Geoflora
concept has recently come under incisive criticism with the realization that plant species
respond individualistically to climatic shifts (Graham, 1972; Wolfe, 1978). Nowhere is this
better exemplified than in late Quaternary macrofossil records of packrat middens (Wells,
1979; Van Devender and Spaulding, 1979).
Recent studies substantiate fluctuating warm periods from the Early Tertiary to Late
Eocene but envision much warmer conditions than the Geoflora model, followed by rapid
climatic deterioration (Wolfe, 1978, 1980; Matthews, 1979). Paratropical" conditions are
thought to have prevailed at higher latitudes over most of the Northern Hemisphere (ibid.).
During this period the most suitable conditions of the Tertiary must have existed for entry of a
S. tropica group taxon into North America.
Physical aspects of a S. tropica group taxon dispersing from Africa into Eurasia and then to
North America prior to the Late Eocene is complicated by two factors. First, the Tethys Sea
isolated Africa from Eurasia during most of the Tertiary, such that faunal exchange was
primarily limited to brief periods of (?filter) connection in the Late Oligocene, Middle and
Late Miocene and Late Pliocene (Cooke, 1972; Hallam, 1973; Azzaroli and Guazzone, 1979).
It is not precluded that other connections of very short duration occurred during which time
Siphona could have entered Eurasia. Alternately, Siphona might have crossed the water
barrier directly, by chance dispersal (a sweepstake route, such as between Madagascar and
Africa). By whatever means a S. tropica group taxon reached Eurasia, timing was apparently
not coincident with a period of substantial mammalian exchange. More important to my
hypothesis than Siphona crossing the Tethys Sea at the most opportune time is a S. tropica
group taxon reaching the climatic corridor before it closed in the Late Eocene.
The second complication pertaining to a Siphona taxon reaching North America is the
Turgai Strait separating Europe and Asia during the Early Tertiary (Adams, 1981). Siphona
could have entered Europe by a route through Gilbraltar and crossed a North Atlantic land
bridge (Matthews, 1979) without ever reaching Asia. Conversely, dispersal could have been via
Asia and Beringia. At least the latter route seems highly probable because the existence of a S.
tropica group member in the Oriental Region (the Philippines) implies that the Turgai Strait
did not prevent the group from dispersing into Asia.
At the end of the Late Eocene, the climate in northern regions cooled rapidly (Wolfe, 1978,
1980, as the “terminal Eocene event”). Because S. tropica group taxa are not known from
America north of Mexico or the Palearctic Region, I suggest they were unable to adapt and so
10a Geoflora is a flora that undergoes little compositional change over a long period of time,
’‘intermediate to tropical and subtropical.
North American species of Siphona Meigen
333
moved southward as conditions changed. This movement could have been in response to shifts
in hosts’ ranges, but I doubt this. Even though Siphona hosts are inadequately known, the
general distribution pattern of Siphona species - habitation of islands and continents far and
wide - suggests to me that hosts are ubiquitous and not crucial to the understanding of Siphona
zoogeography.
It is unfortunately premature to attempt to cover circumstances by which the S. tropica
group became established in Middle America since hosts, habitats and other aspects of extant
species’ way of life are unknown. Habitat information is vital, for the history of Tertiary floras
is complex (Graham, 1972; Axelrod, 1975, 1979; Rosen, 1978). It is significant, however, that
the S. tropica group is not represented in eastern North America. Apparently the S. tropica
group did not become associated with the temperate rain forest widely distributed in the United
States and Middle America by the Middle Tertiary (Graham, 1972; Axelrod, 1975, 1979).
Little data are available for inference of the history of the S', maculata, S. cristata and S.
geniculata groups in the Old World. The former is recognized as the most primitive Siphona
species group, so it presumably arose in Africa prior to all others. Examined specimens of
Siphona taxa from Africa did not include S. maculata group members, so the group may have
been virtually replaced in that region by more derived taxa. In America north of Mexico the S.
maculata, S. cristata and S. geniculata groups have the same pattern of distribution, so
probably have similar histories there. These histories are explored below.
Species distributions are primarily northern and western in America north of Mexico, and
from this several inferences can be drawn. It is likely the S. maculata, S. cristata and S.
geniculata groups have not had a long history in the New World. The distributional pattern
does not indicate an Eocene association with paratropical conditions as shown in the S. tropica
group. Neither are eastern relicts of the Middle Tertiary temperate rain forest evident. Why
the most primitive species group, the S. maculata group, shows a more recent distribution
pattern than the derived S. tropica group, is an anomaly without a satisfactory zoogeographic
explanation.
During the Miocene a cooling trend was evident, as the climate slowly began to approximate
our own. By late Middle Miocene higher latitudes were covered with a diverse microthermal
coniferous forest with a least some floral similarity between Eurasia and North America
(Wolfe, 1980). Similarly, a distinctive flora associated with Upper Tertiary orogenies was
developing in western North America, while drier conditions were experienced in central
regions and the temperate rain forest was retreating eastward and southward (Graham, 1972).
It was probably during this time that the first members of the S. maculata and S. cristata
groups entered North America, across the Bering land bridge12.
A Miocene or younger Siphona invasion of North America might account for certain
western elements in the distribution patterns of the S. maculata and S. cristata groups. It is
possible both S. pacifica and S. pisinnia (or their ancestors) represent Late Miocene - Early
Pliocene associations with young Rocky Mountain environments, particularly with their
inferred status as primitive members of the S. maculata and S. cristata groups, respectively.
The latter might be the older, for its distribution implies a warm temperate to subtropical
adaptation. Neither species seems to represent a center of origin for North American members
12North Atlantic land bridge(s) broken by that time (Matthews, 1979; Eldholm and Thiede,
1980).
Quaest. Ent., 1982, 18 (1-4)
334
O’Hara
of its group, so I infer that subsequent migrations of S. maculata and S. cristata group taxa
from the Palearctic Region took place.
A western distribution of possibly more recent age is that of S. oligomyia. As sister species
to -S. lutea, it almost certainly arose in America north of Mexico. S. lutea is cool temperate in
adaptation and transcontinental in distribution (Fig. 89), and it is apparently parapatric or
narrowly sympatric with the range of the more southerly distributed S. oligomyia (Fig. 90).
Southward shifts in range of S. lutea (or ancestor) during Pleistocene glaciations perhaps lead
to a vicariance of parental range in the west, with new species, S. oligomyia , developing a
unique set of adaptations and structural characteristics.
As mentioned in the “Phylogenetic interpretation” section, the relationship between S.
maculata and S. floridensis is cladistically unresolved (Fig. 103), though these species might
represent sister species, as inferred from their geographic distributions.
S. maculata is widespread across northern North America (Fig. 82), while S. floridensis is
only recorded from Florida and adjacent areas. Assuming S. maculata reached North America
prior to the Holocene, it would have experienced a southward displacement during Pleistocene
glaciations.
Under full glacial conditions of the Wisconsin (18,000ybp) (or an earlier glaciation), S'.
maculata might have reached northern Florida, as did some other boreal and cool temperate
elements of the biota (reviewed in Wright, 1981). Florida was apparently too dry to support
mesic forest during the Wisconsin full-glacial, but xeric oak-pine forest was present (Watts and
Stuiver, 1980). Whatever the exact nature of the biotas, community structures must have been
unique.
Perhaps as the Laurentide ice sheet retreated and the climate warmed, most populations of
S. maculata migrated northward, while others became stranded in pockets of suitable habitat
in the south. Though most of the latter became extinct, S. floridensis might be a living
descendant of one population that survived and adapted as conditions changed. (Similarly,
apparently disjunct populations of S. maculata in Nevada and southern Colorado (Fig. 82)
might also be relicts of the once extensive southern range of this species, though they have not
undergone the phenetic change of S. floridensis.)
Considerable similarity between North American and Palearctic members of the S.
maculata and S. cristata groups indicate faunal exchanges have continued into the Quaternary
period. At least three species in common between regions almost certainly reflects Pleistocene
exchange during interglacials.
The S', geniculata group as well could have entered North America as early as the Miocene,
but there is no indication of this. S. medialis is more southerly distributed than S. hokkaidensis
and more atypical of the group, so its arrival in North America presumably predates that of S.
hokkaidensis. I interpret the extraordinarily varied appearance of S. hokkaidensis as primarily
a Quaternary phenomenon, whereby populations were isolated and diverged. Further
consideration of this taxon must await study of Palearctic members.
The relationship between S. futilis and S. illinoiensis of the S. futilis group is very close,
but other S. futilis group affinities are nebulous. Perhaps the group is very old; in the New
World as old or older than the S. tropica group, since cool temperate taxa are evidently lacking.
