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Quaestiones

Entomologicae

A periodical record of entomological investigations^

published at the Department of Entomology,

University of Alberto, Edmonton, Conodo.

VOLUME 19

1983

CONTENTS

Darling-A Review of the New World Species of Euperilampus (Hymenoptera: Chalcidoidea),

with Notes about Host Associations and Phylogenetic Relationships 1

Lacey and Lacey-Filter Feeding of Simulium fulvinotum (Diptera: Simuliidae) in the Central

Amazon Basin .41

Fredeen-Trends in Numbers of Aquatic Invertebrates in a Large Canadian River during Four

Years of Black Fly Larviciding with Methoxychlor (Diptera: Simuliidae) 53

Ball and Hilchie-Cymindine Lebiini of Authors: Redefinition and Reclassification of Genera

(Coleoptera: Carabidae) 93

Book Review-Malicky, H. 1983. Atlas of European Trichoptera 217

Goulet-The Genera of Holarctic Elaphrini and Species of Elaphrus Fabricius (Coleoptera:

Carabidae); Classification, Phylogeny and Zoogeography . 219

Book Review-Chvala, M. 1983. The Empidoidea (Diptera) of Fennoscandia and Denmark. II.

General part. The families of Hybotidae, Atelestidae and Microphoridae. ... 483

Book Review-Duval, C. T. (Series Editor). 1982. Fauna of New Zealand. . . 486

Book Review-Griffiths, G. C. D. (Editor). 1982. Flies of the Nearctic Region 489

Book Review-Halffter, G. and W. D. Edmonds. 1982. The Nesting Behavior of Dung Beetles

(Scarabaeinae)- an Ecological and Evolutive Approach .491

Editor’s Acknowledgements 494

Index to Volume 19 . . . 495

Quaestiones Entomologicae, Volume 19(1,2)

Ball, G. E. and G. J. Hilchie (1983, 19: 93-216).- Cymindine Lebiini of Authors: Redefinition

and Reclassification of Genera (Coleoptera: Carabidae).

CORRIGENDA

pages

93 Title

“Coleoptera” was misspelled in the title. The corrected title is :CYMINDINE LEBIINI

OF AUTHORS: REDEFINITION AND RECLASSIFICATION OF GENERA

(COLEOPTERA: CARABIDAE).

ADDENDA

204 ACKNOWLEDGEMENTS

Financial support for this study was received through Grant A- 1399, Natural Sciences

and Engineering Research Council of Canada. Funds for publication were received

through Grant A- 1399 and through a grant from the Central Research Fund (NSERC),

University of Alberta. The authors very much appreciate this assistance.

1

j . .-i'vi- - ■ ■ -

1

I

i

Quaest

lones

Entomologicae

H S 0 iV/4^'

I AUG 81983

4/BRARIES

A periodical record of eittomoiogicol investigotions^

published at the Departmeat of Entomology,

University of Alberto, Edmonton, Conado.

VOLUME 19

NUMBERS 1-2

JANUARY-APRIL 1983

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department of

Entomology, University of Alberta, Edmonton, Alberta.

Volume 19 Numbers 1-2 1983

CONTENTS

Darling-A Review of the New World Species of Euperilampus (Hymenoptera: Chalcidoidea),

with Notes about Host Associations and Phylogenetic Relationships 1

Lacey and Lacey-Filter Feeding of Simulium fulvinotum (Diptera: Simuliidae) in the Central

Amazon Basin 41

Fredeen-Trends in Numbers of Aquatic Invertebrates in a Large Canadian River during Four

Years of Black Fly Larviciding with Methoxychlor (Diptera: Simuliidae) 53

Ball and Hilchie-Cymindine Lebiini of Authors: Redefinition and Reclassification of Genera

(Coleoptera: Carabidae) 93

Book Review-Malicky, H. 1983. Atlas of European Trichoptera 217

Euperilampus triangularis Say, female

A REVIEW OF THE NEW WORLD SPECIES OF EUPERILAMPUS (HYMENOPTERA;

CHALCIDOIDEA), WITH NOTES ABOUT HOST ASSOCIATIONS AND

PHYLOGENETIC RELATIONSHIPS.

D. Christopher Darling

Entomology

Cornell University

Ithaca, New York

14853

Quaestiones Entomologicae

19:1-40 1983

ABSTRACT

The genera Perilampus Latreille, Euperilampus Walker, Krombeinius Boucek^

Steffanolampus Peck, and Monacon Waterston form a monophyletic taxon ranked either as a

family (Perilampidae), or as a subfamily (Perilampinae) of the Pteromalidae. Based on

synapomorphic character states of size of postspiracular sclerite, pronotal size, and sculpture

of inner orbits and propodeum, Krombeinius Boucek and Euperilampus Walker are sister

groups, their common ancestor in turn being the derived sister group of the Perilampus

hyalinus species group. The subgenera Euperilampus sensu stricto (type species Perilampus

gloriosus Walker, 1862) and Euperilampoides Girault (type species Euperilampoides

scutellatus Girault, 1915) are invalid taxa in a phylogenetic system, because character states

hypothesized to be synapomorphic for them are more likely homoplasious. The 12 New World

species of Euperilampus are arranged in three groups: the E. tanyglossa group; E. krombeini

Burks; and the E. triangularis group. The E. tanyglossa group includes two Mexican species,

E. tanyglossa new species (type locality— Jalisco, Zapotlanejo), and E. aureicornis, new species

(type locality— Guerrero, Amula). The sister group of these species is hypothesized to be the

Old World species E. mediterraneus Bouctk, and to be related to E. scutellatus Girault, 1915,

another Palearctic species. The E. tanyglossa group + E. mediterraneus + E. scutellatus,

based on synapomorphic features of notauli and mesoscutal sculpture, comprise the sister

group of the stem of E. krombeini + the E. triangularis group. The latter is based on

synapomorphic features of postspiracular sclerite, propodeal and mesoscutal sculpture, and

color of body and wings, and includes nine New World species, of which four are in the E.

brasiliensis complex, and the others are not further classified. Members of the E. brasiliensis

complex are: E. brasiliensis (Ashmead); E. enigma, new species (type locality— Bolivia, Santa

Cruz, Robore): E. ameca, new species (type locality— Mexico, Nayarit, Santa Isabel); and E.

luteicrus (type locality— Mexico, Jalisco, Guadalajara). The unclassified species of the E.

triangularis group are E. triangularis (Say); E. gloriosus (Walker); E. magnus, new species

(type locality— Mexico, Chiapas, El Chorreadero); E. solox, new species (type locality—

Argentina, Tucuman, Tacanas); and E. iodes, new species (type locality— Brazil, Santa

Catarina, Nova Teutonia). Because of a shortage of reliably interpretable characters, a

reconstructed phylogeny of the species of the E. triangularis group is not proposed.

Euperilampus triangularis is a parasitoid of the ichneumonid, Hyposoter fugitivus Say, itself a

parasitoid of the arctiid, Hyphantria cunea (Drury).

2

Darling

RESUME

Les genres Perilampus Latreille, Euperilampus Walker, Krombeinius fioucek, Burksilampus Bouctk, Steffanolampus

Peck, et Monacon Waterston constituent une lignee monophyletique classifiee soit comme famille (Perilampidae), ou

comme une sous-famille (Perilampinae) des Pteromalidae. En considerant les conditions synapomorpiques de la taille du

sclerite postspiraculaire et du pronotum, et de la sculpture de la region intraorbitale et du propodeum, Krombeinius;

Boucek et Euperilampus Walker representent des taxons freres, dont I’ancetre commun est a son tour le taxon derive et

frere du groupe d'especes de Perilampus hyalinus. Les sous-genres Euperilampus sensu stricto (espece rype.Perilampus

gloriosus Walker, 1862) et Euperilampoides Girault (espece type: Euperilampoides scutellatus Girault, 1915) sont des

taxons invalides dans le cadre d’une classification phylogenetique, parce que les conditions des caracteres supposement

synapomorphiques regroupant les deux sont probablement issues de convergence. Les 12 especes i/’Euperilampus du

Nouveau Monde sont arrangees en trois groupes: le groupe ^3f’E. tanyglossa; E. krombeini Burks; et le groupe d’E.

triangularis. Le groupe J’E. tanyglossa comprend deux especes mexicaines: E. tanyglossa, nouvelle espece (localite du

type: Jalisco, Zapotlanejo), et E. aureicornis, nouvelle espece (localite du type: Guerrero, Amula). Le taxon frere de ces

especes est suppose etre E. mediterraneus Boucek, de I’Ancien Monde, et serait lui-meme apparente a E. scutellatus

Girault, 1915, une autre espece eurasienne. Base sur les particularites synapomorphiques des notauli et de la sculpture

du mesoscutum, le groupe ^/’E. tanyglossa forme, avec les especes E. mediterraneus et E. scutellatus, le taxon frere d’E.

krombeini et du groupe d’E. triangularis. Ce dernier est defini a partir de caracteristiques synapomorphiques du sclerite

postspiraculaire, de la sculpture du propodeum et du mesoscutum, et de la couleur du corps et des ailes; il comprend

neuf especes du Nouveau Monde, parmi lesquelles quatre font partie du complexe d’E. brasiliensis, tandis que les autres

ne sont pas classifiees. Les membres du complexe d’E. brasiliensis sont: E. brasiliensis (Ashmead); E. enigma, nouvelle

espece (localite du type: Bolivie, Santa Cruz, Robore); E. ameca, nouvelle espece (localite du type: Mexique, Nayarit,

Santa Isabel); et E. luteicrus, nouvelle espece (localite du type: Mexique, Jalisco, Guadalajara). Les especes non

classifiees du groupe d’E. triangularis sont: E. triangularis (Say); E. gloriosus (Walker); E. magnus, nouvelle espece

(localite du type: Mexique, Chiapas, El Chorreadero); E. solox, nouvelle espece (localite du type: Argentine, Tucuman,

Tacanas); et E. iodes, nouvelle espece (localite du type: Bresil, Santa Catarina, Nova Teutonia). L’auteur ne presente pas

de diagramme phylogenetique pour les especes du groupe d’E. triangularis h cause d’un manque de caracteres

interpretables de facon sure. Euperilampus triangularis est un parasitoide ^/’Hyposoter fugitivus Say, un Ichneumonidae

lui-meme parasitoide de TArctiidae Hyphantria cunea (Drury).

TABLE OF CONTENTS

Introduction 2

Synopsis of the New World Species of Euperilampus 3

Methods and Terms 3

Material 5

Key to New World Euperilampus 7

The New World Species of Euperilampus 8

Phylogenetic Relationships 34

Acknowledgements 38

Literature Cited 38

INTRODUCTION

The genera Perilampus Latreille, Euperilampus Walker, Krombeinius Boucek,

Burksilampus Boucek, Steffanolampus Peck and Monacon Waterston form a well defined

monophyletic taxon regarded as either the family Perilampidae (Graham 1969) or as a

subfamily in the Pteromalidae, the Perilampinae (sensu Boucek 1978). Boucek (1978) has

characterized this taxon and reviewed the arguments concerning the placement of this group in

the higher classification of the Chalcidoidea, No resolution will be possible until phylogenetic

studies are conducted in the Chalcidoidea. The Pteromalidae is most certainly a paraphyletic

group (possible polyphyletic), and relegating Perilampus and related genera to this unnatural

assemblage is unwarranted phylogenetically and practically. I therefore follow Graham (1969)

The New World species of Euperilampus

3

and recognize the family Perilampidae. I exclude Chrysolampinae (sensu Graham 1969) from

this family, again following Graham (1969). Larval structures of Chrysolampus thenae

(Walker) were discussed by Askew (1980), and offer no basis for inclusion of this genus in a

higher taxon with the genus Perilampus. Where known, all species of Perilampus have a

planidial first instar larva (Smith 1912, Clancy 1946, Principi 1947). This type of larva is not

found in Chrysolampus. The larval characters cited by Askew (1980) as indicating a close

relationship between Perilampus and Chrysolampus are widely distributed in many chalcidoid

taxa (see Parker 1924) and are here regarded as plesiomorphies.

Revisions are currently available for all genera of Perilampidae except the large

cosmopolitan genus Perilampus (Boucek 1980, for Monacon; Boucek 1978, all other genera).

In assembling material for a revisionary study of New World Perilampus I have received

undescribed species of Euperilampus from Mexico and South America that provide important

insights into the phylogeny and biogeography of Euperilampus.

In this paper I discuss generic characters of Euperilampus, examine the validity of the

current subgeneric classification, and review and present a key to the New World species. Eight

new species are described, and all New World species except E. gloriosus are redescribed. The

host associations of E. triangularis are also discussed. A cladogram of species groups of

Euperilampus is presented, and the phylogenetic relationships of Krombeinius, Euperilampus

and Perilampus are discussed.

SYNOPSIS OF THE NEW WORLD SPECIES OF EUPERILAMPUS

E. tanyglossa species group

E. tanyglossa n. sp.

E. aureicornis n. sp.

E. krombeini Burks

E. triangularis species group

E. triangularis (Say)

E. gloriosus (Walker)

E. magnus n. sp.

E. solox n. sp.

E. iodes n. sp.

E. brasiliensis complex

E. brasiliensis (Ashmead) n. comb.

E. enigma n. sp.

E. luteicrus n. sp.

E. ameca n. sp.

METHODS AND TERMS

Color. — All New World species of Euperilampus exhibit metallic or iridescent colors that

are difficult to describe. These structural colors are the result of interference patterns, due to

asynchrony between the component wavelengths of entering and returning light. The

predominant color depends primarily on thickness of alternating cuticular lamellae and

distance between successive lamellae and, to a lesser degree, on angle of incidence of incoming

light. A change in angle of incidence from 60 degrees to 90 degrees can result in a 40 nm color

Quaest. Ent., 1983, 19 ((1,2))

4

Darling

shift. This can result in changes in color from violet to blue-violet or from blue to blue-green

(Fox 1979). Metallic colors can therefore only be described in general terms. I use spectral

colors, i.e., violet is preferred to purple, purple being a pigmentary color resulting from a

mixture of reds and blues. In this paper all colors were described when viewed under diffuse

incandescent light.

Care must also be taken in describing metallic colors of specimens subjected to certain

taxonomic procedures. For instance, lacto-phenol (a clearing agent) and thymol (an

anti-fungicide used in relaxing jars) are swelling agents which cause a change to longer

reflected wavelengths, and color changes from blue to green, or to brassy-yellow. These changes

are thought to be reversible (Fox 1979), but specimens treated with lacto-phenol retain the

aberrant colors months after removal from the swelling agent.

Structure and Sculpture. — Morphological terms follow Graham (1969) and Richards

(1977). Hence, ‘postspiracular sclerite’ is used for ‘prepectus’ of authors in the Chalcidoidea.

Sculpture types follow Eady (1968), except ‘coriarious’ is used in preference to ‘coriaceous’,

following the recommendation of Harris (1979, p. 2). Sculpture is best viewed and is here

described under diffuse light. Scanning electron micrographs illustrate the major types of

sculpture.