As sister species, S. futilis and S. illinoiensis conform to a distributional pattern of noted
zoogeographic importance (Martin and Harrell, 1957; Axelrod, 1975; Rosen, 1978; Allen and
Ball, 1980). S. futilis and S. illinoiensis are, respectively, found in Mexico (Fig. 96) and
eastern United States (Fig. 92). It is generally agreed, on the basis of abundant data, that very
North American species of Siphona Meigen
335
little biotal exchange has occurred between Mexico and eastern United States since the
Miocene, when the Neotropical corridor between these areas deteriorated (Martin and Harrell,
1957; Rosen, 1978; Allen and Ball, 1980). The corridor was apparently not re-established even
during Pleistocene full-glacials; at least not with mesic forest elements (reviewed in Wright,
1981).
Despite the apparent rarity of post-Miocene biotal exchange between Mexico and eastern
United States, I am reluctant to propose a Miocene vicariance for S. futilis and S. illinoiensis.
By analogy with diversification of the S. tropica group in Mexico and inferred recent
vicariances of S. lutea and S. oligomyia, and S. maculata and S. floridensis, a time span of ca.
15 million years or more for the vicariance (without major change) of S. futilis and S.
illinoiensis seems too long. A more recent (Pleistocene) vicariance or dispersal is suggested by
phenetic evidence. If this explanation is valid, this example is one of the few that demonstrates
a post-Miocene connection between Mexico and eastern United States for mesic adapted taxa.
Siphona affinities between Middle and South America have not been closely examined.
Most species appear derived from northern elements, with a few evidently autochthonous
species. I expect exchange prior to the Pliocene connection of North and South America
occurred, though infrequently, leading to a small endemic Siphona fauna in South America.
To reiterate, the following are the main conclusions drawn from a historical zoogeographic
analysis of species groups and North American species of Siphona :
1 ) Siphona originated in Africa since the Late Cretaceous,
2) a S. tropica group taxon entered North America through a northern paratropical corridor
connecting the Old and New Worlds before the climate cooled in the Late Eocene,
3) minimum age of Siphona is Late Eocene,
4) the S. maculata, S. cristata and S. geniculata groups entered North America via Beringia
since the Miocene and periodic exchange continued into the Pleistocene, and
5) three species pairs represent Pleistocene vicariant events: S. maculata and S. floridensis, S.
lutea and S. oligomyia , and S. futilis and S. illinoiensis.
The major historical zoogeographic events concerning Siphona species within North
America are summarized in Fig. 108.
One of the elements of a zoogeographic hypothesis founded in part on cladistic relationships
is its predictive power (Ball, 1976). Explicit predictions, like assumptions, contribute to the
clarity and significance of hypotheses.
The assumptions listed earlier are testable and therefore render their own predictions. To
those can be added another. I do not expect taxa belonging to the S. maculata, S. cristata and
S. geniculata groups to be found in South America, for I interpret the ancestors of North
American members of these groups northern in origin and temperate to cold temperate in
adaptation. Thus, representation of one or more of these groups in South America would imply
either an earlier incursion into North America than I hypothesized (ie. during a warm period)
or a shift in adaptation of a member taxon from temperate to subtropical or tropical.
CONCLUDING REMARKS
It should be apparent from the foregoing text that much remains to be learned about the
genus Siphona. The basic classificatory work is inadequate for such regions as South America,
Australia and the Orient, and until the required studies have been completed, it will continue to
hinder attempts such as mine to delve into the historical (phylogenetic and zoogeographic)
Quaest. Ent., 1982, 18 (1-4)
336
O’Hara
aspects of the genus.
With few unique and reliable character states of adults by which to distinguish Siphona
species, identifications are commonly difficult. Since most keys are based exclusively on
morphological differences among species, only discovery of new morphological characters are
likely to improve them. Nevertheless there are other avenues of investigation into which I have
not ventured that might prove useful for species delimitation and phylogenetic analysis. These
include larval characters, host records, electrophoresis and cytological characters.
I have written little about the ecological requirements of Siphona species, for indeed there is
little known. Hosts are unrecorded for most North American species and additional
information (such as habitat preference) is similarly lacking. I can but hope others will examine
the aspects of Siphona species that I have not.
ACKNOWLEDGEMENTS
This paper was originally written in partial fulfillment of a Master’s degree. It is a pleasure
to thank my supervisor, Dr. G.E. Ball, for his thoughtful advice and suggestions throughout the
course of the study. In addition, his careful review of the first draft of the thesis led to
significant improvements in both substance and form. Financial aid from Natural Sciences and
Engineering Research Council grant A- 1399, held by Dr. G.E. Ball, is gratefully
acknowledged.
For first kindling my interest in tachinids, and suggesting Siphona as a thesis topic, I thank
Dr. D.M. Wood of the Biosystematics Research Institute, Ottawa. His knowledge of, and
enthusiasm for, the Tachinidae made discussions with him both informative and pleasurable.
Dr. Wood reviewed the manuscript and provided useful suggestions for its improvement.
Four other tachinid workers also contributed in various ways to this study. Dr. C.W.
Sabrosky, recently retired from the Systematic Entomology Laboratory, USDA, Washington,
was always prompt and helpful in my many dealings with him, and provided useful notes on
Siphona types under his care. Dr. B. Herting of the Staatl. Museum fur Naturkunde,
Stuttgart, provided specimens of Palearctic species of Siphona , and information on the
Palearctic Siphona fauna. Dr. H. Shima of Kyushu University, Ropponmatsu, sent Japanese
specimens of Siphona and commented on affinities among certain species. Special thanks are
extended to Dr. S. Andersen of the Zoologisk Museum, Copenhagen, who provided manuscript
copies of his “Revision of European species of Siphona Meigen” and “Phylogeny and
classification of Old World genera of Siphonini”. These arrived while the thesis was being
prepared for publication, so reference to them has been less than would otherwise have been the
case.
My wife, Wendy, assisted in many ways. As a companion on field trips, she helped collect
tachinid specimens. She also prepared locality data lists and distribution maps, and typed
portions of the manuscript into the computer.
I thank fellow students at the University of Alberta for enlightening discussions of
systematic theory and zoogeography, in particular J.S. Ashe, J.M. Cumming, G.A.P. Gibson
and R.E. Roughley. I also thank G.A.P. Gibson for testing, and recommending changes to, my
keys to adults of North American Siphona species.
I also thank an anonymous reviewer and members of my Advisory Committee (Drs. D.A.
Craig and J.S. Nelson) for critically reviewing the penultimate draft of the thesis, and
suggesting improvements. J.-F. Landry kindly prepared the French abstract.
North American species of Siphona Meigen
337
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North American species of Siphona Meigen
345
Fig. 1 . Terms and measurement of Siphona head features. Head profile of male S. illinoiensis.
Abbreviations: ar, arista; eye, compound eye; eye hgt, eye height; fr pit, frontal plate; fr s, frontal setae; gn,
gena; gn dil, genal dilation; hd hgt, head height; i vt s, inner vertical seta; lab, labella; lgt flgm 1, length of
first flagellomere; orb pit, orbital plate; o vt s, outer vertical seta; pafc, parafacial; pc orb s, proclinate orbital
setae; pip, palpus; pocr, postcranium; prem, prementum; prob, proboscis; re orb s, reclinate orbital setae; rst,
rostrum; subvb s, subvibrissal seta; vb, vibrissa.
Quaest. Ent., 1982, 18 (1-4)
346
O’Hara
Fig. 2. Structure and terms of Siphona thoracic nota, as illustrated by S. macronyx.
Abbreviations: Notal divisions: psct, prescutum; set, scutum; sctl, scutellum. Setae: acr, acrostichal; apl,
apical; bs, basal; dc, dorsocentral; ds, discal; hm, humeral; ial, intra-alar; ipal, intrapostalar; It, lateral; npl,
notopleural; pal, postalar; phm, posthumeral; psl, presutural; sbap, subapical; spal, supra-alar.
North American species of Siphona Meigen
347
3
Fig. 3. Right wing of S. maculata, illustrating average bristling on R4+5 and Siphona venation. Figs. 4-6.
Male foreleg acropods, dorsal aspect. Setae shown on one side only. Claw hooks directed ventrally, hence
not visible in figures. Figs. 4 and 6 illustrate average-sized claws and pulvilli, Fig. 5 large claws and
pulvilli. Fig. 4. S. illinoiensis. Fig. 5. S. intrudens. Fig. 6. S. lutea. Figs. 7-8. Preabdomens, dorsal aspect.
[Median marginal setae foreshortened in figures; subequal in length to laterals.] Fig. 7. S. intrudens ,
illustrating weak marginal setae and absence of median marginal setae on T1+2. Fig. 8. S. medialis,
illustrating strong lateral marginal setae and presence of median marginal setae on T1+2.
Abbreviations: Veins: A,, A2, branches of anal; CuAp cubitus; M, media; R,, R,+3, R4+5, branches of
radius; Sc, subcosta. Crossveins: dm-cu, discal medial-cubital; r-m, radial-medial. Abdomen: lat marg,
lateral marginal seta; med marg, median marginal seta; T]+2, T3, T4, T5, abdominal terga.
Quaest. Ent., 1982, 18 (1-4)
348
O’Hara
12
Fig. 9. Structure and terms of Siphona male genitalia, as illustrated by S. illinoiensis. Fig. 9a. Anterior
view of distiphallus. Fig. 9b. Lateral view of genitalia. Fig. 10. Sternum 5 of S. lutea, illustrating average
condition of postero-lateral margins. Fig. 1 1 . Sternum 5 of S. oligomyia , illustrating hooked condition of
postero-lateral margins. Fig. 12. Structure and terms of Siphona female genitalia, as illustrated by S.
hokkaidensis. Fig. 12a. Ventral view. Fig. 12b. Dorsal view. Fig. 12c. Lateral view. Abbreviations: Male
genitalia: aed apod, aedeagal apodeme; bsph, basiphallus; cere, cerci; dsph, distiphallus; epand, epandrium;
gon, gonopod; hypd apod, hypandrial apodeme; sur, surstylus. Female genitalia: cere, cerci; sg 8, segment
8; st 5, 6, 7 and 10, abdominal sterna. Scale bars = 0.1mm.
North American species of Siphona Meigen
349
Quaest. Ent., 1982, 18 (1-4)
Fig. 13. Flubbs-Hubbs diagram illustrating variation among specimens of Siphona species; eye height: head height. The range for each species is indicated by a horizontal line and
the mean by a short vertical line. A hollow rectangle represents two standard errors to either side of the mean and a solid rectangle represents 1.5 standard deviations to either side
of the mean. Only range and mean values are shown for samples with less than ten specimens. See “Methods" section for further information.
.44
350
O’Hara
Fig. 14. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; length of first flagellomere: head height. See caption for Fig. 13 and “Methods” section
for further information.
2.0 2.5
North American species of Siphona Meigen
351
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Fig. 15. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; proboscis length: head height. See caption for Fig. 13 and “Methods" section for further
information.
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352
O’Hara
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Fig. 16. Hubbs-Hubbs diagram illustrating variation among specimens of Siphona species; length of first flagellomere: eye height. See caption for Fig. 13 and “Methods” section
for further information.
North American species of Siphona Meigen
353
Quaest. Ent., 1982, 18 (1-4)
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Figs. 18-21. Male head profiles of Siphona species. Fig. 18. S. maculata\
Fig. 19. S. floridensis\ Fig. 20. S. intrudens\ Fig. 21. S. lurida ; Scale bars = 0.5mm.
North American species of Siphona Meigen
355
Figs. 22-25. Male head profiles of Siphona species. Fig. 22. S. pacifiea\ Fig. 23. S. pisinnia\ Fig. .24. 5.
cristata: ; Fig. 25. S. multifaria\ Scale bars = 0.5mm.
Quaest. Ent., 1982, 18 (1-4)
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Figs. 26-29. Male head profiles of Siphona species. Fig. 26. S. lutea ; Fig. 27. S. oligomyia\ Fig. 28. S.
geniculata ; Fig. 29. S. medialis\ Scale bars = 0.5mm.
North American species of Siphona Meigen
357
Figs. 30-33. Male head profiles of Siphona species. Fig. 30. S. hokkaidensis , antennal type 1; Fig. 31. S'.
hokkaidensis, antennal type 2; Fig. 32. S', hokkaidensis , antennal type 3; Fig. 33. S. macronyx\ Scale bars
= 0.5mm.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Figs. 34-37. Male head profiles of Siphona species. Fig. 34. S. futilis ; Fig. 35. S. brunnea\ Fig. 36. S.
akidnomyia; Fig. 37. S. longissima. Scale bars = 0.5mm.
North American species of Siphona Meigen
359
Figs. 38-40. Male head profiles of Siphona species. Fig. 38. S. tropica ; Fig. 39. S. rizaba\ Fig. 40. S.
unispina., Fig. 41. Head profile of female S. lurida\ Scale bars = 0.5mm.
Quaest. Ent., 1982, 18 (1-4)
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360
O’Hara
Figs. 42-50. Female head profiles of Siphona species. Fig. 42. S. maculata; Fig. 43. S. floridensis\ Fig. 44.
S. intrudens\ Fig. 45. S. pacifica\ Fig. 46. S. cristata ; Fig. 47. S. multifaria\ Fig. 48. S. lutea\ Fig. 49. S'.
pisinnia ; Fig. 50. S. geniculata\ Scale bars = 0.5mm.
North American species of Siphona Meigen
361
Figs. 51-59. Female head profiles of Siphona species. Fig. 51. S. hokkaidensis\ Fig. 52. S. medialis\ Fig.
53. S. futilis\ Fig. 54. S. illinoiensis\ Fig. 55. S. brunnea\ Fig. 56. S. macronyx; Fig. 57. S. tropica ; Fig.
58. S. rizaba\ Fig. 59. S. akidnomyia\ Scale bars = 0.5mm.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Figs. 60-65. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia.
Fig. 60. S. maculata\ Fig. 61. S. floridensis\ Fig. 62. 5". intrudens\ Fig. 63. S. lurida\ Fig. 64. S. pacifica\
Fig. 65. S. pisinnia ; Scale bars = 0.1mm.
North American species of Siphona Meigen
363
Figs. 66-71. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia;
c. Lateral view of second distiphallus. Fig. 66. S. cristata\ Fig. 67. S. multifaria\ Fig. 68. 5. lutea\ Fig. 69.
S. oligomyia ; Fig. 70. S. geniculata\ Fig. 71. S. hokkaidensis\ Scale bars = 0.1mm.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Figs. 72-77 . Male genitalia of Siphona species: a. Anterior view of distiphalius; b. Lateral view of genitalia.
Fig. 72. S. medialis ; Fig. 73. S. futilis'. Fig. 74. S. brunnea\ Fig. 75. S. macronyx\ Fig. 76. S. akidnomyia\
Fig. 77. S. longissima-. Scale bars = 0.1mm.
North American species of Siphona Meigen
365
Figs. 78-80. Male genitalia of Siphona species: a. Anterior view of distiphallus; b. Lateral view of genitalia.
Fig. 78. S', tropica ; Fig. 79. S. rizaba ; Fig. 80. S. unispina\ Scale bars = 0.1mm.
Quaest. Ent., 1982, 18 (1-4)
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Fig. 81. Known North American distribution of Holarctic species Siphona hokkaidensis , with data on
antennal type and abdominal setae. Dots denote locality records for S. hokkaidensis “b” (Figs. 13-16), and
female specimens for which there were no male associates. Squares denote locality records for male
specimens with antennal type 1, and associated females, from Alberta, Saskatchewan and Manitoba ( S .
hokkaidensis" a”). Rings signify locality records for which data were lacking. Abbreviations: Antennal type
1, Fig. 30; type 2, Fig. 31; type 3, Fig. 32; f, female, Fig. 51. N, number of specimens in sample. No. of
med. marg. on T,+2, number of specimens with 0, 1 or 2 median marginal setae on T1+2: 0, setae absent;
1, one seta on one side only; 2, 1 pair of setae. >1 pr. lat. marg. T3, number of specimens with more than
1 lateral marginal seta on at least one side on T3.
North American species of Siphona Meigen
367
Fig. 82. Known North American distribution of the Holarctic species Siphona maculata Staeger. Fig. 83.
Known distribution of Siphona pisinnia n. sp. Fig. 84. Known distribution of Siphona floridensis n. sp.
Quaest. Ent., 1982, 18 (1-4)
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Fig. 85. Known distribution of Siphona intrudens (Curran). Fig. 86. Known North American distribution
of the Holarctic species Siphona lurida Reinhard. Fig. 87. Known distribution of Siphona pacifica n. sp.
Fig. 88. Known distribution of Siphona macronyx n. sp.
North American species of Siphona Meigen
369
Fig. 89. Known distribution of Siphona lutea (Townsend). Fig. 90. Known distribution of Siphona
oligomyia n. sp. Fig. 91. Known North American distribution of S. geniculata (De Geer) (introduced from
the Palearctic Region). Fig. 92. Known distribution of Siphona illinoiensis Townsend.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Fig. 93. Known distribution of Siphona multifaria n. sp.
North American species of Siphona Meigen
371
Fig. 94. Known distribution of Siphona medialis n. sp. Fig. 95. Known North American distribution. of the
Holarctic species Siphona cristata (Fabricius).
Quaest. Era., 1982, 18 (1-4)
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O’Hara
Figs. 96-101. Known distributions of Siphona species. Fig. 96. Siphona futilis van der Wulp. Fig. 97.
Siphona brunnea n. sp. Fig. 98. Siphona rizaba n. sp. Fig. 99. Siphona akidnomyia n. sp. Fig. 100.
Siphona longissima n. sp. Fig. 101. Siphona tropica (Townsend).
North American species of Siphona Meigen
373
Fig. 102. Character matrix presenting the distribution of character states among North American Siphona
species and African species S. unispina (Mesnil). Characters and states are discussed in “Phylogenetic
interpretation” section.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
° "< < u.