The antennae of Euperilampus are sexually dimorphic. The funicle is stouter and the scape

is expanded distally in males. In some species of Euperilampus the anterior face of the male

scape is roughened (100-200X magnification), and scanning electron microscopy reveals

indentations with pores (Figs. 45-52). Similar sexual characters are found in Perilampus and

Steffanolampus. In males of Perilampus hyalinus (Say) (Fig. 64), the anterior surface of the

scape is covered with large punctures each of which has a single central pore. In Perilampus

these structures have been referred to as ‘sensorial punctures’ (Smulyan 1936). It seems more

likely that the pores are glandular openings and not sensory in function; there is no indication of

a cuticular peg or dome. Note that many of the punctures are filled with material (Fig. 64).

Possibly this substance is the residue of pheromones involved in sexual behavior. Histological

studies will be necessary to adequately characterize these structures. I use the term ‘punctures’

for these structures. In Euperilampus there are two to six pores in each puncture (Figs.

46,48,52) or punctures are absent (Figs. 42-44). Determination of taxonomic importance of

number of pores per puncture must await collection of more material. The distribution of

punctures, however, has proved of considerable value in delimiting species.

Measurements. — The terms length (L), width (W), and height (H), refer to the maximum

value obtained by rotating the specimen. This avoids parallax problems encountered when

measuring three-dimensional objects. It is, however, critical to have both endpoints in focus

when the measurements are taken. Measurements and their abbreviations used in the text are

as follows: EH, eye height, taken in frontal view; MS, length of malar space; A, length of

anellus in dorsal view; FI, length of first funicular segment in dorsal view; SL, scape length;

SW, scape width; HW, head width, in frontal view; HL, head length, in frontal view, from

vertex to lower margin of clypeus; CH, clypeus height; SH, height of supraclypeal area; SW,

width of scrobes; OOL, length of ocular-ocellar line; POL, postocellar line, distance between

posterior ocelli; PN, length of pronotum along midline; MSC, length of mesoscutum along

midline; and SC, length of scutellum along the midline.

The New World species of Euperilampus

5

MATERIAL

This study is based on a total of 603 adults of Euperilampus, as well as representative

material of related taxa. Specimens included are housed in the following collections, which are

indicated in the text by the associated acronyms.

AEI: American Entomological Institute, Ann Arbor, MI, U.S.A. 48105 (H.K. Townes)

BMNH: British Museum (Natural History), London, England SW7 5BD (J.S. Noyes)

CAS: California Academy of Sciences, San Francisco, CA, U.S.A. 94118 (P.H. Arnaud, Jr.)

CNC: Canadian National Collection, Ottawa, Canada K1 A 0C6 (C. Yoshimoto)

CU: Cornell University, Ithaca, NY, U.S.A. 14853 (L.L. Pechuman)

DCD: D. Christopher Darling, personal collection

FSCA: Florida State Collection of Arthropods, Gainesville, FL, U.S.A. 32611 (L. Stange)

lESM: Instituto Entomologico San Miguel 1663, San Miguel, Argentina (M.A. Fritz)

IML: Instituto Miguel Lillo, Universidad Nacional de Tucuman, Tucuman, Argentina (P.

Fidalgo)

KSU: Kansas State University, Manhattan, KS, U.S.A. 66506 (H.D. Blocker)

NHMLAC: Natural History Museum, Los Angeles County, Los Angeles, CA, U.S.A. 90007

(R.R. Snelling)

UA: University of Arkansas, Fayetteville, AK, U.S.A. 72701 (R.G. Chenowith)

UG: University of Guelph, Guelph, Canada NIG 2W1 (D. Pengelly)

UK: Snow Entomological Museum, University of Kansas, Lawrence, KS, U.S.A. 66045 (C.D.

Michener)

UNLP: Universidad Nacional de La Plata, 1900 La Plata, Argentina (L. de Santis)

USNM: United States National Museum, Washington, DC, U.S.A. 20560 (E.E. Grissell)

USU: Utah State University, Logan, UT, U.S.A. 84322 (W.J. Hanson)

Other repositories for specimens of Euperilampus are as follows: American Museum of

Natural History, New York, NY 10024 (M. Favreau); Academy of Natural Sciences of

Philadelphia, Philadelphia, PA 19103 (D. Otte); Carnegie Museum of Natural History,

Pittsburgh, PA 15213 (G. Ekis); Colorado State University, Fort Collins, CO 80521 (H. E.

Evans); Illinois Natural History Survey, Urbana, IL 61803 (W. La Berge); Museum of

Comparative Zoology, Harvard University, Cambridge MA 02138 (A. Newton, Jr.); Ohio

State University, Columbus, OH 43210 (C. Triplehorn); Pennsylvania State University,

University Park, PA 16802 (K. Kim); Southwestern Research Station, Portal, AZ 85632 (V.D.

Roth); S.U.N.Y., Syracuse, Syracuse, NY 13210 (M. O’Brien); Texas A & M University,

College Station, TX 77843 (S. Merritt); University of Alberta, Edmonton, Canada T6G 2E3

(G.E. Ball); University of Arizona, Tucson, AZ 85721 (F.G. Werner); University of Georgia,

Athens, GA 30602 (C. Smith); University of Michigan, Ann Arbor, MI 48109 (T. Moore); and

University of Minnesota, St. Paul, MN 55108 (P. Clausen).

I thank all who allowed me to study the material under their care. The curators of the Snow

Entomological Museum (UK) and Utah State University Collection (USU) were particularly

generous in allowing holotypes described from their material to be deposited in the Smithsonian

Collection (USNM). This was requested to allow amalgamation of type material to facilitate

further study of the group. The result is that six of the ten extant primary types are at the

USNM, with single holotypes in BMNH, CAS, CNC, and IML.

Quaest. Ent., 1983, 19 ((1,2))

6

Darling

GENUS EUPERILAMPUS

Euperilampus Walker, 1871: 67. Type Species: Perilampus gloriosus Walker, 1862: 375, by monotypy and original

designation.

Euperilampus {Euperilampus): Boucek 1972:90 [as subgenus].

Euperilampoides Girault, 1915: 308. Type Species: Euperilampoides scutellatus Girault, 1915, by monotypy and original

designation [synonymy by Riek, 1966: 1227].

Euperilampus {Euperilampoides)-, Boucek 1972:90 [as subgenus].

Nesoperilampus Rohwer, 1923: 349. Type species: Nesoperilampus typicus Rohwer 1923, by monotypy and original

designation [synonymy by Riek, 1966: 1227].

Diagnosis. — Euperilampus is reliably distinguished from other perilampid genera by

having the postspiracular selerite a narrow triangle (Figs. 35-37), mueh less than half as wide

as the adjacent pronotal collar, and by having the marginal vein distinctly shorter than the

postmarginal vein (Figs. 57-59). Perilampus (Figs, 60, 61), Steffanolampus, Burksilampus,

and Monacon have the postspiracular selerite at least as wide as the adjacent pronotal collar.

All genera except Euperilampus have the marginal vein longer than the postmarginal vein

(Fig. 62).

The genus Euperilampus has been characterized by Riek (1966), Burks (1969) and Boucek

(1978). Boucek (1978) has presented a key to world species. Two subgenera were recognized,

following Boucek (1972, 1978): Euperilampus sensu stricto, bright metallic species with

midlobe of mesoscutum and entire scutellum with coarse cross-arcuate rugae. New World; and

Euperilampoides Girault, dark metallic to black species with thoracic dorsum generally

punctate-reticulate. Old World.

Character states of two new species described in this paper {E. tanyglossa and E.

aureicornis) refute this classification. These New World species have the punctate-reticulate

sculpture and slightly indicated notauli of the Old World subgenus Euperilampoides but are

metallic blue-green in color rather than black. Hence, these species are contradictory at the

first couplet of Boucek’s (1978) key, where the subgenera of Euperilampus are separated. As

will be discussed, E. tanyglossa and E. aureicornis are more closely related to Euperilampus

(Euperilampoides) mediterraneus than to other New World species. Phylogenetic relationships

within the genus, discussed in detail at the end of the paper, are not consistent with recognition

of New World and Old World subgenera, Euperilampus and Euperilampoides

(SYNONYMY, REVISED STATUS).

Comparative morphological studies of the Perilampidae have revealed that male genitalia

and structure of the labrum (both sexes) characterize Euperilampus.

Figures 1 and 2 illustrate major features of the male genitalia. All New World species with

the exceptions of E. gloriosus, E. aureicornis, E. magnus (male unknown), and E. ameca (male

unknown) and the Old World species E. scutellatus were examined and allow the following

characterization: distinct parameres lacking, basiparamere (Bp) with a patch of strong setae

(Ls) distributed on transparent areas (Ld) laterad of ventral lobe (VI). In Perilampus

(Domenichini 1953), Steffanolampus and Krombeinius (Darling, unpublished) the parameres

are well developed, with the strong setae distributed on these lobes. The presence of distinct

parameres is here regarded as the plesiomorphic state for the Chalcidoidea, because this state is

widely distributed in many taxa (see Domenichini 1953).

Figure 3 illustrates form of the highly distinctive labrum of Euperilampus adults.

Euperilampus triangularis, E. krombeini, E. scutellatus (males and females), and E.

tanyglossa (male) were examined and allow the following characterization: 8-digitate with a

deep median incision, each digitus with a strong terminal seta, and with a pair of smaller.

The New World species of Euperilampus

1

sessile setae located below the level of the digiti. This arrangement differs from that of

Perilampus (Riek 1966, Domenichini 1969) and Steffanolampus salicetum (Darling,

unpublished): 10 or 12-digitate, with one digit arising more toward the base of the structure,

and the labrum not deeply excised medially, and Krombeinius eumenidarum (Darling,

unpublished): a single narrow central stalk, with the seven digiti arising apically.

New World species of Euperilampus are metallic in color and are distributed from eastern

Canada to southern Brazil.

KEY TO NEW WORLD EUPERILAMPUS

1 Entire mesoscutum and scutellum punctate-reticulate (similar in sculpture to

the pronotum), notauli indistinctly indicated (Fig, 33); labio-maxillary complex

elongate, protruded far beyond the closed mandibles (Fig. 4) 2

V Midlobe of mesoscutum and scutellum with transverse rugae or costae, notauli

distinct (Fig. 14); labio-maxillary complex not conspicuously protruded beyond

closed mandibles (Fig. 9) 3

2 (1) Postspiracular sclerite with three or four weak foveae (Fig. 35); frontal carina

weakly divergent and following inner eye margin, inner orbits not markedly

narrowed at level of antennal toruli (Fig. 5) [Female funicle dark brown, male

funicle yellow] E. tanyglossa n. sp., p. 8

1' (1) Postspiracular sclerite with a single weak fovea, below fovea with coriarious

sculpture; frontal carina oblique, convergent towards inner eye margin, inner

orbits markedly narrowed at level of antennal toruli (Fig. 6) [Female and male

funicle yellow] E. aureicornis n. sp., p. 10

3 (L) Apex of scutellum broadly rounded (Fig. 12); lateral wall of scrobes merged

smoothly with face (not angulate in lateral view), the inner orbits without well

developed longitudinal costae or rugae (Fig. 8) E. krombeini Burks, p. 1 1

3' (L) Apex of scutellum acuminate (Figs. 13-16); lateral wall of scrobes merged

abruptly with face at level of antennal toruli (angulate in lateral view), inner

orbits with well developed longitudinal costae or rugae (Fig. 9)

{E. triangularis species group) 4

4 (30 Scutellum abruptly produced into lanceolate spine, much longer than wide (see

Boucek 1978, Fig. 9); metasoma and apex of scutellum bright coppery to golden

in color E. gloriosus (Walker), p. 15

4' (30 Scutellum without abrupt lanceolate spine; entire scutellum and metasoma

metallic violet, blues and greens, in some specimens with black areas, not

coppery in color 5

5 (40 Mesoscutum with distinct contrasting glossy black areas on sidelobe, along

notauli, black areas smooth, not roughened (Fig. 14) 6

5' (40 Mesoscutum without distinct contrasting black areas on sidelobe along notauli;

areas along notauli roughened in many specimens 9

6 (5) Fore and mid tibiae yellow, concolorous with tarsi; male antennal scape with

distinct punctures on anterior surface, surface roughened (Fig. 45) [Mexican,

Female unknown] E. luteicrus n. sp., p. 21

6' (5) Fore and mid tibiae dark, brown or metallic; male antennal scape with

punctures fewer and distinctly separated (Fig. 49,50) 7

Quaest. Ent., 1983, 19 ((1,2))

8

Darling

7 (6') Females [antennal scape not widened apically, Figs. 5-7]

E. brasiliensis complex

7" (6') Males [antennal scape widened apically, Figs. 42,45] 8

8 (7') Scape without distinct punctures on anterior surface (Fig. 49), surface smooth;

digiti of genitalia without large recurved teeth (Fig. 75)

E. brasiliensis (Ashmead), p. 19

8' (7') Scape with distinct punctures on anterior surface (Fig. 50), surface roughened;

digiti of genitalia with large, recurved teeth (Fig. 74)

E. enigma n. sp., p. 20

9 (50 Axillula with distinct costae, ventral costa merged with posterior margin of

axillula (Fig. 53); sculpture on scutellum reduced medially, rugae incomplete

(Fig. 16) E. triangularis (Say), p. 13

9' (50 Axillula smooth or with indistinctly defined costae, not merged with posterior

margin of axillula (Fig. 54); sculpture on scutellum not reduced medially, rugae

or costae complete (Fig. 13, 15) 10

10 (90 Midlobe of mesoscutum with regular cross-arcuate costae (Figs. 13,14) 11

10" (9") Midlobe of mesoscutum with irregular rugae (Fig. 15) 12

11 (10) Margin of scrobes, sinuous in frontal view, markedly flared at level of antennal

toruli (as in Fig. 11); postspiracular sclerite with large centrally located fovea

co-extensive with most of upper postspiracular sclerite; sidelobe of mesoscutum

with narrow black areas along notauli E. ameca n. sp., p.21

11" (10) Margin of scrobes, smoothly curved in frontal view, not markedly flared at level

of antennal toruli (Fig. 10); postspiracular sclerite with small fovea located

anteriorly, leaving large, smooth triangular area posteriorly (Fig. 31); sidelobe

of mesoscutum without black areas along notauli E. iodes n. sp., p. 17

12 (10") Postspiracular sclerite with distinctly circular fovea, not co-extensive with entire

upper postspiracular sclerite (as in Fig. 37), smaller puncture below; scutellum

relatively short, SC:MSC = 1.24; margin of scrobes not flared at level of

antennal toruli (Fig. 7); large, about 7 mm E. magnus n. sp., p. 16

12" (10") Postspiracular sclerite with large fovea co-extensive with entire upper

postspiracular sclerite (Fig. 32); scutellum longer SC:MSC = 1.34-1.50;

margin of scrobes flared at level of antennal toruli (as in Fig. 11); smaller,

maximum length 6 mm E. solox n. sp., p. 16

THE NEW WORLD SPECIES OF EUPERILAMPUS

Euperilampus tanyglossa n. sp.