Fig. 103. Cladogram illustrating inferred relationship between the S. maculata group and the other North
American Siphona species groups, with a hypothetical phylogeny of North American species of the S.
maculata species group. Dots denote apotypic character states, rings plesiotypic states. For explanation and
discussion of characters and states see Fig. 102 and “Phylogenetic interpretation” section.
North American species of Siphona Meigen
375
Figs. 104-107. Cladograms illustrating hypothetical phylogenetic relationships among North American
species of Siphona species groups. Dots denote apotypic character states, rings plesiotypic states. For
explanation and discussion of characters and states see Fig. 102 and “Phylogenetic interpretation” section.
Fig. 104. The S. cristata group; Fig. 105. The S. geniculata group; Fig. 106. The S. tropica group; Fig.
107. The S. futilis group.
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
Fig. 108. Generalized historical zoogeography of North American species of Siphona, as inferred from a
cladistic analysis and species distributions.
(1) 5. tropica group - migration into North America through Beringia or across a North Atlantic land
bridge prior to the Late Eocene during Tertiary climatic optimum, with subsequent diversification in
Mexico, (a) Late Tertiary to Pleistocene migration of taxa into Central and South America.
(2) S. maculata, S. cristata and S. geniculata groups - first representatives arrived in North America
during the Miocene; exchange continued into Pleistocene, (a) Oldest taxa of these groups might have
become associated with the developing Rocky Mountain biota (?Late Miocene - Pliocene), (b)
Transcontinental, cool to cold temperate taxa, indicative of later invasions (?Pliocene - Pleistocene), (c)
Vicariance of S. lutea and 5. oligomyia (Pleistocene), (d) Vicariance of S. maculata and S. floridensis
(Pleistocene).
(3) S. futilis group - place and age of origin uncertain. Vicariance of S. futilis and S. illinoiensis
(Pleistocene).
North American species of Siphona Meigen
377
INDEX TO NAMES OF TAXA
(Synonyms in italics)
FAMILIES AND TRIBES
Bombyliidae, 329
Noctuidae, 264
Oestridae, 329
Piophilidae, 340
Siphonini, 278, 309, 314
Tachinidae, 314, 326, 329, 330, 336
Tipulidae, 264
GENERA
Aphantorhapha Townsend, 274, 314
Asiphona Mesnil, 274, 314
Bucentes Latreille, 275, 278
Ceranthia Robineau-Desvoidy, 274, 314,
319
Ceranthia 5. lat., 274, 314, 315, 316, 326
Ceranthia 5. str., 274
Crocuta Meigen, 275
Dischistus Loew, 329
Peribaea Robineau-Desvoidy, 314
Phantasiosiphona Townsend, 274, 275
Protopiophila Duda, 329
Pseudosiphona Townsend., 314, 315
Siphonopsis Townsend, 273, 274, 314, 315
SPECIES AND SUBSPECIES
, cinereus Latreille, Bucentes, 275
abbreviata (Villeneuve), Siphona, 278
abbreviata Villeneuve, Bucentes, 278
abdominalis Robineau-Desvoidy,
Ceranthia, 317
akidnomyia new species, Siphona, 278,
285, 286, 306, 308, 311, 312, 313, 317,
325, 326, 328
albocincta (Villeneuve), Siphona, 278
albocincta Villeneuve, Bucentes, 278
alticola (Mesnil), Ceranthia, 275
alticola (Mesnil), Siphona, 275
alticola Mesnil, Crocuta, 214, 275
amoena (Mesnil), Siphona, 278
amoena Mesnil, Crocuta, 278
amplicornis Mesnil, Siphona, 278
analis Meigen, Siphona, 278, 299
analis Robineau-Desvoidy, Siphona, 280
angusta Mesnil, Siphona, 278
antennalis (Mesnil), Siphona, 278
antennalis Mesnil, Crocuta, 278
anthomyformis Lynch Arribalzaga,
Siphona, 280
atricapilla Mesnil, Siphona, 278
bevisi Curran, Siphona, 278, 318
bilineata (Mesnil), Siphona, 278
bilineata Mesnil, Crocuta, 278
boreata Mesnil, Siphona, 278, 325
brasiliensis Townsend, Siphonopsis, 273,
274
brunnea new species, Siphona, 278, 285,
286, 305,306,312,325,328
brunnescens (Villeneuve), Asiphona, 318
candelisequa Schiffermiiller, Agrotis, 273
capensis Curran, Siphona, 278
ceres (Curran), Siphona, 278, 305
ceres Curran, Bucentes, 278, 305
chaetolyga Rondani, Siphona, 278, 292
cinerea (Latreille), Siphona, 219
cinerea Latreille, Bucentes, 279, 299
cinerea Meigen, Siphona, 219
clausa Robineau-Desvoidy, Siphona, 280
collini Mesnil, Siphona, 278, 322
conata Reinhard, Siphonopsis, 273, 274
confusa Mesnil, Siphona, 273, 278, 323
consimilis Robineau-Desvoidy, Siphona,
280
cothurnata (Mesnil), Siphona, 278
cothurnata Mesnil, Crocuta, 278
crassulata (Mesnil), Ceranthia, 274, 275
crassulata (Mesnil), Siphona, 275
crassulata Mesnil, Crocuta, 274, 275
creberrima (Speiser), Siphona, 278
creberrima Speiser, Crocuta, 21 8
cristata (Fabricius), Siphona, 267, 272,
273, 278, 281, 284, 292, 293, 294, 295,
296, 297, 298, 299, 303, 307, 308, 313,
319, 322, 323, 324, 328, 330, 331, 333,
334, 335
cristata Fabricius, Stomoxys, 278, 292
Quaest. Ent., 1982, 18 (1-4)
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O’Hara
cuthbertsoni Curran, Siphona, 274, 278,
298, 319
delicatula Mesnil, Siphona , 279
diluta (Wulp), Clausicella, 275
diluta Wulp, Siphona , 273, 275
dorsalis Brauer and Bergenstamm,
Siphona , 280
efflatouni Mesnil, Siphona, 278, 322
flavifrons Staeger, Siphona, 273, 278, 322,
324
flavipes (Coquillett), Ceranthia, 317
floridensis new species, Siphona, 278, 282,
286, 287, 288, 289, 316, 321, 322, 323,
326.328.334.335
foliacea (Mesnil), Siphona, 278
foliacea Mesnil, Crocuta , 278
fuliginea cerina Mesnil, Siphona, 278
fuliginea Mesnil, Siphona, 278
fuliginea rubea Mesnil, Siphona, 278
fuscicornis Robineau-Desvoidy, Siphona,
280
futilis Wulp, Siphona, 273, 278, 285, 286,
305, 306, 307, 308, 310, 311, 312, 324,
325.328.334.335
gedeana Wulp, Siphona, 278
geniculata (De Geer), Siphona, 264, 273,
275, 276, 277, 278, 282, 299, 300, 301,
303, 304, 307, 322, 323, 324, 326, 327,
328.330.331.333.334.335
geniculata De Geer, Musca, 275, 299, 300
gigantea Schrank, Tipula, 273
gracilis (Mesnil), Siphona, 279, 325
gracilis Mesnil, Crocuta , 279
grandistyla Pandelle, Siphona, 279, 318,
324, 325
hokkaidensis Mesnil, Siphona, 267, 268,
273, 276, 279, 282, 283, 284, 300, 301,
302, 303, 304, 316, 323, 324, 328, 334
humeralis Robineau-Desvoidy, Siphona,
280
illinoiensis Townsend, Siphona, 277, 279,
283, 284, 304, 306, 307, 308, 324, 325,
328, 334, 335
infuscata (Mesnil), Siphona , 280
infuscata Mesnil, Crocuta , 280
intrudens (Curran), Siphona, 267, 277,
279, 282, 283, 287, 288, 289, 290, 291,
297, 303, 308, 321, 322, 326, 327, 328
intrudens Curran, Bucentes, 279, 288
irrorata Meigen, Tipula, 273
janssensi (Mesnil), Siphona, 279
janssensi Mesnil, Crocuta , 279
jocularis Mesnil, Actia, 274
kuscheli (Cortes), Siphona, 279
kuscheli Cortes, Phantasiosiphona , 274,
279
laticornis Curran, Siphona, 279
lindneri Mesnil, Siphona, 279
longissima new species, Siphona, 279, 285,
306, 309, 310, 311, 312, 313, 325, 328
lurida Reinhard, Siphona, 276, 279, 282,
286, 287, 289, 290, 291, 298, 316, 317,
321,322,326,327,328
lutea (Townsend), Siphona, 275, 276, 279,
282, 292, 293, 294, 296, 297, 298, 303,
308, 313, 317, 318, 322, 323, 325, 328,
334,335
lutea Townsend, Crocuta , 279, 296
macronyx new species, Siphona, 279, 283,
284, 311, 312, 313, 317, 325, 326, 328
maculata griseola Mesnil, Siphona, 279
maculata Staeger, Siphona, 267, 272, 273,
279, 282, 283, 286, 287, 288, 289, 291,
297, 298, 303, 316, 317, 319, 321, 322,
326, 327, 328, 329, 331, 333, 334, 335
maculipennis Meigen, Siphona, 279
malaisei (Mesnil), Actia, 275
malaisei (Mesnil), Siphona , 275