(Figs. 4, 5, 18, 25, 28, 33, 34, 35, 41, 42, 56, 58, 69, 70)

Type Locality. — Mexico, Jalisco, Zapotlanejo.

Type Material. — Holotype (Female, USNM No. 100317): Mexico, Jalisco, Zapotlanejo (Oct. 3 1966,

G.E./A.S. Bohart) [Specimen donated to USNM by USU]. Paratypes: Female, four Males, all from Mexico: Morelos, 10

mi E Cuernavaca, (Sept. 15 1972, Hanson/Poff) [Female, USU]; Zacatecas, 5 mi N Tabasco, (Sept. 18 1970; G.E./R.M.

Bohart) [2 males: USNM, BMNH]; Jalisco, 15 mi NE Guadalajara, (Sept. 17 1970, G.E./R.M. Bohart) [Male, USU];

Morelos, 6 mi E Cuernavaca, (Sept. 1 1970, Bohart/ Hanson) [Male, DCD].

Diagnosis. — Combination of an elongate labio-maxillary complex (Fig. 4) and

postspiracular sclerite with three or four indistinct foveae (Fig. 35) distinguish this species from

The New World species of Euperilampus

9

all New World species. E. aureicornis, which also has an elongate labio-maxillary complex, has

a single, indistinct fovea on the upper postspiracular sclerite and the funicle and clava of the

antenna yellow in both sexes {E. tanyglossa: female, dark brown; male, bright orange-yellow).

E. tanyglossa is distinguished from E. krombeini and E. triangularis group species by the

elongate labio-maxillary complex and the punctate-reticulate sculpture of the mesoscutum

(Fig. 33; cf. Fig. 12, E. krombeini and Fig. 16, E. triangularis).

Geographical distribution. — This species is distributed in the Central Plateau region of

Mexico. The six specimens were collected in five localities and in three different years. All

specimens have been collected between 1 September and 3 October. It is likely that this species

is widely distributed in the highlands of central Mexico but is rarely collected because adults

are present for only about one month of the year. A similar seasonal abundance pattern is found

in E. krombeini. The host is unknown. The elongate labio-maxillary complex suggests

associations with long-corolla flowers.

Derivation of specific epithet. — From the Greek {tany, ‘long’ and glossa, ‘tongue’), a

reference to the extremely elongate labio-maxillary complex.

Description. —

Female: Length, 4. 8-5. 8 mm. Head dark metallic green and violet; antennal scape metallic green, pedicel and anellus

brown, funicle and clava dark brown above, underside with light brown areas; labio-maxillary complex dark brown;

mandible reddish in middle, dark at base and apex. Mesosoma metallic green and violet; wings strongly darkened

throughout; coxae, trochanters and femora dark violet, tibiae dark brown without metallic reflections, tarsi yellow, pretarsi

dark brown. Metasoma metallic green.

Head: length of malar space 0.25-0.28 eye height; OOL 0.96-1.0 POL; frontal carina narrowly divergent, parallel with

inner eye margin, inner orbits not markedly narrowed at level of antennal toruli (Fig. 5); maximum width of scrobes about

one-third (0.33-0.35) head width; head transverse, width:height = 1.24-1.28; gena well developed, head widest across

genae; vertical costae of inner orbits (parascrobal spaces) short and irregular, surface thus wrinkled, extended onto face

and convergent with well developed orbital costae on clypeus as less distinct cross-arcuate costae; clypeus with well

developed transverse costae; clypeus transverse, width:height = 1.65-1.74, with sparse short setae except for patch of setae

(seven to nine) at each lateral ventral margin, upper margin straight, lower margin emarginate, without tentorial pits;

ocular-ocellar region and vertex costate; ocellar triangle almost smooth; supraclypeal area 0.59-0.60 clypeus height,

polished with two parallel lines from upper margin of clypeus to antennal toruli; margin of scrobes, in lateral view, merged

smoothly with face (Fig. 4); lower tooth of mandible rounded at apex; labio-maxillary complex extremely elongate (Figs. 4,

56). Antennae: pedicel and funicular segments subequal in length; funicular segments transverse, except elongate FI;

anellus 0.40 length of FI; scape narrowly linear, length 4.4-5. 1 maximum width.

Mesosoma: PN:MSC = 0.33-0.36; SC:MSC = 1.13-1.18; dorsum of pronotum and entire mesoscutum

punctate-reticulate (Fig. 33), punctures well defined, distinctly circular (Fig. 25) and coalesced in from of transverse rugae

only anteriorly on pronotum and along meson of mesoscutum; notauli indistinct; scutellum with short irregular transverse

rugae medially, punctate-reticulate laterally; apex of scutellum with distinct and indistinctly septate marginal rim (Fig.

34); sides of scutellum rounded, convergent at an angle of about 70 degrees; underside of scutellum smooth; propodeum

vertical, with wide but indistinctly impressed median area, submedian areas coriarious with transverse costae, callus

reticulate-rugose (Fig. 18); postspiracular sclerite gradually narrowed ventrally (Fig. 35), not sinuous as in E. triangularis

group, with three or four indistinct foveae; pronotum with smooth area laterally, below level of foveae on postspiracular

sclerite; axilla punctate-reticulate above, below with irregular costae; axillula with well developed longitudinal rugae.

Forewing: stigmal vein equal to or slightly longer than marginal vein, postmarginal about 3 times length of marginal vein

(Fig. 58).

Metasoma: smooth and shining without punctures; setae sparse; T2 with abrupt median concavity and Y-shaped

groove (Fig. 28), border between T2 and T3 sinuous and indistinct; T3 more quadrate than in E. triangularis group, length

about one-half maximum width.

Male: Length, 5. 2-6. 5 mm. Color as in female; except funicle and clava bright orange-yellow and underside of funicle

with dark transverse markings. Structure and sculpture as in female except; Head: length of malar space, 0.16-0.22 eye

height; head width:height = 1.20-1.24; clypeus width:height = 1.68-1.78; lateral wall of scrobes slightly more developed;

antennal scape, in frontal view, expanded only slightly, length 3. 5-4.0 maximum width, without distinct punctures (Fig.

42), in lateral view expanded apically with strong setae on outer surface, punctures well developed, surface distinctly

roughened (Fig. 41), inner surface with indistinct punctures; pedicel quadrate; funicle stouter. Mesosoma: PN:MSC =

0.34-0.37; SC:MCS = 1.10-1.21. Metasoma: T3 more transverse. Subgenital plate (Fig. 69): elongate, sides gradually

divergent, widest along sternite 8, width 1.43-1.50 length along midline [n = 2]. Genitalia (Fig. 70); digiti with four or five

large teeth and single smaller tooth; ventral lobe triangular, apex broadly rounded; lateral demelanized areas of

basiparamere large, pigmented median area much longer than length of digiti [n = 2].

Quaest. Ent., 1983, 19 ((1,2))

10

Darling

Euperilampus aureicornis n. sp.

(Fig. 6)

Type Locality. — Mexico, Guerrero, Amula [Almolonga on recent maps].

Type Material. — Holotype (Female, BMNH). Mexico, Guerrero, Amula 6000 ft., (Sept., H. H. Smith).

Godman-Salvin Coll. 1904.-1. Paratype: Male, same label data as holotype [BMNH].

Diagnosis. — Combination of elongate labio-maxillary complex (as in Fig. 4) and

postspiracular sclerite with a single fovea distinguish this species from all other New World

species. E. aureicornis is very similar to E. tanyglossa which also has the elongate

labio-maxillary complex, but differs in having only a single fovea on the postspiracular sclerite

(cf. three or four in E. tanyglossa) and funicle and clava of the antennae yellow in the female

(cf. brown in E. tanyglossa). E. aureicornis is distinguished from E. krombeini and E.

triangularis group species by the elongate labio-maxillary complex and the punctate-reticulate

sculpture of the mesoscutum (as in Fig. 33; cf. Fig. 12, E. krombeini and Fig. 16, E.

triangularis).

Geographical distribution. — This species is known from a single locality at 1829 m. in the

Sierra Madre del Sur. The host is unknown.

Derivation of specific epithet. — From the Latin {aureus, ‘golden’ and cornus, ‘horn’)

referring to the yellow antennae (funicle and clava) in both males and females.

Description. —

Female: Length, 4.1 mm. Head metallic green, scrobal cavity black; antennal scape metallic green, pedicel dark

brown, anellus, funicle and clava golden yellow; labio-maxillary complex dark brown; mandible yellow-brown in middle,

dark at base and apex. Mesosoma metallic green with violet reflections on pleurae; wings darkened throughout; coxae with

metallic violet reflections, femora, trochanters and tibiae deep brown, tarsi brown. Metasoma dark metallic green, with

bronzy reflections.

Head: length of malar space 0.23 eye height; OOL equal to POL; frontal carina oblique, convergent toward inner eye

margin, inner orbits markedly narrowed at level of antennal toruli (Fig. 6); maximum width of scrobes about one-half

(0.46) head width; head transverse, width:height = 1.30; gena well developed, head widest across genae; vertical costae of

inner orbits short and irregular, surface wrinkled, extended onto face and convergent with well developed outer orbital

costae at clypeus as less developed cross-arcuate costae; clypeus transverse, width:height = 1.54, evenly covered with

sparse short setae, smooth and shining with indistinct punctures, transverse costae only at extreme lateral margins, with

indistinct tentorial pits at about midpoint of lateral margins of clypeus (Fig. 6); ocular-ocellar region and vertex costate;

ocellar triangle almost smooth; supraclypeal area 0.62 clypeus height; margin of scrobes, in lateral view, merged smoothly

with face (as in E. tanyglossa, Fig. 4); lower tooth of mandible tapered to sharp point; labio-maxillary complex extremely

elongate (as in E. tanyglossa. Fig. 4); Antennae: pedicel and funicular segments subequal in length; funicular segments

transverse except elongate FI; anellus 0.38 length of FI; scape narrowly linear, length 4.8 maximum width.

Mesosoma: PN:MSC == 0.36; SC:MSC = 1.11; dorsum of pronotum and entire mesoscutum punctate-reticulate,

punctures coalesced in form of transverse rugae only anteriorly on pronotum and along meson of mesoscutum (as in E.

tanyglossa. Fig. 33); notauli indistinct; scutellum with short irregular transverse rugae medially, punctate-reticulate

laterally; apex of scutellum with distinct and weakly septate marginal rim; sides of scutellum rounded, convergent at angle

of about 70 degrees; underside of scutellum smooth; propodeum vertical, with shallowly impressed median area, with

indistinct transverse costae, raised submedian areas coriarious with transverse costae more dense than in E. tanyglossa (cf.

Fig. 18), callus reticulate-rugose; postspiracular sclerite gradually narrowed ventrally (as in E. tanyglossa. Fig. 35), with

single large fovea, and coriarious sculpture below fovea; pronotum with coriarious area laterally, below level of fovea on

postspiracular sclerite; axilla punctate-reticulate above, below with irregular rugae; axillula with well developed oblique

rugae, similar to E. triangularis (cf. Fig. 53). Forewing: stigmal vein slightly longer than marginal, postmarginal about

three times length of marginal vein (as in E. tanyglossa. Fig. 58).

Metasoma: smooth and shining without punctures; setae sparse; T2 with abrupt median concavity, border between T2

and T3 sinuous and indistinct; T3 more transverse than in E. tanyglossa length about one-third maximum width.

Male: Length, 3.75 mm. Color as in female. Structure and sculpture as in female except; Head: length of malar space,

0.20 eye height; head width:height = 1.35; clypeus width:height = 1.49, tentorial pits much deeper and larger; antennal

scape, in frontal view, expanded only slightly, length 4.15 maximum width, without distinct punctures, in lateral view

expanded apically with strong setae on lateral suface but without well developed punctures, surface smoother than in E.

tanyglossa (cf. Fig. 42), inner surface of scape with punctures more distinct and larger than in E. tanyglossa-, pedicel

quadrate. Mesosoma: PN:MSC = 0.36; SC:MSC = 1.11. Genitalia and subgenital plate not examined.

The New World species of Euperilampus

11

Euperilampus krombeini Burks

(Figs. 8, 12, 17, 29, 36, 38, 43, 59, 68, 73)

Euperilampus krombeini Burks, 1969: 79, (Figs. 6,9).

Type Locality. — U.S.A., Arizona, Tucson.

Type Material. — Holotype (female, USNM No. 69937) [examined]. Paratypes, 3 Females, 6 Males (USNM)

[examined; Allotype, USNM No. 69937, misidentified as male; Male paratype deposited in UK].

Material Examined. — U.S.A. (39 Females, 28 Males): Arizona (Cochise, Pima Cos.), New Mexico (Hildago

Co.). Mexico (6 Females, 2 Males): Sonora, Sinaloa, Chihuahua, Baja California Sur.

Diagnosis. — This is the only New World species of Euperilampus that lacks well developed

longitudinal costae or rugae on the inner orbits (parascrobal spaces) (Fig. 8). This species is

distinguished from species of the E. triangularis group by the lateral walls of the scrobe, which

merge smoothly with the face (Fig. 8; cf. Fig. 9, E. triangularis), and from E. tanyglossa and

E. aureicornis by the short labio-maxillary complex and the cross-arcuate sculpture of the

midlobe of the mesoscutum (Fig. 12; cf. Fig. 33, tanyglossa).

Geographical distribution. — This species has been collected primarily in the Sonoran and

Chihuahuan desert regions. All specimens have been collected in August and September. The

host is unknown and many specimens have been collected on flowers.

Description. — E. krombeini is redescribed primarily to allow comparison with other species,

and to include characters not in the original description. Boucek (1978) figured the apex of the

scutellum (his Fig. 11). Measurements presented in this redescription are based on 5 paratype

males and 5 females (Holotype, 2 paratypes, 2 specimens from the locality of these paratypes.

Continental, AZ.) [USNM].

Female: Length, 4. 2-5.0 mm. Color metallic violet and green, blues rarely seen, generally green with violet reflections,

except metasoma which is dark metallic green. Antennal scape metallic blue-green, pedicel, anellus, funicle and clava

brown; labio-maxillary complex brown; mandible reddish in middle, dark at base and apex, base with violet reflections;

wings darkened throughout; coxae, trochanters, femora and tibiae brown to reddish-brown, usually with distinct metallic

reflections, tibiae yellow apically, pretarsi dark brown.

Head: length of malar space 0.23-0.27 eye height; OOL 0.64-0.79 POL; maximum width of scrobes 0.34-0.37 head

width; head transverse, width:height = 1.23-1.34; gena well developed, head widest across genae; inner and outer orbits

(parascrobal spaces) without distinct costae or rugae (Figs. 8,38), costulae only on genae and on face laterad of clypeus;

clypeus transverse, width:height = 1.53-1.65, evenly covered with long setae, with indistinct punctures, surface polished,

upper margin straight, lower margin emarginate, without tentorial pits; ocular-ocellar region and vertex smooth, with

indistinct punctures, and glabrous area laterad of each posterior ocellus; vertex with well developed costae at posterior

margin; supraclypeal area 0.46-0.57 clypeus height; lateral wall of scrobes merged smoothly with face (Fig. 8); lower tooth

of mandible rounded at apex; base of mandible with distinct punctures; labio-maxillary complex short. Antennae: pedicel

and funicular segments subequal in length; funicular segments transverse, except elongate FI; anellus 0.31-0.40 length of

FI; scape narrowly linear, length 4. 0-4. 8 maximum width.

Mesosoma: PN:MSC = 0.36-0.37; scutellum short, slightly longer than mesoscutum, SC:MSC = 1.04-1.17; dorsum

of pronotum punctate-reticulate, punctures coalesced in form of indistinct irregular transverse costulae medially, lateral

punctures distinctly circular; midlobe of mesoscutum with incomplete irregular transverse rugae, reticulate along notauli,

sculpture in many specimens less distinct mesad (Fig. 12); sidelobe of mesoscutum completely sculptured, with punctures

anteriorly along notauli, laterally reticulate-rugose to rugose posteriorly; notauli distinct; scutellum with well developed

cross-arcuate costae, underlying surface with punctures along axillula; apex of scutellum without distinctly septate

marginal rim; scutellum broadly rounded (Fig. 12); underside of scutellum smooth; propodeum vertical, with distinct

median furrow, submedian areas polished with dense transverse costulae, callus reticulate-rugose (Fig. 17); postspiracular

sclerite abruptly narrowed ventrally, sinuous, with shallow fovea coextensive with most of upper postspiracular sclerite, in

many specimens with faint punctures below fovea (Fig. 36); axilla reticulate-rugose above, below with irregular rugae;

axillula with one to three well developed oblique costae (Fig. 36). Forewing: stigmal vein slightly shorter than marginal,

postmarginal about three times length of marginal vein (Fig. 59).

Metasoma: T2 with median longitudinal row of closely spaced punctures extended two-thirds distance to T2/T3

border and with lateral arcuate lines of punctures joined to apex of median row (Fig. 29); T2 with sparse setae, without

punctures, border between T2 and T3 sinuous and indistinct; T3 more quadrate than in E. triangularis group, length about

one-half maximum width (0.52-0.57), with lateral patches of setae, and distinct punctures; T4 and T5 with well developed

punctures extended transversely across anterior surface of terga.

Quaest. Ent., 1983, 19 ((1,2))

12

Darling

Male: Length, 3. 7-4. 6 mm. Color as in female. Structure and sculpture as in female, except; Head: antennal scape,

in frontal view, expanded only slightly, length 3. 9-4.4 maximum width, without punctures, outer surface with strong

setae and roughened with punctures (Fig. 43), inner surface without distinct punctures; pedicel quadrate; anellus

relatively shorter, 0.10-0.26 length of FI; funicle stouter. Mesosoma: PN:MSC = 0.33-0.40; SCiMSC = 1.08-1.26.

Metasoma: T3 more transverse, length about one-third width. Subgenital plate (Fig. 68): transverse, sides gradually

divergent, widest along sternite 8, width 2.05-2.23 length along midline [n = 4]. Genitalia (Fig. 73): digiti with three or

four large teeth and single smaller tooth; ventral lobe acuminate, apex broadly rounded; lateral demelanized areas of

basiparamere large, pigmented median area about equal in length to digiti.

Euperilampus triangularis group

Diagnosis. — This species group is characterized by massive scrobal walls which are sharply

angulate at the level of the antennal toruli and merge abruptly with the face (Fig. 9). The apex

of the scutellum is acuminate (Figs. 13-16), not broadly rounded as in E. krombeini (Fig. 12).

The labio-maxillary complex is short (Fig. 9), not elongate as in E. tanyglossa and E.

aureicornis (Fig. 4).

Geographical distribution. — This species group is widely distributed in eastern North

America, into central Quebec, and south to Florida; in the montane regions of Mexico and in

the montane regions of southern Brazil, northern Argentina, eastern Bolivia and Paraguay.

There are no specimens from northern South America or Central America. This disjunction

may simply be due to a paucity of montane collections.

Description. — A group description is provided, based on the presently included species: E.

triangularis, E. gloriosus, E. brasiliensis, E. enigma, E. luteicrus, E. ameca, E. iodes, E.

solox, and E. magnus. Measurements and ratios are presented in general terms and the ranges

of numerical values are listed in Table 1 for the species represented by multiple specimens. The

group descriptors are not repeated in the descriptions of the included species; a full description

of each species is the group description with the appropriate numerical values from Table 1 and

the species description.

Female-. Length, 3. 0-7. 5 mm. Head, mesosoma and metasoma metallic (iridescent) violets, blues and greens with violet

reflections, in some species with contrasting glossy black areas on head and mesosoma; antennal scape metallic green,

pedicel brown with metallic reflections, anellus, funicle and clava dark brown, labio-maxillary complex dark brown;

mandible, basally metallic, distally reddish-brown; wings darkened throughout; coxae, trochanters, femora and tibiae dark

with metallic reflections (fore and mid tibiae yellow in E. luteicrus)-, distal end of tibiae light brown, tarsi yellow, pretarsi

dark brown.

Head-. Length of malar space about one-third eye height; lower margin of eyes extended to top of clypeus; base of

mandible with distinct punctures; OOL approximately equal to POL; maximum width of scrobes one-third to one-half

head width; head transverse, width:height = 1.0- 1.2; vertical costae of inner orbits (parascrobal spaces) well developed,

extended onto face and convergent with outer orbital costae on clypeus (Fig. 9); clypeus transverse, width about 1.5 times

height, with costae at lateral margins, evenly covered with long setae; upper margin concave or straight, lower margin

emarginate, without tentorial pits; vertex with welt developed carina(e) at posterior margin and glabrous area laterad of

each posterior ocellus; supraclypeal area about one-half clypeus height, glabrous and polished; lateral wall of scrobes

merged abruptly with face, at level of antennal toruli (Fig. 9); lower tooth of mandible tapered to sharp point;

labio-maxillary complex short (Figs. 9,55), extended just beyond mandibles. Antennae: pedicel and funicular segments

subequal in length; funicular segments transverse except elongate FI; relative length of anellus to FI diagnostic; scape

narrowly linear, length about 5-6 times maximum width.

Mesosoma-. sculpture diagnostic; pronotum one-third to one-half length of mesoscutum; scutellum longer than

mesoscutum, SC:MSC diagnostic; notauli distinct; apex of scutellum with marginal rim; propodeum vertical, with median

impressed foveae and raised submedian areas with transverse costae, callus reticulate-rugose (Fig. 19); postspiracular

sclerite abruptly narrowed ventrally, sinuous, with single large fovea dorsally, size and shape of which is diagnostic, with

some smaller punctures below fovea in some species. Forewing: stigmal vein shorter than marginal, postmarginal vein

about three times length of marginal vein (Fig. 57).

Metasoma-, smooth and shining, T2, T4, and T5 evenly covered with setae, T3 with lateral patches of setae, T4 with

indistinct punctures at base of setae, punctures on T5 indistinct or well developed; T2 smoothly concave (Fig. 27), border

between T2 and T3 sinuous and indistinct; T3 transverse, length about one-third maximum width (0.33-0.39).

The New World species of Euperilampus

13

Male: Length, 2.0^. 0 mm. Color as in female. Structure and sculpture as in female except; Head; relative length of

malar space to eye height smaller, antennal scape, in frontal view, expanded slightly apically, length about four times

maximum width; distribution of punctures and setae diagnostic; pedicel quadrate, shorter than FI; relative length of

anellus to FI shorter; funicle stouter. Mesosoma: costae on submedial areas of propodeum more prominent. Metasoma:

T3 more transverse. Subgenital plate: subquadrate, abruptly expanded along sternite 8 (Fig. 67), width:length,

1.30-1.83 (n=15). Genitalia: diagnostic.

Euperilampus triangularis (Say)

(Figs. 1, 2, 3, 9, 1 1, 16, 19, 20, 21, 27, 37, 39, 44, 53, 55, 57, 67)

Perilampus triangularis Say, 1828:78. [Type lost].

Euperilampus triangularis-, Crawford, 1914:69.

Type Locality.— U.S.A., Indiana.

Material Examined. — U.S.A. (310 Females, 182 Males): Alabama, Arkansas, Connecticut, District of

Columbia, Florida, Illinois, Indiana, Iowa, Kansas, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri,

New Hampshire, New Jersey, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, South Dakota, Texas,

Vermont, Virginia, West Virginia, Wisconsin. Canada (8 Females, 6 Males): Quebec; Aylmer, Lac Chicobi, Montreal.

Ontario: Ottawa, Point Pelee, Ruby.

Notes about synonymy. — Burks (1969) compared E. triangularis with E. krombeini and

illustrated the head of E. triangularis (his Fig. 1 1) as £. hyalinus. {lapsus calami) in the figure

legends. Boucek (1978) illustrated the apex of the scutellum of E. triangularis (his Fig. 10). E.

triangularis sensu Boucek is coextensive with my E. triangularis species group.

Diagnosis. — This species is reliably distinguished from other members of the E.

triangularis group by having a concolorous mesoscutum, i.e., without contrasting black areas

on the sidelobe, and axillulae with distinct costae, the most ventral of these merging with the

posterior border of the axillulae (Fig. 53; cf. Fig. 54, E. brasiliensis). Sculpture on the

scutellum is less developed along the midline, the rugae incomplete (Fig. 16; cf. Fig. 13, E.

iodes and Fig. 14, E. brasiliensis and Fig. 15, E. solox).

Geographical distribution. — This species is widely distributed in eastern North America,

extending to western South Dakota (Custer Co., Lawrence Co.), eastern Kansas (Riley Co.),

and east Texas (Galveston Co.). The distribution matches closely the extent of broadleaf

deciduous forests. Western extremes in the range are along tributaries of the Missouri River:

riparian areas with broadleaf deciduous forests. Representative collection dates are as follows:

New York, June 30 - September 1; Massachusetts, June 24 - September 10; Quebec, July 1 -

August; Kansas, June 20 - July 1.

E. triangularis has been collected frequently in Highlands Co., Florida at the Archbold

Research Station [FSCA]. This is an area of needleleaf evergreen forest and violates the

distributional correlates established above. These specimens show some morphological

peculiarities (see ‘Variation’ section). Collection dates also differ dramatically from the

northeastern material and range from March 2 - May 30 (based on 70 specimens). Other

specimens from Florida (all northern Florida) also were collected between these dates. This

species has not been recorded from Georgia, South Carolina, Mississippi, Tennessee, Kentucky

or Louisiana. Excluding Florida, North Carolina is the most southern record of this species

along the east coast, and specimens have been taken May 16 (Columbus Co., coastal). May 20

(Black Mts.), July 27 (Haywood Co., Blue Ridge Mts.), September 23 (Buncombe Co., Blue

Ridge Mts.).

The temporal separation of E. triangularis into spring and late summer collection dates is

probably related to overwintering of hosts in northern and montane areas. Direct development

of the hosts in the southern extremes of the range would explain occurrence of specimens in

April and May. Unfortunately the host(s) of Euperilampus triangularis in Florida is unknown.

Quaest. Ent., 1983, 19 ((1,2))

14

Darling

Boucek examined North American material and specimens from Santa Catarina, Brazil

(Nova Teutonia). Boucek (1978) lists material from Colorado, as does Peck (1951, 1963), and

Burks (1979), apparently in reference to Ashmead (1890). I have seen no specimens from the

Rocky Mountains or from Colorado.

Host associations. — Euperilampus triangularis is a secondary parasite (hyperparasite) of

the fall webworm, Hyphantria cunea (Drury) [Arctiidae]. Warren and Tadic (1970) reared E.

triangularis as a parasitoid of Hyposoter fugitivus (Say) [Ichneumonidae] which attack the

fall webworm as primary parasitoids (13 females, 28 males). This material was misidentified as

Perilampus hyalinus, although P. hyalinus was also reared from Hyposoter (six females, two

males) and other primary parasitoids [series examined, UA]. Rearings of Hyphantria cunea in

New York (Darling, unpublished) have also yielded both P. hyalinus and E. triangularis.

Other parasitoids obtained in the New York rearings included only Tachinidae [Eusisyropa

blanda (O.S.), Blondelia hyphantriae (Tothill) and Merida ampelus (Wlk.)] and

Ichneumonidae [Therion spp. and Sinophorus validus (Cresson) complex] [CU, UG, DCDj.

The host of Euperilampus triangularis in New York has yet to be determined. I also have seen

a specimen of E. triangularis reared from H. cunea from Ruby, Ontario, Canada [CNCj.

Description. — The redescription and measurements are based on 17 fem.ales and 18 males,

three specimens of each sex from the following six localities, except where noted otherwise:

Florida, Highlands Co., Archbold Biological Station [FSCA]; Virginia, Arlington Co.,

Kearney Sta. [USNMj; Arkansas, Washington Co., Fayetteville, Ex: Hyposoter spp, parasite

of Hyphantria cunea [UA]; Canada, Quebec, various localities, and 1 Female from Ruby, Ont.

[CNC, USNM]; Kansas, various localities (only 2 Females examined) [UK, KSU, USNM];

New York, Seneca Co., Geneva, Ex: culture of Hyphantria cunea [DCD, UG].

Female: Length, 2.9-6. 3 mm. Color ranging from blue-violet to blue-green, with violet reflections on mesoscutum and

green reflections on metasoma, without contrasting black areas on sidelobe of mesoscutum.

Head: maximum width of scrobes 0.28-0.35 head width; margin of scrobes, in frontal view, sinuous and flared at level

of antennal toruli (Fig. 1 1); inner orbital costulae irregularly spaced, wavy to rugose (Fig. 9), convergent toward posterior

ocellus and not extended through ocular-ocellar region (Fig. 21); outer orbital costae various, restricted to malar region or

extended to vertex, less developed above; vertex almost completely sculptured (punctures and indistinct rugae), except for

glabrous area laterad of each posterior ocellus; clypeus with indistinct arcuate costae at extreme lateral margins, punctures

well developed and dense, surface appearing roughened. Antennae: anellus 0.18-0.25 length of FI, relatively shorter than

in other species (see Table 1).

Mesosoma: scutellum longer than mesonotum, SC:MSC = 1.42-1.60; dorsum of pronotum punctate-reticulate,

punctures coalesced in form of transverse rugae anteriorly (as in Fig. 26), punctures along midline polygonal, not distinctly

circular; midlobe of mesoscutum with transverse rugae, more irregular (reticulated) posteriorly, sidelobe of mesoscutum

posteriorly rugose, anteriorly along notauli roughened with irregular punctures, laterally punctate-reticulate; scutellum

with arcuate rugae, less developed along midline, and longitudinally rugose-reticulate laterally (Fig. 16); apex of scutellum

with distinctly septate marginal rim; scutellum acuminate, sides convergent at about 60 degrees; underside of scutellum

sculptured (Fig. 39); median area of propodeum with deeply impressed foveae, in form of distinct X-shape (Fig. 19),

submedian areas polished with cross-arcuate costulae throughout; postspiracular sclerite with large, round,

centrally-located fovea (Fig. 37); axilla reticulate-rugose above, rugose below; axillula smooth and shining, with one to

three short oblique costae, ventral costa in most specimens merged with posterior border of axillula (Fig. 53).