malaisei Mesnil, Crocuta, 274, 275
maxima Poda, Tipula, 273
medialis new species, Siphona, 276, 277 ,
279, 281, 303, 304, 308, 313, 323, 328
meigenii (Lepeletier and Serville),
Siphona , 279, 300
meigenii Lepeletier and Serville, Bucentes,
279
melania (Bezzi), Siphona, 279
melania Bezzi, Bucentes, 279
melanocera Robineau-Desvoidy, Siphona,
280
melanura Mesnil, Siphona, 279
minuta (Fabricius), Stomoxys, 279, 300
Quaest. Ent., 1982, 18 (1-4)
North American species of Siphona Meigen
379
multifaria new species, Siphona, 267, 272,
273, 279, 281, 284, 285, 286, 288, 292,
293, 294, 295, 297, 298, 299, 303, 307,
308,319,322,323,328
munroi Curran, Siphona, 279
murina (Mesnil), Siphona, 279
murina Mesnil, Crocuta, 279
nigricans (Villeneuve), Siphona, 279, 324,
327
nigricans Villeneuve, Bucentes , 279
nigripalpis (de Meijere), Siphona , 278
nigrohalterata Mesnil, Siphona , 278
nigroseta Curran, Siphona, 279
nigrovittata Meigen, Siphona , 279, 300
nobilis (Mesnil), Siphona, 279
nobilis Mesnil, Crocuta , 279
obelisea Schiffermiiller, Agrotis, 273
obesa (Mesnil), Siphona, 279
obesa Mesnil, Crocuta , 279
obscuripennis Curran, Siphona, 279
oculata Pandelle, Siphona , 279, 292
oleracea L., Tipula, 273
oligomyia new species, Siphona, 274, 277,
279, 282, 296, 297, 298, 313, 318, 319,
322, 323,325,326,328,334,335
pacifica new species, Siphona, 279, 283,
284, 286, 289, 291, 298, 313, 321, 323,
328,333
palpina Zetterstedt, Siphona , 278, 292
paludosa Meigen, Tipula, 264, 273
paludosa Mesnil, Siphona, 279, 324
patellipalpis (Mesnil), Siphona, 279
patellipalpis Mesnil, Crocuta , 279
pauciseta Mesnil, Siphona , 280
pauciseta Rondani, Siphona, 279, 292, 323
pellex (Mesnil), Actia, 275
pellex (Mesnil), Siphona , 275
pellex Mesnil, Crocuta , 274, 275
persilla Coquillett, Siphona , 280
phantasma (Mesnil), Siphona, 279
phantasma Mesnil, Crocuta , 279
pigra Mesnil, Siphona, 279
pisinnia new species, Siphona, 276, 280,
281, 283, 284, 285, 286, 292, 293, 294,
298, 299, 306, 322, 323, 327, 328, 333
plusiae (Coquillett), Siphonopsis, 273,
274, 305
pseudomaculata Blanchard, Siphona, 280
pusilla Robineau-Desvoidy, Siphona, 280
quadricincta Rondani, Siphona , 280
quadrinotata Robineau-Desvoidy,
Siphona, 280
reducta (Mesnil), Siphona, 280
reducta ludicra Mesnil, Siphona, 280
reducta Mesnil, Crocuta , 280
rizaba new species, Siphona, 280, 285,
286, 306, 309, 310, 311, 312, 325, 328
rossica Mesnil, Siphona, 280, 324
rubrapex Mesnil, Siphona, 280
rubrica (Mesnil), Siphona, 280
rubrica Mesnil, Crocuta , 280
setinerva (Mesnil), Siphona, 280, 318
setinerva Mesnil, Crocuta , 280
setosa Mesnil, Siphona, 280
seyrigi Mesnil, Siphona, 280, 321, 327
silvarum Herting, Siphona , 279, 301
silvatica Robineau-Desvoidy, Siphona, 280
simulans (Mesnil), Siphona, 280
simulans Mesnil, Crocuta , 280
singularis (Weidemann), Siphona, 280
singulars Wiedemann, Tachina, 280
siphonosoma Malloch, Actia, 274
sola Mesnil, Siphona, 280
spinulosa (Mesnil), Siphona, 280
spinulosa Mesnil, Crocuta , 280
tachinaria Meigen, Siphona , 279, 300
taiwanica (Baranov), Siphona, 280
taiwanica Baranov, Crocuta , 280
tenuis Curran, Siphona , 279, 296
testacea Robineau-Desvoidy, Siphona, 280
trichaeta (Mesnil), Siphona, 280
trichaeta Mesnil, Crocuta , 280
triseta Mesnil, Actia, 274
tristis Robineau-Desvoidy, Siphona, 280
tropica (Townsend), Siphona, 276, 277 ,
280, 285, 286, 308, 309, 310, 311,312,
—318, 325, 327, 328, 331, 332, 333, 334,
335
tropica Townsend, Phantasiosiphona , 274,
275, 280, 309
unispina (Mesnil), Siphona, 280, 318, 325,
331
380
O’Hara
unispina Mesnil, Crocuta , 280
urbanis (Harris), Siphona , 279
urbanis Harris, Musca, 279, 300
vittata Curran, Siphona, 280
vixen Curran, Siphona, 280
wittei (Mesnil), Siphona, 280
wittei Mesnil, Crocuta , 280
Quaest. Ent., 1982, 18 (1-4)
381
BOOK REVIEW
SPENCER, K.A. 1981. A revisionary study of the leaf-mining flies (Agromyzidae) of
California. Division of Agricultural Sciences, University of California, Special Publication
3273. iv + 489 pages containing 655 figures. $20 U.S.
This work is essentially a regional fauna, with description of new species and revisionary
comments included as necessary. It raises the number of agromyzid species recorded in
California to 252, of which 132 are described as new. Californian dipterists will find Spencer’s
treatment a sound foundation for further studies, and I hope they will be inspired to make it out
of date as soon as possible. At 252 species the total of known Californian species is surely still
underdocumented in comparison with the well over 300 species now known in the British Isles. I
would expect the real total of Californian species to be in excess of 500 in view of the great
diversity of climates and habitats represented in that state.
Two new taxa of the genus-group are proposed in this work: Galiomyza (type-species
Agromyza morio Brishke) for certain leaf-miners of Rubiaceae formerly included in a
heterogenous concept of Praspedomyza, and Annimyzella (type-species Agromyza maculosa
Malloch) as a subgenus of Amauromyza. These proposals seem to me justified.
In general the taxonomic treatment is up-to-date, but the synonymy of Metopomyza
griffithsi Sehgal with M. scutellata (Fallen) proposed on page 336 has been refuted in the
special study of Metopomyza contained in Tschirnhaus’ (1981) work “Die Halm- und
Minierfliegen im Grenzbereich Land-Meer der Nordsee” (Spixiana, suppl. 6). Spencer’s figure
of the aedeagus of a Californian specimen (Fig. 487) agrees with that of M. griffithsi described
from Alberta. The true M. scutellata has not been demonstrated to occur in North America.
The generic position of the new species described as Phytomyza minutissima Spencer needs
further study. This species does not run to Phytomyza in Spencer’s generic key because of the
lack of orbital setulae. Reduction of the aedeagal sclerotization makes the relationship of the
species difficult to interpret. On the basis of the information given, it could alternatively belong
to Paraphytomyza or Gymnophytomyza as well as to Phytomyza.
The 655 figures were mostly prepared by the author’s wife Ann and are of her usual good
quality. Some figures have been reproduced from previous publications, but most are original.
Scale lines have unfortunately been omitted.
While the content of this work meets with my approval and I have no hesitation in
recommending it to students of North American Diptera, there are some shortcomings in the
presentation. Offset printing has been used without reduction on pages of the standard size for
typing paper (8.5 x 1 1 inches). This results in an unnecessarily large volume which is too heavy
in relation to the strength of its (paper) binding. Libraries and regular users of the work will
need to have their copies rebound with hard covers. It would have been to the benefit of all
concerned if the page size had been reduced and stronger binding used. The figures were
intended for 1 /3 reduction, and considerable reduction in the space occupied by text could have
been achieved by use of single spacing. The proof reading has also been remiss, as evidenced by
the tantalizing references to “p. ” which appear frequently in the text. Page 7 has been
duplicated (appearing again as p. 12). More careful editing of future publications in this series
is called for.