Metasoma: T5 with indistinct punctures.

Male: Length, 2. 3-5. 2 mm. Color as in female. Structure and sculpture as in female except. Head: antennal scape, in

frontal view, expanded slightly apically, smooth (Fig. 44); anellus relatively smaller, 0.10-0.15 length of FI, relatively

shorter than other species (see Table 1). Mesosoma: propodeum more coarsely sculptured with more prominent

cross-arcuate costae. Genitalia (Figs. 1,2): digiti with three, four, or five large teeth and two or three smaller teeth; ventral

lobe rounded, not distinctly acuminate but in few specimens notched at apex; lateral demelanized areas of basiparamere

large and quadrate, not reduced laterally, pigmented median area shorter than length of digiti [n-17].

Variation. — The reared series of E. triangularis from Arkansas differs considerably from

wild caught specimens. The Arkansas series consists of extremely small individuals about 3 mm

in length. In these specimens the sculpture of the propodeum is quite distinctive with the

median triangle about half the height of the propodeum (compare Fig. 20, reared from

The New World species of Euperilampus

15

Hyposoter and Fig. 19, wild caught specimens from Archbold Research Station). Of the total

material examined, only five wild caught specimens (1.1%) are as small as this reared material

(MSC < .75 mm). In these wild caught specimens, the propodeum is of the standard

configuration, with the median triangle quite small. It is possible that the rearing conditions

were sub-optimal, resulting in these abnormally small individuals, with the associated variation

in propodeum structure. All wild caught specimens from Arkansas (n = 3) are of normal size

(5.00-5.83 mm) and the propodeal triangle is not enlarged. A single small specimen from Ruby,

Ontario [CNC], again reared from H. cunea, has the unusually large propodeal triangle.

Florida specimens also differ from northeastern material. Inner orbital costae and transverse

rugae of the midlobe of the mesoscutum are very irregular, especially in males. Florida

specimens also have relatively longer scutella, SC:MSC, than specimens from other areas

(males, 1.69-1.75; all other localities, 1.42-1.60. females, 1.55-1.56; all other localities,

1.42-1.55). Male genitalia of New York specimens (n = 6), Florida (n = 6), Virginia (n = 2),

and Arkansas, reared from Hyposoter (n = 2), show no consistent differences.

The apex of the scutellum is quite aberrant in four specimens. Variants include deeply cleft

and bilobed apices, and asymmetrical developments of the normal acuminate apex. The most

striking variant is a deeply bilobed apex which is bent under the vaulted part of the scutellum so

that the marginal rim is not visible in dorsal view. This specimen, a female from Highlands Co.,

Florida, was among four normal specimens from the same Malaise trap collection [FSCA]. In

addition, the reared series from Arkansas have many specimens with truncate scutella,

shallowly cleft at the apex.

The status of the Florida population will have to be reconsidered when the host

association(s) is determined. Possibly this population represents a sibling species related to E.

triangularis. Description of such a sibling species from eastern North America will present

‘exceptional circumstances’, as outlined by Article 75 of the International Code of Zoological

Nomenclature. A neotype will have to be designated for Say’s species, to fix the name with the

northern population. A suitable topotypic specimen is in the Cornell collection, [Female:

Indiana, Madison, July 29 1957, H.E. Evans]. This specimen agrees with all the particulars of

this redescription.

Euperilampus gloriosus (Walker)

Perilampus gloriosus Walker, 1862:375.

Euperilampus gloriosus-. Walker, 1871:67.

Euperilampus gloriosus-, Boucek, 1978:304 [lectotype designation].

Type Locality. — Mexico.

Type Material. — Lectotype (male, BMNH) [not examined].

Diagnosis.— The following is based on notes of A.B. Gahan [USNM], Burks (1969), and

Boucek (1978). This species can readily be distinguished from all other New World species of

Euperilampus by the scutellum, which is rather abruptly produced into a lanceolate spine (Fig.

9 in Boucek 1978) and by the bright coppery to fiery golden color of the apex of the scutellum

and the metasoma.

The species is known only from the type material. The specimen was collected by M. Salle;

the exact locality in Mexico is apparently unknown.

Quaest. Ent., 1983, 19 ((1,2))

16

Darling

Euperilampus magnus n. sp.

(Fig. 7)

Type Locality. — Mexico, Chiapas, Chiapa de Corzo.

Type Material. — Holotype (Female, CAS): Mexico, Chiapas, Municipio Chiapa de Corzo, El Chorreadero, 670

m (Aug 16 1976, DE/JA Breedlove).

Diagnosis. — The relatively short scutellum,. SCiMSC = 1.24, separates the holotype from

all species except the E. brasiliensis complex. The specimen differs from E. brasiliensis

complex females by the absence of contrasting black areas on the sidelobe of the mesoscutum

and by the margin of the scrobes not being flared at the level of the antennal toruli (Fig. 7).

The reliability of size in recognizing this species must await further collecting. This specimen is

1.2 times larger than any other New World specimen.

Geographical distribution. — The type locality is in the interior highlands of Chiapas and is

the only Mexican locality for Euperilampus, east of the Isthmus of Tehuantepec.

Derivation of specific epithet. — From the Latin, with reference to the large size of the

holotype.

Description. —

Female-. Length, 7.5 mm. Color ranging from blue-violet to blue-green, with violet and green reflections, and glossy

black posteriorly on vertex, but without contrasting black areas on sidelobe of mesoscutum.

Head-. Length of malar space 0.36 eye height; OOL = 0.83 POL; maximum width of scrobes 0.35 head width; margin

of scrobes, in frontal view, smoothly curved, not flared at level of antennal toruli (Fig. 7); gena very well developed, head

widest across genae; head transverse, width:height = 1.2; inner orbital costulae irregularly spaced, convergent on posterior

ocellus and not extended through ocular-ocellar region (as in E. triangularis. Fig. 21); outer orbital costulae extended past

point of maximum width of eye but not to vertex; vertex almost smooth, posteriorly with seven transverse costulae; clypeus

transverse, widthiheight = 1.55, with indistinct arcuate costae at extreme lateral margins, punctures distinct and dense,

surface appearing roughened; supraclypeal area 0.47 clypeus height. Antennae: anellus 0.18 length of FI; scape narrowly

linear, length 4.6 times maximum width.

Mesosoma: PN:MSC = 0.50; scutellum slightly longer than mesoscutum, SC:MSC = 1.24; dorsum of pronotum

punctate-reticulate, interspaces widened and punctures distinctly circular; midlobe of mesoscutum rugose, transverse

rugae less wavy than in E. solox (cf. Fig. 15), sidelobe of mesoscutum posteriorly rugose, anteriorly along notauli

roughened with irregular punctures, laterally punctate-reticulate; scutellum with arcuate rugae, less distinet on disk; apex

of scutellum with distinctly septate marginal rim; scutellum acuminate, sides convergent at about 70 degrees; underside of

scutellum smooth; median area of propodeum shallowly impressed, without deep foveae, submedian areas with swirling

costae; postspiracular sclerite with large, round, centrally-located fovea, and smaller puncture below; axilla

reticulate-rugose above, rugose below; axillula smooth and shining, with one to three short oblique costae not merged with

posterior border.

Metasoma-. T5 with distinct punctures.

Male-. UNKNOWN.

Euperilampus solox n. sp.

(Figs. 15, 32, 47,48, 76)

Type Locality. — Argentina, Tucuman, Tacanas.

Type Material. — Holotype (Female, IML): Argentina, Tucuman, Tacanas (Nov. 5 - 30 1968, L. Stange).

Paratypes: (4 Females, 2 Males) all from Argentina. Tucuman, Trancas to Tacanas (Nov. 1-30, Stange) [Female,

BMNH], (Jan., Arnau) [Female, USNM]. Salta: Alemanla (April, Stange/ Porter) [Female, CNC, identified as

Euperilampus triangularis (Say) in Fidalgo 1980: 194]. Tucuman: San Pedro Colalao (Foerster), [Female, lESMj. Salta:

Cerro San Bernardo (Feb., Monros/ Willink). [Male, IML]. Tucuman: Trancas, San Pedro Colalao (Feb., Arnau) [Male,

CNC].

Diagnosis. — E. solox is recognized by the reticulate-rugose sculpture of the midlobe of the

mesoscutum (Fig. 15). The mesoscutum does not have contrasting black areas, which

distinguishes the species from the E. brasiliensis complex. The fovea of the postspiracular

sclerite is large but indistinctly impressed (Fig. 32) and the margins of the scrobes are flared at

The New World species of Euperilampus

17

the level of the antennal toruli (as in Fig. 11), distinguishing this species from E. iodes (cf. Fig.

10. E. iodes). E. solox does not have the reduced sculpture on the disk of the scutellum, which

is characteristic of E. triangularis (Fig. 15; cf. Fig. 16. E. triangularis).

Geographical distribution. — E. solox is known only from northern Argentina. The host is

unknown.

Derivation of specific epithet. — From the Latin, solox, for ‘coarse or rough’ with reference

to the irregular sculpture of the mesoscutum.

Description. —

Female-. Length, 5. 4-6. 2 mm. Color predominantly blue-green, with violet reflections, eontrasting black areas

restricted to vertex, not on mesoscutum.

Head: maximum width of scrobes 0.41-0.43 head width; margin of scrobes, in frontal view, sinuous and flared toward

eye margin at level of antennal toruli (as in E. triangularis. Fig. 11); inner orbital costae rugose, extended to ocular-ocellar

region; outer orbital costulae extended to vertex, less distinct above; vertex completely sculptured (coarse punctures and

indistinct rugae), except for glabrous area laterad of each posterior ocellus; clypeus with indistinct arcuate costae at

extreme lateral margins, punctures distinct and dense, surface appearing roughened. Antennae: anellus 0.26-0.35 length of

FI.

Mesosoma: scutellum longer than mesoscutum, SC:MSC = 1.38-1.48; dorsum of pronotum punctate-reticulate,

punctures not coalesced in form of distinct transverse rugae anteriorly, punctures circular or polygonal, interspaces

widened; mesoscutum reticulate-rugose, with very irregular rugae anteriorly on midlobe (Fig. 15) and posteriorly on

sidelobe, sculpture reduced along notauli; scutellum with irregular cross-arcuate rugae, laterally reticulate-rugose; apex of

scutellum with distinctly septate marginal rim; scutellum acuminate, sides convergent at about 70 degrees; underside of

scutellum sculptured at apex; median area of propodeum with deeply impressed foveae, in form of distinct V-shape,

submedian areas polished, with seven or eight well developed and evenly distributed cross-arcuate costae; postspiracular

sclerite with shallow circular fovea co-extensive with most of upper postspiracular sclerite (Fig. 32); axilla

reticulate-rugose above, rugose below; axillula smooth and shining, without costae extended to posterior border (as in E.

brasiliensis, Fig. 54).

Metasoma: T5 with well developed punctures, stronger than in other species of E. triangularis group.

Male: Length, 3. 7-4. 2 mm. Color as in female. Structure and sculpture as in female except. Head: Antennae: anellus

relatively smaller; scape, in frontal view, expanded apically, with distinct punctures on anterior and inner surfaces, outer

surface with strong setae, surface roughened (Figs. 47,48). Genitalia (Fig. 76): digiti with two to five large teeth and two

smaller teeth; ventral lobe broadly rounded, notched in single specimen; lateral demelanized areas of basiparamere large

and quadrate, not reduced laterally, pigmented median area slightly shorter than digiti [n = 2].

Variation. — The sculpture of the mesonotum is less distinct in some specimens. The female

specimen (USNM: Trancas to Tacanas) is a deep emerald green color due to cleaning the

specimen in lacto-phenol to remove a fungal coating.

Euperilampus iodes n. sp.

(Figs. 10, 13, 23, 24,31,51,52,72)

Type Locality.— Brazil, Santa Catarina, Nova Teutonia.

Type Material. — Holotype (Female, CNC No. 17004): Brazil, Santa Catarina, Nova Teutonia, 300 - 500 m,

(Feb. 1968, F. Plaumann). Paratypes (4 Females, 2 Males), same locality and collector as holotype (Aug., Sept., Oct.)

[BMNH,CNC, IML].

~ Additional Material Examined. — Brazil, Sao Paulo, Barreiro, Serra de Bocaina, 1650 m (Sept., Alvarenga

and Seabra) [Female, AEI]. Mexico, Jalisco, 15 mi. S. Lagos de Moreno (Aug., 1962) [Female, NHMLAC].

Diagnosis. — This is the only species with the fovea of the postspiracular sclerite small and

located on the anterior portion of the sclerite, leaving a large smooth triangular area posteriorly

(Fig. 31). The concolorous mesoscutum and the relatively longer scutellum distinguish E. iodes

from the E. brasiliensis group (see Table 1). This species differs from E. solox is having the

margin of the scrobe smoothly curved, not flared at the level of the antennal toruli (Fig. 10; cf.

Fig. 11, flared at level of toruli), and more regular sculpture on the mesoscutum (Fig. 13; cf.

Fig. 15, E. solox). This species has the sculpture of the scutellum complete (Fig. 13), not

reduced on the disk of the scutellum as in E. triangularis, (Fig. \6). E. iodes is the only species

Quaest. Ent., 1983, 19 ((1,2))

18

Darling

with costae extending through the ocular-ocellar region along the eye margin (Figs. 23, 24; cf.

Fig. 21, iF. triangularis and Fig. 22. E. brasiliensis).

Geographical distribution. — E. iodes is sympatric with E. brasiliensis at Nova Teutonia,

Brazil. Specimens are known only from Mexico and Brazil, and the host is unknown. The

description and measurements are based on the type material.

Derivation of specific epithet. — From the Greek, iodes, ‘violetlike’, with reference to color

of adults of this species.

Description. —

Female: Length, 4. 2-5. 4 mm. Color predominantly blue-violet, with green reflections on scrobal cavity, supraclypeal

area, pleuron of mesosoma and metasoma; without contrasting black areas on vertex and mesoscutum.

Head: maximum width of scrobes 0.38-0.40 head width; margin of scrobes, in frontal view, smoothly curved, not flared

at level of antennal toruli (Fig. 10); inner orbital costae regularly spaced, outermost costulae extended around top of eye,

through ocular-ocellar region (Figs. 23, 24); outer orbital costulae extended to vertex, less distinct above, but much more

distinct along eye margin; vertex almost smooth, posteriorly with transverse costae; clypeus with well developed costae at

lateral margins, punctures indistinct and sparse, surface smooth. Antennae: anellus 0.27-0.32 length of FI.