Graham C.D. Griffiths
Department of Entomology
The University of Alberta
Edmonton, Alberta T6G 2E3
EDITOR’S ACKNOWLEDGEMENTS
In times of financial uncertainty, such as these, editors, and especially publishers, need to
scrutinize their operations- for obvious reasons. But it would seem that, at this time,
examination of editorial procedures and consciences is a more pressing need. Exponential
increase in scientific publication has reached crisis proportions, seemingly as a result of factors
having rather little to do with desire of authors to add to our store of knowledge (see Broad,
W.J., 1982, Bioscience 32 (8): 645-647). Associated with this increase has been an increase in
cost, an increase in presentation of false results, an increase in plagiarism, an increase in
repetition of content, an increase in multiple authorship with a corresponding decrease in
responsibility for material reported, a decrease of information per publication (the ultimate
being publication of “least publishable units”, or LPU’s), a possible decrease in quality of
content, and major problems for working scientists to cope with the volume of published pages.
Mr. Broad, a reporter for Science, concludes that scientific papers seem to have become more
tools for careerists to advance their own causes, rather than principal vehicles for transmission
of knowledge. He refers to the situation as “the disease of paper inflation”.
It seems to me that responsibility for this state of affairs is partly that of editors, and this is
why an examination of conscience is called for. Each editor can contribute to the needed cure
for paper inflation by first, identification of how his journal might have contributed to the
disease, and second, by a vigorous attempt to expunge those procedures deemed undesirable.
While not wishing to be self-righteous, I hasten to point out that Quaestiones Entomologicae
cannot be accused of contributing to paper inflation by encouraging publication of “LPU’s”,
and I am confident that our authors are not guilty of the other types of offenses listed. In
general we have sought extensive, original, treatments of varied entomological topics. I think
this volume is a fair sample of the sorts of papers published over the years, and I thank the
authors for choosing this journal.
As I write these lines, preparation of Volume 18 is nearing completion, and by resorting to
the expediency of publishing the entire year’s offering between two covers only, it seems likely
that we will achieve our goal of again issuing an entire volume in the same year that is
designated on the cover. Mrs. Suseela Subbarao, Publications Manager, is in large part
responsible for this, and I am grateful to her for her dedicated effort.
Although an editor is responsible for what appears in his journal, he seeks from others the
advice he must have about quality and accuracy of content of submitted manuscripts. He is not
compelled to accept all of the advice given, though it is probably in his best interests and that of
his journal to do so. Whether or not a reviewer’s advice is accepted, it is appropriate to
acknowledge assistance rendered. It is in that spirit that the following are acknowledged, and
the appearance of their names here does not imply endorsement of the contents of this volume.
Be that as it may, I am grateful for the assistance of:
H. E. Evans, Department of Zoology and Entomology, Colorado State University,
Fort Collins, Colorado;
J. R. Spence, Department of Entomology, University of Alberta;
D. D. Wilder, Insect Identification laboratory, Agriculture Research Service, U. S. D. A.,
United States National Museum, Washington, D. C.;
D. M. Wood, Biosystematics Research Institute, Research Branch, Agriculture
384
Canada, Ottawa, Ontario;
G. C. Williams, Center for Advanced Study in the Behavioral Sciences, Stanford,
California.
D. A. Craig served as editor during part of the summer, and undertook a substantial portion
of the chores associated with this volume. J. F. Landry prepared French translations of several
of the abstracts. J. H. Cumming and D. J. Williams read proof.
I am grateful to all of these individuals, and I look forward to receiving their cooperation in
preparation of future volumes.
George E. Ball
September, 1982.
INDEX TO VOLUME 18
385
Acilius sulcatus (L.), 3
adusta Navas, Povilla , 43, 45, 52
Afronurus Lestage, 33
albopilosus Cresson, Philanthus, 94, 95,
96, 98,99, 103, 104, 105, 106, 112,
113, 116, 118, 119, 120, 121, 123, 124
albrighti Traver, Ametropus, 44
album (Say), Ephoron, 43
Alcock, J., 93, 99, 105
Allen, R.K., 21,30
Alouf, N .J., 44
alpinus (Pictet), Baetis, 26
Ameletus Eaton, 43, 44
Ameletus inopinatus Eaton, 43, 53
americana Edmunds and Traver, Dolania,
44
Ametropodidae, 44
Ametropus albrighti Traver, 44
Ametropus ammophilus Allen and
Edmunds, 44
Ametropus eatoni Brodskij, 44
Ametropus fragilis Albarda, 44
Ametropus neavei McDunnough, 44
ammophilus Allen and Edmunds,
Ametropus , 44
Andersen, T., 46
Anderson, N.H.,
see Lehmkuhl, D.M., 34
Andrenidae, 93, 104
Aneletris Edmunds, 43
Anepeorus McDunnough, 33, 54
Aphilanthops frigidus (F. Smith), 103,
106
Aphilanthops Patton, 103
Armitage, K.B., 93, 103, 104, 105
Arthroplea Bengtsson, 33
Ashwell, I.Y., 95
Atalophlebiodes Phillips, 34
atratus McDunnough, Tricorythodes, 44
aurivillii Bengtsson, Ephemerella, 31
Axelrod, D.I., 122
Baetidae, 23, 27, 45, 47, 54
Baetiella, 27
Baetis alpinus (Pictet), 26
Baetis fuscatus (L.), 26
Baetis Leach, 20, 23, 27, 45, 54
Baetis macani bundyae, 26
Baetis macani Kimmins, 26
Baetis muticus (L.), 26
Baetis rhodani (Pictet), 23, 26
Baetis scambus Eaton, 26, 45
Baetis tricaudatus Dodds, 26
Baetis vagans McDunnough, 26
Baetis vernus (Curtis), 26
Baetisca bajkovi Neave, 44
Baetisca obesa (Say), 44
Baetisca rogersi Berner, 44
Baetisca Walsh, 44
Baetiscidae, 44
Baetodes Needham and Murphy, 27
bajkovi Neave, Baetisca , 44
Bartholomae, P.G., 43
basale (Walker), Siphloplecton , 34
Bates, H.W., 8
Behningiidae, 44
belgica Lestage, Torleya, 31
Bembix , 1 1 2
Bembix littoralis Turner, 1 12
Bembix texana Cresson, 1 12
Benech, V., 23
Benke, A.C., 47, 48
Berner, L., , 35
see also Edmonds, G.F., 21
see also Hall, R.J., 44
see also Scott, D.C., 43
see also Trost, L.M.B., 52
Bidwell, A., 43
bifidum (Bengtsson), Procloeon, 27
bilineata (Say), Hexagenia, 32, 49
Bohart, R.M., 98, 103
Bohle, H.W., 31,46
borealis Eaton, Metretopus, 35
Brachycercus Curtis, 29
Bretschko, G., 27, 43
Brittain, J.E., 27, 34
Brown, W.L.jr., 96, 119
Bryson, R.A., 95
Butler, A.G., 8
C.A., Menzie, 48
Caenidae, 27, 29, 47
Caenis horaria (L.), 29
Caenis macrura Stephens, 52
386
Index to Volume 18
Caenis rivulorum Eaton, 47
Caenis robust a Eaton, 52
Caenis Stephens, 27, 29, 54
Callibaetis Eaton, 27, 54