Mesosoma: scutellum longer than mesoscutum, SC:MSC = 1.38-1.50; dorsum of pronotum punctate-reticulate,

punctures coalesced in form of distinct transverse rugae anteriorly, punctures along midline polygonal, not distinctly

circular; midlobe of mesoscutum and entire scutellum with incomplete but regular cross-arcuate costae, sidelobe of

mesoscutum posteriorly roughened, anteriorly along notauli with indistinct irregular punctures, laterally

punctate-reticulate to rugose posteriorly (Fig. 13); apex of scutellum with distinctly septate marginal rim; scutellum

acuminate, sides convergent at about 70 degrees; underside of scutellum sculptured at apex; median area of propodeum

with deeply impressed foveae, in form of distinct V-shape, submedian areas smooth and shining, with indistinct

cross-arcuate costulae dorsally; postspiracular sclerite with small fovea extended about one-half upper postspiracular

sclerite, fovea anterior on postspiracular sclerite, with smooth triangular area in upper posterior corner (Fig. 31), axilla

rugose; axillula smooth and shining, many specimens with 1-3 short costae.

Metasoma: T5 with indistirict punctures.

Male: Length, 3. 7-3. 9 mm. Color as in female, but contrasting black areas on vertex. Structure and sculpture as in

female except. Head: Antennae: anellus relatively smaller, 0.14-0.25 length of FI; scape, in frontal view, expanded slightly

apically, punctures on anterior and inner surfaces, outer surface with strong setae, surface roughened (Figs. 51, 52).

Mesosoma: sculpture of sidelobe of mesoscutum either female condition or completely rugose along notauli; propodeum

more coarsely sculptured with more prominent cross-arcuate costae and with fovea on submedian areas. Genitalia (Fig.

72): digiti with five large teeth and two smaller teeth; ventral lobe acuminate, not broadly rounded; lateral demelanized

areas of basiparamere large and quadrate, pigmented median area shorter in length than digiti [n = 2].

Variation. — The specimen from Barreiro, Brazil, differs from the type series in having the

fovea of the postspiracular sclerite somewhat larger, and the inner orbital costae less developed

along the top of the eye. The Mexican specimen is similar to the Barreiro specimen, but the

lateral wall of the scrobe is not as prominent in lateral view and the wings are hyaline, not

distinctly darkened.

Euperilampus brasiliensis complex

Diagnosis. — All species have black areas on the sidelobe which contrast with the otherwise

metallic color. These areas are quite large and occupy the anterior one-half of the sidelobe,

except in E. ameca (reduced to a narrow band along the notauli). Scutella are relatively shorter

than in all other species of the E. triangularis group (Fig. 14), SC:MSC ranging from 1.19 to

1.31. The only exception is the holotype of E. luteicrus, SC:MSC = 1.45. The short scutella

result in the sides of the scutellum being convergent at an angle of about 75 degrees (60 - 70

degrees in other species of the E. triangularis group). The upper postspiracular sclerite has a

single large fovea which is not distinctly circular (Fig. 30).

Taxonomic note. — This complex is defined to encompass species related to E. brasiliensis

as diagnosed above {E. enigma, E. luteicrus, and E. ameca). The paucity of material has

presented many problems; all species are allopatric, and the three new species are based on

single specimens. Association of sexes is further complicated by the fact that the type material

The New World species of Euperilampus

19

of E. brasiliensis is represented only by females.

Females of this complex from central South America are indistinguishable, but two distinct

forms of males are present. A series of seven females from Nova Teutonia, Santa Catarina,

Brazil, are here regarded as conspecific with the type of E. brasiliensis [type locality-

Chapada, Brazil]. Three males from Nova Teutonia appear to be conspecific and are here

described as the male of E. brasiliensis. A very distinctive male from Robore, Bolivia, is

described as E. enigma n. sp. This could be the male of E. brasiliensis', if so, the Nova Teutonia

material would represent a new species. I decided to treat the Bolivian male as a new species,

since this is a more conservative solution. The other possibility would be to associate the

Bolivian male with the type material of E. brasiliensis, and to describe a new species for the

Nova Teutonia material based on differences from the associated, Bolivian male. As treated

here, E. brasiliensis and E. enigma are separated by the Paraguay River.

This complex is represented in Mexico by two new species, each based on a single specimen.

The hosts are not known for any species of this complex.

Euperilampus brasiliensis (Ashmead) n. comb.

(Figs. 14, 22, 26, 30, 40, 49, 54, 75)

Perilampus brasiliensis Ashmead, 1904: 467 (Plate 34, Fig. 4).

Type Locality. — Brazil, Chapada [Matto Grosso]. [Holland (1919:482) discusses the various expeditions of

H.H. Smith. The 1881-1886 trip to Brazil spent considerable time along the upper waters of the Rio Paraguay and Rio

Guapore in western Brazil near Chapada and Matto Grosso. Much of this material went to the Carnegie Museum and was

studied by Ashmead.]

Type Material. — Lectotype (Female, USNM No. 56960) [Present designation]: Chapada, April, H.H. Smith

Coll. Paralectotype (Female, USNM Paratype No. 56960): Chapada, Nov.

Material Examined. — Brazil (12 Females, 4 Males); Santa Catarina, Nova Teutonia 300 - 500 m. (Jan. -

June, August, September; F. Plaumann) [AEI, BMNH, CNC, UK]. Argentina (1 Female, 1 Male) Misiones: Loreto

(Dec.) [Female, UNLP], Iguazu (March) [Male, IML; identified as Euperilampus triangularis (Say) in Fidalgo 1980:

194]. Paraguay (1 Female): Independencia (Sept.) [UNLP].

Diagnosis. — Only the male of this species can be distinguished from other members of the

E. brasiliensis complex. The male genitalia lack enlarged, recurved teeth on the digitus (Fig.

75; cf. Fig. 74, E. enigma), and the scape of the antenna has very weak punctures on the inner

surface (Fig. 49; cf. Fig. 50, E. enigma'. Figs. 45, 46, E. luteicrus).

Description. — The redescription and measurements are based on the type material and the

specimens from Nova Teutonia.

Female: Length, 4. 6-6. 3 mm. Color predominantly blue-green, with violet reflections. Vertex, midlobe and sidelobe of

mesoscutum, with contrasting glossy black areas.

Head: maximum width of scrobes 0.37-0.44 head width; margin of scrobes, in frontal view, sinuous and flared at level

of antennal toruli (as in E. triangularis. Fig. 11); head widest across eyes; inner orbital costae not convergent on posterior

ocellus or extended around top of eye, ended abruptly at ocular-ocellar region (Fig. 22); outer orbital costae short, largely

confined to malar region; vertex almost smooth, posteriorly with transverse costae; clypeus with indistinct arcuate costae at

extreme lateral margins, punctures indistinct, surface appearing smooth and polished. Antennae: anellus 0.23-0.33 length

of FI.

Mesosoma: scutellum slightly longer than mesoscutum, SC:MSC = 1.19-1.31; dorsum of pronotum

punctate-reticulate, punctures coalesced in form of transverse rugae anteriorly, punctures along midline polygonal, not

distinctly circular (Fig. 26); midlobe of mesoscutum and entire scutellum with incomplete but regular cross-arcuate costae

(Fig. 14), less distinct along scutellar sulcus, sidelobe of mesoscutum smooth anteriorly along notauli (black areas),

laterally punctate-reticulate to rugose behind; apex of scutellum with distinctly or indistinctly septate marginal rim;

scutellum slightly acuminate, sides convergent at about 75 degrees; underside of scutellum almost smooth (Fig. 40);

median area of propodeum with deeply impressed foveae in form of distinct V-shape, submedian areas coriarious above

and smooth below with one or two transverse costae; postspiracular sclerite with very large fovea extended entire length of

upper postspiracular sclerite (Fig. 30); axilla rugose; axillula smooth and shining, costae not extended to posterior border

(Fig. 54).

Quaest. Ent., 1983, 19 ((1,2))

20

Darling

Metasoma: T5 with indistinct punctures.

Male: Length, 3. 7-4. 6 mm. Color as in female except, dark violet area can extend along entire midlobe of

mesoscutum and scutellum. Structure and sculpture as in female except: Head: Antennae: anellus relatively smaller,

0.15-0.23 length of FI; scape, in frontal view, expanded slightly apically, smooth, punctures indistinct and restricted to

inner surface, outer surface with strong setae, surface roughened (Fig. 49). Mesosoma: Propodeum more coarsely

sculptured with more prominent transverse costae. Genitalia (Fig. 75): digiti with three, four, or five large teeth and

two smaller teeth; ventral lobe broadly rounded; lateral demelanized areas of basiparamere reduced laterally, pigmented

median area shorter than digiti [n = 4].

Variation. — The Paraguayan specimen differs in a number of characteristics: sculpture of

the midlobe of the mesoscutum is not reduced along the scutellar suture, and the axillula is not

completely smooth but has a complete carina which parallels the ventral margin and defines a

narrow finger-like region. This sculpture is different from that of E. triangularis which does

not have a single carina but one to three oblique carinae (Fig. 53).

Euperilampus enigma n. sp.

(Figs. 50, 74)

Type Locality. — Bolivia, Santa Cruz, Robore.

Type Material. — Holotype (Male, USNM No. 100318). Bolivia, Santa Cruz, Robore (October 1959) [DCD

Slide No. 73] [specimen donated to the USNM by UK]. See discussion under E. brasiliensis complex.

Diagnosis. — The holotype differs from all E. brasiliensis complex males by having large,

recurved teeth on the digiti (Fig. 74; cf. Fig. 75, E. brasiliensis and Fig. 71, E. luteicrus). The

distinct punctures on the anterior surface of the antennal scape (Fig. 50; cf. Fig. 49, E.

brasiliensis), and the relatively large pronotum (PN:MSC = 0.58) further distinguish the male

of this species from those of E. brasiliensis [PN:MSC = 0.46-0.53]. There are fewer punctures

on the scape than in E. luteicrus (Figs. 45, 46).

Taxonomic note. — A female specimen from Bolivia is tentatively associated with E.

enigma: Female, Bolivia, Santa Cruz, Buena Vista (July 12 1971, Porter/Stange) [IML,

identified as Euperilampus triangularis (Say) in Fidalgo 1980: 194; DCD Slide Nos. 135, 136,

mouthparts, ovipositor]. This female is indistinguishable from E. brasiliensis and is not

described.

Derivation of specific epithet. — From the Greek for ‘something obscure, a riddle’; an

allusion to the uncertain systematic affiliation of the holotype.

Description. —

Male: Length, 4.2 mm. Color predominantly blue-green, with violet reflections. Vertex, midlobe and sidelobe of

mesoscutum with contrasting glossy black areas.

Head: Length of malar space 0.25 eye height; OOL = 0.87 POL; maximum width of scrobes 0.44 head width; margin

of scrobes, in frontal view, sinuous, slightly flared at level of antennal toruli; head widest across eyes; head transverse,

width:height = 1.17; inner orbital costae not convergent on posterior ocellus or extended around top of eye, ended abruptly

at ocular-ocellar region (as in E. brasiliensis. Fig. 22); outer orbital costae short, largely confined to malar region; vertex

almost smooth, posteriorly with transverse costae; clypeus transverse, width:height = 1.56, with indistinct arcuate costae

at extreme lateral margins, punctures indistinct, surface appearing smooth and polished; supraclypeal area 0.52 clypeus

height. Antennae: anellus 0.20 length of FI; scape, in frontal view, expanded slightly apically, length 4.53 times maximum

width, with indistinct punctures on anterior and inner surfaces, outer surface with strong setae, surface roughened (Fig.

50).

Mesosoma: PN:MSC = 0.58; scutellum longer than mesoscutum, SC:MSC = 1.27; dorsum of pronotum

punctate-reticulate, punctures coalesced in form of transverse rugae medially, punctures along midline polygonal, not

distinctly circular; midlobe of mesoscutum and entire scutellum with incomplete but regular cross-arcuate costae (as in E.

brasiliensis. Fig. 14), smooth along scutellar sulcus, sidelobe of mesoscutum with wide smooth area anteriorly along

notauli (black areas), laterally punctate-reticulate to rugose behind; apex of scutellum with indistinctly septate marginal

rim; scutellum slightly acuminate, sides convergent at about 75 degrees; underside of scutellum almost smooth (as in E.

brasiliensis. Fig. 40); median area of propodeum with deeply impressed foveae in form of distinct V-shape, submedian

areas distinctly coriarious with one or two transverse costae; postspiracular sclerite with very large fovea co-extensive with

entire upper postspiracular sclerite; axilla rugose; axillula smooth and shining, costae not extended to posterior border (as

The New World species of Euperilampus

21

in E. brasiliensis. Fig. 54).

Metasoma: T5 with indistinct punctures. Genitalia (Fig. 74): digiti with three very large and three smaller teeth,

teeth of right digitus recurved distally; ventral lobe broadly rounded; demelanized area of basiparamere large and

quadrate, not reduced laterally, pigmented median area less than one-half length of digiti.

Female: UNKNOWN

Euperilampus luteicrus n. sp.

(Figs. 45,46,71)

Type Locality. — Mexico, Jalisco, Guadalajara.

Type Material. — Holotype (Male, USNM No. 100319): Mexico, Jalisco, 15 mi. NE Guadalajara (Sept. 17

1970, GE/RM Bohart) [DCD Slide No. 170, genitalia] [specimen donated to USNM by USUj.

Diagnosis. — This holotype is the only New World specimen of Euperilampus with yellow

fore and mid tibiae, concolorous with the tarsi. I expect the yellow tibiae will be diagnostic for

the as yet unknown female. The antennal scape is covered with punctures (Figs. 45, 46), denser

than in E. enigma (Fig. 50), and E. brasiliensis (Fig. 49). The scutellum is relatively longer

than all other species in the E. brasiliensis complex, SCiMSC = 1.45, but the contrasting

black areas on the sidelobe of the mesoscutum suggest affinities with E. brasiliensis.

Geographical distribution. — The type locality is in the Central Plateau region of Mexico.

Extent of range is unknown.

Derivation of specific epithet. — From the Latin {luteus, ‘yellow’ and crus, ‘leg or shank’),

with reference to the yellow fore and mid tibiae.

Description. —

Male: Length, 3.75 mm. Color predominantly blue-green, with violet reflections. Vertex, midlobe and sidelobe of

mesoscutum with contrasting glossy black areas; fore and mid tibiae yellow, concolorous with tarsi.

Head: Length of malar space 0.25 eye height; OOL = POL; maximum width of scrobes 0.43 head width; margin of

scrobes, in frontal view, sinuous, slightly flared at level of antennal toruli; head widest across eyes; head transverse,

width:height = 1.19; inner orbital costae convergent on posterior ocellus, not extended around top of eye (as in E.

triangularis. Fig. 21): outer orbital costae short, largely confined to malar region; vertex almost smooth, posteriorly with

transverse costae; clypeus transverse, width:height = 1.60, with indistinct arcuate costae at extreme lateral margins,

punctures indistinct, surface appearing smooth and polished; supraclypeal area 0.58 clypeus height. Antennae: anellus 0.12

length of FI; scape, in frontal view, expanded slightly apically, length 4.10 times maximum width, punctures dense and

well developed on both anterior and inner surfaces, outer surface with strong setae, surface roughened (Figs. 45,46).

Mesosoma: PN:MSC = 0.57; scutellum longer than mesoscutum, SC:MSC = 1.45, relatively longer than other E.

brasiliensis complex species (range, 1.19-1.31); dorsum of pronotum punctate-reticulate, punctures coalesced in form of

transverse rugae anteriorly, punctures along midline distinctly circular; midlobe of mesoscutum and entire scutellum with

incomplete but regular cross-arcuate costae (as in E. brasiliensis. Fig. 14), smooth along scutellar sulcus, sidelobe of

mesoscutum with wide smooth area anteriorly along notauli (black areas), laterally punctate-reticulate to rugose behind;

apex of scutellum with distinctly septate marginal rim; scutellum slightly acuminate, sides convergent at about 75 degrees;

underside of scutellum almost smooth; median area of propodeum with deeply impressed fovea in form of distinct V-shape,

submedian areas smooth with few transverse costae and finer costulae; postspiracular sclerite with very large fovea

co-extensive with entire upper postspiracular sclerite; axilla rugose; axillula smooth and shining, costae not extended to

posterior border (as in E. brasiliensis. Fig. 54).

Metasoma: T5 with indistinct punctures. Genitalia (Fig. 71): digiti with three or four large teeth and two smaller

teeth; ventral lobe broadly rounded; demelanized lateral areas of basiparamere large and oval, not reduced laterally,

pigmented median area notched laterally and shorter than length of digiti.

Female: UNKNOWN

Euperilampus ameca n. sp.

Type Locality. — Mexico, Nayarit, Santa Isabel.

Type Material. — Holotype (Female, USNM No. 100320): Mexico, Nayarit, 9 mi. NW Santa Isabella [ =

Isabel] (March 10 1972, Parker/Miller).

Quaest. Ent., 1983, 19 ((1,2))

22

Darling

Diagnosis. — The holotype female differs from E. brasiliensis by having sidelobe of the

mesoscutum almost completely sculptured and with the contrasting black areas very small and

adjacent to the notauli. The short scutellum, SC:MSC = 1.30, establishes this species as a

member of the E. brasiliensis complex.

Geographical distribution. — The type locality is in the Sierra Madre Occidental in the

drainage of the Ameca River.

Derivation of the specific epithet. — A noun in apposition, from Ameca River.

Description. —

Female: Length, 5 mm. Color predominantly blue-green, with violet reflections. Vertex and midlobe of mesoscutum

with contrasting glossy black areas, sidelobe of mesoscutum with black areas restricted to narrow band along anterior

portion of notauli (much smaller than in other E. brasiliensis complex species).

Head: Length of malar space 0.29 eye height; OOL=0.94 POL; maximum width of scrobes 0.41 head width; margin

of scrobes, in frontal view, sinuous and flared at level of antennal toruli (as in E. triangularis. Fig. 11): head widest across

eyes; head transverse, width;height = 1.24; inner orbital costae not convergent on posterior ocellus or extended around top

of eye, ended abruptly at ocular-ocellar region (as in E. brasiliensis. Fig. 22); outer orbital costae short, largely confined to

malar region; vertex almost smooth, posteriorly with transverse costae; clypeus transverse, width:height = 1.59, with

indistinct arcuate costae at extreme lateral margins, punctures indistinct, surface appearing smooth and polished;

supraclypeal area 0.51 clypeus height. Antennae: anellus 0.25 length of FI; scape narrowly linear, length 4.53 times

maximum width.

Mesosoma: PN:MSC = 0.43; scutellum longer than mesoscutum, SC:MSC = 1.30; dorsum of pronotum

punctate-reticulate, punctures coalesced in form of transverse rugae medially, punctures along midline polygonal, not

distinctly circular; midlobe of mesoscutum and entire scutellum with incomplete but regular cross-arcuate costae, sidelobe

of mesoscutum without conspicuous smooth area anteriorly along notauli, this area roughened by indistinct rugae, laterally

punctate-reticulate to rugose behind; apex of scutellum with distinctly septate marginal rim; scutellum slightly acuminate,

sides convergent at about 75 degrees; underside of scutellum almost smooth (as in E. brasiliensis. Fig. 40); median area of

propodeum with deeply impressed foveae in form of distinct V-shape, submedian areas coriarious with single transverse

costa dorsally; postspiracular sclerite with very large fovea co-extensive with entire upper postspiracular sclerite; axilla

rugose; axillula smooth and shining, costae not extended to posterior border (as in E. brasiliensis. Fig. 54).

Metasoma: T5 with indistinct punctures.

Male: UNKNOWN

The New World species of Euperilampus

23

Table 1: Metric descriptors, for species of the Euperilampus triangularis species group, known

from multiple specimens (see discussion of E. triangularis species group). Measurements as

defined in ‘Methods and Terms’. Diagnostic characters indicated by asterisk (*). [ F, females;

M, males.]

Quaest. Ent., 1983, 19 ((1,2))

24

Darling

Figs. 1-3. Generic characters of Euperilampus. 1. Male genitalia E. triangularis, dorsal. 2. Male genitalia E. triangularis,

ventral [ Ad, adeagus; Bp, basiparamere; Dg, digitus; Ld, lateral demelanized area of basiparamere; Ls, lateral setae; Mp,

median pigmented area; VI, ventral lobe of basiparamere ]. 3. Labrum E. triangularis, dorsal view. Scale line 0.1 mm.

The New World species of Euperilampus

25

Figs. 4-7. Heads. 4. E. tanyglossa, female paratype; lateral. Inset; apex of labio-maxillary complex, anterior. 5. E.

tanyglossa, female paratype; frontal. 6. E. aureicornis, female holotype; frontal. 7. E. magnus, female holotype; frontal.

Scale line 1 mm.

Quaest. Ent., 1983, 19 ((1,2))

26

Darling

Figs. 8-16. 8-1 1. Heads. 8. E. krombeini; lateral. 9. E. triangularis; lateral. 10. E. iodes\ frontal. 1 1. £. triangularis;

frontal. 12-16. Mesosomata, dorsal. 12. E. krombeini. 13. E. iodes. 14. E. brasiliensis. 15. E. solox. 16. E. triangularis

[AL, axilla; AX, axillula; ML, midlobe of mesoscutum; MR, marginal rim of scutellum; MSC, mesoscutum; NO, notaulix;

PN, pronotum; SC, scutellum; SL, sidelobe of mesoscutum].

The New World species of Euperilampus

27

mmrn

Figs. 17-24. 17-20. Propodea. 17. E. krombeini. 18. E. tanyglossa. 19. E. triangularis, Florida; length = 5.4 mm. 20. E.

triangularis reared from Hyposoter, Arkansas, length = 2.9 mm. 21-24. Heads, dorsal. 21. E. triangularis. 22. E.

brasiliensis. 23. E. iodes. 24. E. iodes\ higher magnification, illustrating costae extending through ocular-ocellar region

along eye margin.

Quaest. Ent., 1983, 19 ((1,2))

28

Darling

Figs. 25-32. 25.-26. Sculpture types. 25. Punctate-reticulate, pronotum of E. tanyglossa. 26. Punctate-reticulate,

punctures coalesced to form irregular rugae, pronotum of E. brasiliensis. 27-29. Second metasomal tergites. 27. E.

triangularis. 28. E. tanyglossa. 29. E. krombeini. 30-32. Mesosomata, lateral. 30. E. brasiliensis. 31. £. iodes. 32. E.

solox [P, postspiracular sclerite].

The New World species of Euperilampus

29

Figs. 33-40. 33-34. Mesosoma, E. tanyglossa. 33. Pronotum and mesoscutum, dorsal. 34. Scutellum, dorsal. 35-37.

Mesosomata, lateral. 35. E. tanyglossa. 36. E. krombeini. 31. E. triangularis', 38. Head, frontal; E. krombeini. 39-40.

Scutellum, underside. 39. E. triangularis. 40. E. brasiliensis [AX, axillula; MR, marginal rim of scutellum; P,

postspiracular sclerite].

Quaest. Ent., 1983, 19 ((1,2))

30

Darling

Figs. 41-52. Male antennal scapes. A\. E. tanyglossa, outer surface, 280x. 42-52. Anterior surfaces. 42. E. tanyglossa,

233x. 43. E. krombeini, 263x. 44. E. triangularis, 202x. 45. E. luteicrus, 233x. 46. E. luteicrus, l,190x. 47. E. solox, 233x.

48. E. solox, l,190x. 49. E. brasiliensis, 210x. 50. E. enigma, 233x. 51. E. iodes, 233x. 52. E. iodes, l,190x.

The New World species of Euperilampus

31

Figs. 53-59. 53 - 54. [AL, axilla; AX, axillula]. 53. E. triangularis. 54. E. brasiliensis. 55-56. Labio-maxillary complexes

[C, cardo, S, stipes and G, galea or galea/lacinia of maxilla; PR, prementum; L, ligula of labium]. 55. E. triangularis. 56.

E. tanyglossa. 57 -59. Forewings. 57. E. triangularis. 58. E. tanyglossa. 59. E. krombeini [M, marginal, PM,

postmarginal, S, stigmal and SM, submarginal veins].

Quaest. Ent., 1983, 19 ((1,2))

Darling

Figs. 60-66. Character states in Perilampus. 60-61. Mesosomata, lateral. 60. P. chysopae. 61. P. platygaster [sic auct.].

62. Forewing, P. hyalinus. 63. Propodeum, P. platygaster. 64. Male antennal scape, anterior surface, P. hyalinus. 65.

Head, lateral, P. hyalinus. 66. Mesosoma, dorsal, P. hyalinus [FC, frontal carina; MS, malar sulcus; P, postspiracular

sclerite; R, median ridge of propodeum; S, submedian area of propodium].

The New World species of Euperilampus

33

Figs. 67-76. Male genitalic structures of Euperilampus. 67-69. Subgenital plates. 67. E. triangularis. 68. E. krombeini.

69. E. tanyglossa. 70-76. Digit! and apex of basiparamere, ventral view (compare with Fig. 2). The demelanized areas of

the basiparamere are outlined and compared with the stippled areas in Fig. 2. The lateral setae (Ls) are not drawn. Scale

line 0.1 mm. 70. E. tanyglossa. 1\. E. luteicrus. 12. E. iodes. 73. E. krombeini. 74. E. enigma. 75. E. brasiliensis. 76. E.

solox.

Qmest.Ent.. 1983, 19 ((1,2))

34

Darling

PHYLOGENETIC RELATIONSHIPS

Hypotheses of phylogeny can be developed following the methods of Hennig (1966).

Reference to an outgroup, ideally the sister group of the taxon under consideration, allows

character states to be hypothesized as either plesiomorphic (ancestral) or apomorphic

(derived). The degree of relationship, defined as the relative recency of common ancestry, can

then be assessed based on shared derived characters (synapomorphies).

Character states and polarities are summarized in Table 2. The basis for the polarity

decisions is outlined for the individual characters [ ] in the following discussion.

Intergeneric relationships within the Perilampidae are unresolved. All recognized genera

show affinities with the speciose cosmopolitan genus Perilampus (Boucek 1978). It is quite

possible that various species groups within Perilampus share a more recent common ancestor

with other recognized genera than with other species of Perilampus.

A natural classification of the Perilampidae, i.e., one recognizing hypothesized phylogenetic

relationships, must be based on recognition of monophyletic species groups within Perilampus.

Smulyan (1936), in a revision of the nearctic species of Perilampus, recognized the ‘carinate

species group’, based on possession of a distinct frontal carina (Fig. 65) and finger-like axillulae

(Fig. 61). Both of these characters are here regarded as apomorphic in the Perilampidae. The

frontal carina is absent from Steffanolampus salicetum (Steffan), and from species related to

Perilampus micans Dalman [ groups I regard as early derivatives of Perilampus on the basis of

the structure of the postspiracular sclerite and labrum] . The frontal carina is also absent from

widely divergent species of Perilampus. However, in addition to the New World species of

Perilampus, the frontal carina is also developed in Indo-Pacific species related to P.

punctiventris Crawford (e.g., P. singaporensis Rohwer, P. nesiotes Crawford), and in

Krombeinius and Euperilampus. The frontal carina [1] is here regarded as a synapomorphy

uniting these groups.

Albeit inappropriately named, Smulyan’s ‘carinate group’ is regarded as monophyletic on

the basis of the finger-like axillulae; this character is only found in the New World carinate

species and is therefore regarded as apomorphic. This group is perhaps better referred to as the

"Perilampus hyalinus group’, based on the widespread New World species.

The Perilampus hyalinus group shares an apomorphic character with species of

Krombeinius and Euperilampus: the malar sulcus is obliterated by oblique striation (costae)

[2], as noted by Boucek (1978:302).

Krombeinius and Euperilampus are united on the basis of the following synapomorphies:

postspiracular sclerite reduced to a narrow triangle [3]; pronotum massive dorsally [12]; inner

orbits with well developed raised costae or rugae [13] (characters noted by Bou5ek 1978); and

the propodeum lacking a distinct median ridge [14]. The plesiomorpic states are found in all

species of Perilampus-, however, the raised costae on the inner orbits are slightly developed in

some species of the Perilampus hyalinus complex.

Euperilampus is defined on the basis of the following apomorphies: male genitalia with

reduced parameres [8]; labrum 8-digitate [11]; marginal vein shorter than postmarginal [5];

scutellum with a distinct marginal rim [9]; and metasomal T3 transverse, shorter than T2 [10].

Autapomorphies have yet to be documented for Krombeinius. The strikingly modified labrum

(see under discussion of genus Euperilampus, examined in a single specimen) and the host

association (primary parasites of solitary wasps, known only for the type species, K.

eumenidarum’, host: Paraleptomenes mephitis (Cameron)) are possibly autapomorphies.

The New World species of Euperilampus

35

The preceding hypotheses of relationships allow polarity determinations for character states

found in Euperilampus. A character state is regarded as plesiomorphic in Euperilampus if it is

present in Krombeinius. If both character states are present in Krombeinius then the state of

the character in the P. hyalinus group is considered as plesiomorphic. A cladogram can then be

constructed to show the distribution of derived character states.

I have extended the cladistic analysis only to species groups of Euperilampus. The New

World species groups are as outlined in the ‘Synopsis’, and three terminal taxa are used in the

analysis: E. krombeini, E. tanyglossa group and E. triangularis group. No attempt was made

to present a cladogram of E. triangularis group species. The diagnostic characters are mainly

sculpture and color differences. I expect considerable homoplasy in these characters

(convergences and reversals), and hence the resultant cladogram would be suspect. Old World

species are represented in the analysis by E. mediterraneus Boucek, and E. scutellatus

(Girault). I have examined a long series of specimens of E. scutellatus [USNM]. E.

mediterraneus is well illustrated and described (Boucek 1972), and character states can be

determined. E. hymenopterae (Risbec), E. beharae (Risbec), E. sinensis Bou5ek and E. spina

Boucek are not included in the analysis; for the characters listed in Table 2 these species appear

to be identical with E. scutellatus.