Callibaetis floridanus Banks, 52
Campsuarus Eaton, 54
canadense (Walker), Stenacron , 33
Cazier, M.A., 93
Centroptilum Eaton, 27
Centroptilum elsa Traver, 47
Centroptilum luteolum (Muller), 27
Centroptilum pennulatum (Eaton), 52
Cerceris Latareille, 103
Chadwick, H.W, 94
Chapman, R.N, 102
Choroterpes mexicanus Allen, 34
Christiansen, E.A.,
see Whitaker, S.H., 93
Cicindela , 123
Cicindelidae, 123
Cinygma Eaton, 33, 54
Cinygmula McDunnough, 33
Clayton, J.S.,
see Mitchell, J., 95
Clifford, H.F., 27, 34, 46, 47, 52, 54
Cloeon dipterum (L.), 27
Cloeon Leach, 27, 45
Cloeon simile Eaton, 27
Coleman, M.J., , 47
see also Hynes, H.B.N., 47
Colletidae, 104
Coloburiscus humeralis (Walker), 44
Cook, E.F.,
see Hall, R.J., 44
Corbet, P.S., 43
Corbet, S.A., 43, 52
Coupland, R.T., , 94
see Hulett, G.K., 94
crabroniformis Smith, Philanthus , 93, 99
cryptandrus A. Gray, Sporobolus , 94
cupida (Say), Leptophlebia, 34, 47, 53
D’Orville, H., 8
dalecarlica Bengtsson, Heptagenia, 33
Dalke, P.D.,
see Chadwick, H.W., 94
danica Muller, Ephemera , 31, 52
Dannella Edmunds, 54
Darwin, C., 2,5,6, 9, 10, 12
Darwin, E., 4
Darwin, F., 7, 9
Dawkins, R., 12
de Beer, G., 5
debilis (Walker), Paraleptophlebia, 34, 45
deficiens (Morgan), Serratella, 47
Demoulin, G., 52
diaphana Navas, Rithrogena , 32
dipterum (L.), Cloeon , 27
dispar (Curtis), Ecdyonurus, 33
Dix, R.L.,
see Hulett, G.K., 94
Dolania americana Edmunds and Traver,
44
dorothea Needham, Ephemerella, 47
Drunella Needham, 31
Dumas, J.,
see Lavandier, P., 21, 27
eatoni Brodskij, Ametropus, 44
Ecdyonurus dispar (Curtis), 33
Ecdyonurus Eaton, 32
Ecdyonurus lateralis (Curtis), 33
Ecdyonurus quadrilineatus Landa, 47
Ecdyonurus submontanus Klapalek, 33
Ecdyonurus torrent is Kimmins, 33
Ecdyonurus venosus (Fabricius), 33
Edmunds, C.H.,
see Edmunds, G.F. Jr., 52
Edmunds, F.H., 95
Edmunds, G.F., 21
Edmunds, G.F. Jr., 44, 52
Elassoneuria insulicola Demoulin, 35
Elliott, J.M., 30,31,46
elsa Traver, Centroptilum , 47
Epeorus Eaton, 33
Ephemera danica Muller, 31, 52
Ephemera Linneaeus, 31, 32
Ephemera simulans Walker, 31
Ephemerella , 21
Ephemerella aurivillii Bengtsson, 31
Ephemerella dorothea Needham, 47
Ephemerella ignita (Poda), 23, 30, 31, 45,
46
Ephemerella mucronata Bengtsson, 31, 47
Index to Volume 1 8
387
Ephemerella notata Eaton, 47
Ephemerella subvaria McDunnough, 47
Ephemerella Walsh, 30, 31, 45, 54
Ephemerellidae, 29, 30, 45, 47, 54
Ephemeridae, 31, 45, 54
Ephoron album (Say), 43
Ephoron virgo (Oliver), 43
Eucerceris Cresson, 103
Eumenidae, 104
Eurylophella funeralis (McDunnough),
31
Eurylophella Tiensuu, 31
Eurylophella versimilis (McDunnough),
47
Euthypolciidae, 44
Evans, H.E., 93, 96, 99, 102, 103, 104,
105, 106, 112
Fiance, S.B., 31
Fisher, R.A., 12
Fjellheim, A.,
see Andersen, T., 46
Flannagan, J.F., 32
Flint, R.F., 94, 95, 121
floridanus Banks, Callibaetis, 52
Flowers, R.W., 53
Formica Linnaeus, 103
fragilis Albarda, Ametropus, 44
Freitag, R., 93, 94, 122
Fremling, C.R., , 32
see also Thomforde, L.L., 32
Frenzel, B., 94
Frey, D.G.,
see Wright, H.E. jr., 94
frigidus (F. Smith), Aphilanthops, 103,
106
funeralis (McDunnough), Eurylophella ,
31
fuscatus (L.), Baetis , 26
fuscogrisea (Retzius), Heptagenia, 47
George, W.A., 8
germanica Eaton, Rhithrogena , 47
gibbosus (Fabricius), Philanthus , 93, 94,
99
gloriosus Cresson, Philanthus , 94
Gordon, E.L., 34
Grissell, E.E.,
see Bohart, R.M., 98, 103
Habroleptoides modesta (Hagen), 34
Habroleptoides Schoenemund, 54
Habrophlebia Eaton, 34
Habrophlebia lauta McLachlan, 34
Halictidae, 93, 104
Halictus rubicundus Christ., 103, 106
Hall, R.J., 44
Hamilton, A.L., 48
Hamilton, H.,
see also Clifford, H.F., 30, 34, 47
Hartland-Rowe, R., 43
Harvey, R.S., 44
Headley, F.W., 9
Heptagenia dalecarlica Bengtsson, 33
Heptagenia fuscogrisea (Retzius), 47
Heptagenia lateralis (Curtis), 33
Heptagenia sulphurea (Muller), 33, 53
Heptagenia Walsh, 33
Heptageniidae, 32, 33, 45, 47, 54
hercynia Landa, Rhithrogena , 47
Heterocloeon McDunnough, 54
Hexagenia bilineata (Say), 32, 49
Hexagenia limbata Serville, 32
Hexagenia Walsh, 31, 32, 45, 54
Hilsenhoff, W.L.,
see Flowers, R.W., 53
Hirsch, A.,
see Scott, D.C., 43
Homoeoneuria Eaton, 54
horaria (L.), Caenis, 29
Hudson, P.L., 32
Huff, B.L.,
see McCafferty, W.P., 33
Hulett, G.K, 94
humeralis (Walker), Coloburiscus, 44
Humpesch, U.H., 33, 46
Hunt, B.P., 32
Huxley, J.S., 1 1
hymenoides Ricker, Oryzopis , 94
Hynes, H.B.N., 47
Hynes, J.D., 21
lersel, J.J.A. van, 1 12
ignita (Poda), Ephemerella , 23, 30, 31,
45,46
388
Index to Volume 18
lilies, J., 26, 31
incertus (Traver), Tortopus, 43
inopinatus Eaton, Ameletus, 43, 53
insulicola Demoulin, Elassoneuria, 35
interpunctatum (Say), Stenacron, 33
Ironodes nitidus (Eaton), 33
Irvine, W., 4
Isonychia Eaton, 43, 54
J., Hunter, 3
Jensen, S.L.,
see also Edmonds, G.F., 21
Kelly, E.G.,
see Chapman, R.N., 102
Killins, B.A.,
see Clifford, H.F., 30, 34
Killnis, B.A.,
see Clifford, H.F., 47
King-Hele, D., 4
Kirby, W., 3, 4, 5
Klassen, R.W., 95
Kuhn, T., 5
lacustris (Eaton), Siphlornurus , 43
lanceolata Pursh, Psorealea , 94
Landa, V., 21, 31, 33,-43, 47
Lapchin, L.,
see Neveu, A., 23, 26, 31, 32
Larsen, R.,
see Andersen, T., 46
lateralis (Curtis), Ecdyonurus , 33
lateralis (Curtis), Heptagenia , 33
lauta McLachlan, Habrophlebia, 34
Lavandier, P., 21, 27
Lehmkuhl, D.M., 33, 34, 47, 53
Leptohyphes Eaton, 54
Leptophlebia cupida (Say), 34, 47, 53
Leptophlebia marginata (L.), 34, 47, 53
Leptophlebia vespertina (L.), 34, 49, 53
Leptophlebia Westwood, 34, 53
Leptophlebiidae, 33, 34, 45, 47, 54
Lesser, M., 5
limbata Serville, Hexagenia , 32
Lin, C.S.,
see Evans, H.E., 104
Lithobranchia McCafferty, 54
littoralis Turner, Bembix , 1 12
Love, K., 94, 95
Lorenz, K., 93
loyalae Navas, Rhithrogena, 32
luteolum (Muller), Centroptilum, 27
Macan, T.T., 17
Macan, T.T., 27, 52
macani bundyae, Baetis, 26
macani Kimmins, Baetis , 26
MacArthur, R.H., 118
Macdunnoa Lehmkuhl, 33
Mackey, A.P., 52
macrura Stephens, Caenis, 52
maculatus Kimmins, Tricorythus , 44
major (Klapalek), Torleya, 31
manuelito new subspecies, Philanthus,
121, 122, 123
Marchant, J., 7
marginata (L.), Leptophlebia , 34, 47, 53
Martin, P.S., 94, 122
Masaki, S., 46
Matthews, R.W.,
see Evans, H.E., 96, 112
Maudsley, R.,
see Macan, T.T., 27
Mayer-Oakes, W.J., 95
McCafferty, W.P., , 33, 43
see also Morihara, D.K., 20
McClure, R.G., 34
McLean, J.A., 44
Meier, P.G.,
see Bartholomae, P.G., 43
Mengel, R.M., 94
Menke, A.S.,
see Bohart, R.M., 103
Metretopodidae, 34, 54
Metretopus borealis Eaton, 35
Metretopus Eaton, 35, 54
mexicanus Allen, Choroterpes, 34
Mickel, C.E.,
see Chapman, R.N., 102
Microbembex, 104, 123
Microbembex monodonta (Say), 104
Miller, G.E.,
see Chapman, R.N., 102
Minshall, G.W.,
see Newell, R.L., 44
minutus Traver, Tricorythodes, 44
Index to Volume 18
389
Mitchell, J., 94, 95
Mivart, St. G., 9, 1 1
modest a (Hagen), Habroleptoides, 34
monodonta (Say), Microbembex, 104
Montgomery, T., 10, 1 1
Morgan, T.H., 1 1
Morihara, D.K., 20
morrisoni (Banks), Rhithrogena, 47
Mortenson, M.A.,
see Cazier, M.A., 93
Moss, H.C.,
see Mitchell, J., 95
mucronata Bengtsson, Ephemerella, 31,
47
muticus (L.), Baetis, 26
myops (Walsh), Potamanthus , 43
Nageli, C., 12
neavei McDunnough, Ametropus , 44
Neoephemera McDunnough, 54
Neoephemeridae, 44, 54
Neoephemeropsis Ulmer, 44
Neveu, A., 23,31,32
Newell, R.L., 44
nitens (Banks), Philanthus, 93, 113
nitidus (Eaton), Ironodes, 33
notata Eaton, Ephemerella , 47
obesa (Say), Baetisca, 44
occidentalis Eaton, Siphlornurus , 47
Oligoneuriella rhenana (Imhoff), 35, 46
Oligoneuriidae, 35, 45, 54
Oniscigaster wakefieldi McLachlan, 43
ornatum Tshernova, Procloeon, 27
oronti Alouf, Prosopistoma , 44
Oryzopis hymenoides Ricker, 94
Otto, C.,
see Andersen, T., 46
pacificus Cresson, Philanthus , 93
Paley, W„ 4
Palingenia Burmeister, 31, 44, 54
Palingeniidae, 44, 54
Paraleptophlebia debilis (Walker), 34, 45
Paraleptophlebia Lestage, 34, 54
Paraleptophlebia submarginata
(Stephens), 34
Parameletus Bengtsson, 43
Parker, J.R.,
see Chapman, R.N., 102
Parvella , 104
Peckham, E.G.,
see Peckham, G.W., 1 1, 93
Peckham, G.W., 11,93
pennulatum (Eaton), Centroptilum , 52
Pentagenia Walsh, 54
Pescador, M.L., 35, 44
Peters, W.L.,
see also Pescador, M.L., 35, 44
Petr, T., 43
Philanthus , 93, 94, 98, 103, 104, 124
Philanthus albopilosus Cresson, 94, 95,
96, 98,99, 103, 104, 105, 106, 112,
113, 116, 118, 119, 120, 121, 123, 124
Philanthus crabroniformis Smith, 93, 99
Philanthus Fabricius, 103, 113
Philanthus gibbosus (Fabricius), 93, 94,
99
Philanthus gloriosus Cresson, 94
Philanthus manuelito new subspecies,
121, 122, 123
Philanthus nitens (Banks), 93, 1 13
Philanthus pacificus Cresson, 93
Philanthus politus Say, 113
Philanthus psyche Dunning, 94, 123
Philanthus pulcher Dalla Torre, 93
Philanthus solivagus Say, 94
Philanthus triangulum Fabricius, 93
Philanthus ventilabris Fabricius, 94
picteti Sowa, Rithrogena, 32
Pleskot, G., 26
Poliomintha, 104
politus Say, Philanthus , 113
Polymitarcidae, 43, 54
Potamanthidae, 43
Potamanthodes Ulmer, 43
Potamanthus myops (Walsh), 43
Potamanthus Picket, 43
Poulton, E.B., 12
Povilla adusta rsfavas, 43, 45, 52
Pritchard, G., 54
Proboscidoplocia sikorai Vayssiere, 44
Procloeon bifidum (Bengtsson), 27
Procloeon ornatum Tshernova, 27
390
Index to Volume 18
Prosopistoma Latreille, 44, 54
Prosopistoma oronti Alouf, 44
Prosopistomatidae, 44, 54
Pseudiron .McDunnough, 33, 54
Pseudocloeon Klapalek, 27
Psorealea lanceolata Pursh, 94
psyche Dunning, Philanthus, 94, 123
pulcher Dalla Torre, Philanthus, 93
Puthz, V., 21
quadrilineatus Landa, Ecdyonurus , 47
Ramanankasina, R.E., 35, 44
Rau, N.,
see Rau, P., 93
Rau, P., 93
Reinhard, E.G., 93, 99, 104
rhenana (Imhoff), Oligoneuriella , 35, 46
Rhithrogena Eaton, 32, 33, 54
Rhithrogena germanica Eaton, 47
Rhithrogena hercynia Landa, 47
Rhithrogena loyalae Navas, 32
Rhithrogena morrisoni (Banks), 47
Rhithrogena semicolorata (Curtis), 32
rhodani (Pictet), Baetis , 23, 26
Richardson, M.Y., 33
Rithrogena diaphana Navas, 32
Rithrogena picteti Sowa, 32
rivulorum Eaton, Caenis, 47
Robertson, M.R.,
see Clifford, H.F., 52
robust a Eaton, Caenis, 52
rogersi Berner, Baetisca, 44
Romanes, J.G., 9, 10
Rosenberg, D., 17
Ross, H.E., 94, 122
rubicundus Christ., Halictus, 103, 106
Rutter, R.P., 32
scambus Eaton, Baetis, 26, 45
Scott, D.C., 43
Sellick, R.D.,
see Corbet, S.A., 43, 52
semicolorata (Curtis), Rhithrogena, 32
Serratella deficiens (Morgan), 47
Shapiro, A.M., 52
sikorai Vayssiere, Proboscidoplocia, 44
simile Eaton, Cloeon, 27
simulans Walker, Ephemera, 31
Siphlonisca Needham, 43
Siphlonuridae, 43, 47, 54
Siphloplecton basale (Walker), 34
Siphloplecton Clemens, 34
Siphloplecton speciosum Traver, 34, 35
Siphlornurus Eaton, 43
Siphlornurus lacustris (Eaton), 43
Siphlornurus occidentalis Eaton, 47
Smith, H.T.U., , 122
see also Thorp, J., 122
solivagus Say, Philanthus, 94
Sowa, R., 26, 32
speciosum Traver, Siphloplecton, 34, 35
Spence, W.,
see Kirby, W., 3, 4, 5
Sphecidae, 104
Spinadis Edmunds and Jensen, 54
Sporobolus cryptandrus A. Gray, 94
Steinbeck, J., 7
Stenacron canadense (Walker), 33
Stenacron interpunctatum (Say), 33
Stenacron Jensen, 54
Stenonema Traver, 33, 54
Stenonema tripunctatum (Banks), 33
Stewart, K.W.,
see McClure, R.G., 34
Stolzmann, J., 10, 1 1
Strnadtmann, R.W., 98
submar ginata (Stephens),
Paraleptophlebia, 34
submontanus Klapalek, Ecdyonurus, 33
subvaria McDunnough, Ephemerella, 47
sulcatus (L.), Acilius, 3
sulphurea (Muller), Heptagenia, 33, 53
Svensson, B., 31, 52
Swanson, G.A., , 32
see also Hudson, P.L., 32
Sweeney, B.W., , 47
see also Harvey, R.S., 44
Tamarix, 104
Tarter, D.C.,
see Richardson, M.Y., 33
Tauber, C.A.,
see Tauber, M .J., 19
Tauber, M .J., 19
texana Cresson, Bembix, 1 12
Index to Volume 18
391
Thalerosphyrus Eaton, 33
Thibault, M., 23,31,32, 47
Thomforde, L.L., 32
Thorp, J., 122
tiger beetles, 123
Tinbergen, N., 93, 104, 105
tinctus Kimmins, Tricorythus, 44
Tjonneland, A., 44
Torleya belgica Lestage, 31
Torleya major (Klapalek), 31
torrentis Kimmins, Ecdyonurus, 33
Tortopus incertus (Traver), 43
Tortopus Needham and Murphy, 54
Traver, J. R., 27
Traverella Edmunds, 54
triangulum Fabricius, Philanthus, 93
tricaudatus Dodds, Baetis, 26
Tricorythidae, 44, 45, 54
Tricorythodes atratus McDunnough, 44
Tricorythodes minutus Traver, 44
Tricorythodes Ulmer, 44
Tricorythus maculatus Kimmins, 44
Tricorythus tinctus Kimmins, 44
tripunctatum (Banks), Stenonema , 33
Trost, 52
Ulfstrand, S., 26, 33, 35, 53
vagans McDunnough, Baetis , 26
Van Dyck, W., 10, 11
Vannote, R.L.,
see also Sweeney, B.W., 47
see Harvey, R.S., 44
see Sweeney, B.W., 47
venosus (Fabricius), Ecdyonurus , 33
ventilabris Fabricius, Philanthus , 94
vernus (Curtis), Baetis, 26
versimilis (McDunnough), Eurylophella ,
47
vespertina (L.), Leptophlebia , 34, 49, 53
Vignes, J.C.,
see Neveu, A., 23,26,31,32
virgo (Oliver), Ephoron , 43
Vorzimmer, P., 7
Vuilleumier, B.S., 118
wakefieldi McLachlan, Oniscigaster, 43
Wallace, A.R., , 5, 8, 10, 11
see also Darwin, C., 5, 6
Wallace, Alex, 8
Waters, T.F., 19,47,48
Weir, J., 8
Weismann, A., 12
Whelan, K.F., 31,53
Whitaker, S.H., 93
Whitehead, D.R., 120
Williams, G.C., 12
Willis, H.J., 120, 123
Willoughby, N.G.,
see Corbet, S.A., 43, 52
Wilson, E.O.,
see Brown, W.L. jr., 95, 96, 1 19
see MacArthur, R.H., 1 18
Winterbourn, M.J., 34
Wise, E.J., 33
Wisely, B„ 44
Wissing, T.E.,
see Rutter, R.P., 32
Wright, H.E.jr., 94, 95
Zelt, K.A.,
see Clifford, H.F., 52
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
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