E. tanyglossa group {E. tanyglossa, E. aureicornis) (both from Mexico) and E.

mediterraneus (southern Bulgaria) constitute a monophyletic group on the basis of the

following synapomorphies: labio-maxillary complex elongate [20]; and stigmal vein longer than

marginal vein [7]. The labio-maxillary complex is elongated in a similar way in E.

mediterraneus (Boucek 1972, Fig. 2) and in the E. tanyglossa group: the galea/lacinia of

maxilla and prementum, and ligula of the labium are greatly lengthened. The maxillary palps

are also elongate in all three species. These similarities in detail suggest that this elaboration is

homologous, and occurred only once, in the common ancestor of the E. tanyglossa group and E.

mediterraneus. In E. krombeini, the stigmal vein is subequal in length to the marginal vein [6]

(Fig. 59). This character state is phenotypically intermediate between the plesiomorphic and

apomorphic states of character 7. I do not regard this as a linear transformation series or

morphocline (stigmal vein shorter than marginal, stigmal vein equal to marginal, stigmal vein

longer than marginal vein) on the basis of other characters. Seven synapomorphies indicate that

E. krombeini and E. mediterraneus + tanyglossa group are not sister groups [4,15,17,22,18].

Only two characters support a sister group relationship between these taxa, and these are color

characters [19,21], and are discussed later as convergences.

The long-tongued species of Euperilampus are united with E. scutellatus by the following

characters: notauli indistinctly indicated [18]; and the mesoscutum uniformly sculptured,

without a more weakly sculptured area along the notauli [22]. The alternative character states

(see Table 2) are found in Krombeinius and Perilampus and therefore considered

plesiomorphic for Euperilampus.

E.. triangularis group and E. krombeini are united by the presence of cross-arcuate rugae or

costae on the mesoscutum [17]; the deep median foveae on the propodeum [15]; and the

postspiracular sclerite abruptly narrowed ventrally [4].

Figure 77 is the most parsimonious cladogram based on the characters in Table 2. This

analysis is not free of homoplasy. Metallic colors [19] and infuscate wings [21] appear to have

arisen convergently in the two lineages of Euperilampus. It is interesting to note that all New

World species have metallic colors and all Old World species are dark black. There is also a

reversal indicated by this analysis. The raised costae on the inner orbits [13] are reduced in E.

Quaest. Ent., 1983, 19 ((1,2))

36

Darling

Table 2: Polarity of character states in Euperilampus and related genera, based on outgroup

comparisons (see text for discussion).

No.Attribute Plesiomorphic State Apomorphic State

(continued on next page)

The New World species of Euperilampus

37

Table 2 (continued)

krombeini.

This preliminary analysis documents the monophyly of E. tanyglossa + aureicornis +

mediterraneus; this relationship cannot be retrieved from a classification recognizing the

subgenera Euperilampus (New World) and Euperilampoides (Old World). Further

phylogenetic studies wilt be required for the placement of the remaining Old World species and

should serve to test the present cladogram.

If the above hypothesized relationships of the Perilampus hyalinus group, Krombeinius and

Euperilampus are corroborated by the placement of additional species, by additional

characters, and the elucidatioon of monophyletic species groups within Perilampus, then the

recognition of these two genera would render Perilampus paraphyletic. The genus would not

include all species descended from a common ancestor. Species would be placed in Perilampus

if they belonged to the Perilampidae but lacked the autapomorphies of the other genera. In the

final analysis it may be prudent to retain Krombeinius and Euperilampus and to describe new

genera, where needed, for monophyletic species groups of Perilampus. Clearly, the first priority

for the development of a natural classification of the Perilampidae is the definition of

monophyletic species groups within the core genus, Perilampus.

ACKNOWLEDGEMENTS

I thank the following for comments on the manuscript: Z. Boucek, W. L. Brown, Jr., L.

Burnham, J. M. Carpenter, G. C. Eickwort, G. A. P. Gibson, E. E, Grissell, N. F. Johnson, S.

Nichols, M. J. Sharkey, M. Schauff, Q. D. Wheeler, and C. M. Yoshimoto.

The frontispiece was skillfully prepared by Jim Miller. Figures 10, 13, 15, 23, 24 and 31

were prepared from scanning electron micrographs of uncoated specimens; I thank H. F.

Howden and L. E. C. Ling of Carleton University, Ottawa for these photographs.

This research was supported in part by a National Science Foundation Dissertation

Improvement Grant. The Griswold Fund Committee, Department of Entomology, Cornell

University assisted with the publication costs.

Special thanks are extended to Dr. J. G. Franclemont, for numerous discussions of all

aspects of the project.

Quaest. Ent., 1983, 19 ((1,2))

38

Darling

Quaest. Ent., 1983, 19 ((1,2))

Fig. 77. Cladogram showing the distribution of shared derived characters (synapomorphies)jn genera of Perilampidae and

in species and species groups of Euperilampus (see text for discussion of species groups), x - synapomorphy, x

^ ^ ^ , • 1 . in Tjihlp 7

The New World species of Euperilampus

39

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Fox, D.L. 1979. Biochromy. University of California Press, Berkeley. 248 pp.

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Parker, H.-L. 1924. Recherches sur les formes postembryonnaires des chalcidiens. Annales de

la Societe Entomologique de France, 93: 261-379.

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and H.K. Townes. Flymenoptera of America North of Mexico. Synoptic Catalog.

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Entomological Society of London, London. 100 pp.

Riek, E. F. 1966. Australian Hymenoptera Chalcidoidea, Family Pteromalidae, Subfamily

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of the Maclurian Lyceum of the Arts and Sciences, 1: 67-83.

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in America north of Mexico. Proceedings of the United States National Museum, 83:

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of Arkansas Agricultural Experiment Station Bulletin, 759. 106 pp.

FILTER FEEDING OF SIMULIUM FULVINOTUM (DIPTERA: SIMULIIDAE) IN THE

CENTRAL AMAZON BASIN

Lawrence A. Lacey'

Insects Affecting Man and Animals Research Laboratory

U. S. D. A., A.R.S.. P.O. Box 14565

Gainesville, FL 32604

U.S.A.

Quaestiones Entomologicae

19:41-51 1983

Joyce M. Lacey

California Department of Water Resources

Northern District

2440 Main Street, P.O. Box 607

Red Bluff CA 96080

ABSTRACT

Filter feeding of larval Simulium fulvinotum Cerqueira and Mello was studied near

Manaus, Brazil, using particulate fluorescent dye in small black water streams. Transit time

of particles through the midgut was 29.2 min in one stream with a current velocity of ca. I

mjsec and 0.37 mg/l seston, but was only 26.1 min in another stream with 1. 5-2.0 mjsec

velocity and 2.01 mg/l seston. Penultimate instars ofS. fulvinotum held the cephalic fans open

for an average of 3.3 min before they were closed for removal of particles. Food of larvae

included a variety of algae, detritus, bacteria and insect parts. Methods for obtaining

adequate nutrition in the habitats of S. fulvinotum are: less frequent cleaning of fans, an

efficient filtering mechanism, and location of larvae in swift current.

RESUMO

A alimentaQao atraves de filtracao das larvas de Simulium fulvinotum Cerqueira e Mello foi estudada perto de

Manaus, Brasil, utilizando uma tinta fluorescente em peguenos igarapes de dgua preta. O tempo de passagem atraves do

intestino medio foi 29,2 min. em uma igarape com correnteza de I mjseg e 0,37 mgj\ de particulas em suspensao. Em

outro igarape com correnteza de 1, 5-2,0 mjseg e 2,01 mgj\ de particulas em suspensao, o tempo de passagem foi apenas

26,1 min. Individuos da penUltima etapa de S. fulvinotum mantiveram os filtros cefdlicos abertos durante uma media de

3,3 min. antes de retrai-los para remover as particulas. A alimentacdo larval incluiu uma variedade de algas, detritos,

bacterias e pedacos de insetos. As estrategias para obter nutricao adequada nos habitats de S. fulvinotum incluem

movimentos mais lentos de alimentacdo, um mechanismo eficiente de filtracao, e localizacdo das larvas em correnteza

rdpida.

Most biological research on the black flies of the Amazon Basin of Brazil has been on the

adults of vector species with very little attention paid to the biology and ecology of larvae. One

species which has been studied with respect to larval ecology and life history is Simulium

‘Formerly with the Institute Nacional de Pesquisas da Amazonia, Divisao de Ciencias Medicas,

Manaus, Brazil

42

Lacey and Lacey

Figure 1. Typical larval habitat of Simulium fulvinotum, Igarape Acara Reserve Ducke, near Manaus, Brazil.

Filter feeding of Simulium fulvinotum

43

fulvinotum Cerqueira and Mello (1968); habitats were characaterized by Dellome (1978);

number of instars was determined by Gorayeb (1981); and predators of larvae were identified

by Gorayeb and Finger (1978).

Several studies have been conducted on the dynamics of filter feeding of other black fly

species. Until now this aspect of the larval ecology of S. fulvinotum has not been investigated.

In addition to providing basic information on the use of nutrients by simuliids in black water

streams, information on larval feeding could also be useful when planning control of vector

simuliid species in this habitat. Kershaw et al. (1968) and Helson and West (1978)

demonstrated that particulate insecticides were active against target black flies with minimal

effect on other aquatic organisms which did not filter feed. Feeding rate and retention time also

will have a major influence on the efficacy of perorally active microbial control agents, such as

Bacillus thuringiensis var. israelensis (Gaugler and Molloy 1980).

This paper presents information about filter feeding activity of larvae of S. fulvinotum in

small streams near Manaus, Brazil.

THE LARVAL HABITAT

S. fulvinotum larvae characteristically inhabit very swift current in small, shallow, black

water streams (as defined by Sioli 1964; tannin-coloured, nutrient-poor water, draining

podsols). The lip of the water drop in Figure 1 typifies the usual larval attachment site. In our

study, habitats investigated were in primary and secondary forest 20-25 km northeast of

Manaus, Brazil. In addition to typical larval habitats where current velocities of 1. 5-2.0 m/sec

(ca. 1 m^/sec discharge) were recorded, larvae were also found in one atypical stream (I) in

current as low as 1 m/sec (.5 m^/sec discharge). Most streams where S. fulvinotum was found

were heavily shaded. Stream I, however, was exposed to full sun because of recent

deforestation.

Temperatures of 24.3-25.8°C and pH of 5.4-5. 7 have been recorded for S. fulvinotum

larval habitats (Cerqueira and Mello 1968, Dellome 1978). Water temperatures were

24.5-25.7°C during the course of our investigation.

METHODS AND MATERIALS

Transit time of particles passing through the larval midgut was determined using a

fluorescent dye (Hercules® Radiant Fluorescent Dye; orange WD 16).‘ Approximately 10 g of

dye were suspended in a liter of water by vigorous shaking, then evenly poured into the river

upstream of the larvae for 5-10 sec. After 10, 20, 30, and 40 min, leaves with attached larvae

were carefully removed from the stream, so that larvae on adjacent leaves were not disturbed,

and then stored in 70% ethanol. Ultimate larval instars were dissected to determine position of

the dye plug in the midgut. Measurements of position of the dye and length of midgut were

made with an ocular micrometer in a Zeiss dissecting microscope.

Gut filling times were recorded in Igarape da Pedreira (Stream II) and Stream I. Both

streams are located in the Cacao Research Plantation (CEPLAC)) 25 km northeast of

’Mention of a commercial or proprietary product does not constitute an endorsement by the

U.S.D.A.

Quaest. Ent., 1983, 19 (1,2)

44

Lacey and Lacey

Manaus. Temperature in each was measured with a mercury thermometer. Current velocity at

the surface was determined using a float and stop watch. Suspended particles were measured

by taking 3 one-litre samples of water from each site and individually vacuum filtering each

through previously weighed millipore cellulose filters (0.45 um pores, 47 mm diameter filter at

< 40 cm Hg suction). The filters were then dried for 15 hrs at 65°C. After cooling and

equilibrating for 6 hrs in the same conditions under which the original filter weights were

taken, they were weighed on a Mettler H 34 balance ( ± 0.1 mg accuracy).

Gut contents of late instars collected from Stream I on two separate occasions were

analyzed. Larvae were placed on ice soon after collecting until they were dissected in the

laboratory later in the same day. The dissected food columns were teased apart in distilled

water and observed with the aid of a light microscope.

Feeding behaviour was observed in Stream I by viewing the larvae through the bottom of a

600 ml glass beaker which was lowered into the stream within a few centimetres of the attached

larvae.

Data about transit time of particles through the midgut were analyzed with regression

analysis (least squares method). The 20 min samples for the two streams were compared using

Student’s t-test.

Primary cephalic fans of ultimate instar S. fulvinotum (preserved in 70% ethanol) were

prepared for scanning electron microscopy by: dehydrating in ethanol and freon; critical point

drying in freon; and gold coating. Scanning electron micrographs were made with a Hitachi

H-600 electron microscope.

RESULTS

Mean quantities of suspended particles in Streams I and II were 0.37 and 2.01 mg/liter,

respectively. Stream velocities were ca 1.0 and 1. 5-2.0 m/sec respectively. Temperature in both

streams was 25.2°C.

Table 1. Posterior displacement of dye plug in the midgut of Simulium fulvinotum in Stream I

(suspended particles 0.37 mg/1; velocity 1.02 m/sec).

Min. after exposure

No. larvae

Mean % displacement ± S.E.*

38.18 ± 2.01

64.38 ± 2.73

103.0*± 1.75

'Calculated by position of dye in mid- and hindgut.

Filter feeding of Simulium fulvinotum

45

Table 2. Posterior displacement of dye plug in the midgut of S. fulvinotum in Stream

II (suspended particles 2.01 mg/1; velocity 1. 5-2.0 m/sec).

Data used to determine time for particles to pass through midguts of S. fulvinotum larvae

from Streams I and II are presented in Tables 1 and 2. Predicted transit times for the two

streams were 29.2 and 26.1 min, respectively. Displacement of dye plugs 20 min after initial

exposure in the two populations was significantly different (p < 0.05). The mean for the 30

min sample from Stream I was calculated from the position of dye in both the mid- and

hindguts of 11 larvae. The low variance in position of dye plugs observed in the 10 and 20 min

samples in each of the streams indicates rather uniform feeding rates at each location.

Larvae observed feeding in situ held the cephalic fans open for an average of 3.3 ± .3 min

(range: 1.8-4. 8; n = 10). Fans were held closed for between 2 and 25 sec for removal of

particles.

Analysis of gut contents revealed a high percentage of algae (more than 50%) and detritus.

The various algal taxa represented were: Chlorophyta {Oedogonium sp., Ankistrodesmus sp.,

Cosmarium sp. and an unidentified filamentous species); Chrysophyta {Melosira spp.,

Tabellaria spp., Fragilaria sp., Nitzschia sp., and several genera of unidentified pennate

diatoms); and Cyanophyta {Chorococcus sp., Oscillatoria sp., Spirulina sp., and an

unidentified filamentous species). In addition to living diatoms, many empty diatom frustules

were found. Also present were unidentified bacteria, insect parts and sand.

DISCUSSION

The dynamics of filter feeding by black fly larvae have been elucidated by a number of

investigators and summarized by Wallace and Merritt (1980). Particle transit times of from 20

min to more than 24 hr have been recorded (Davies and Syme 1958, Ladle, Bass and Jenkins

1972, Mulla and Lacey 1976, Elouard and Elsen 1977, Wotton 1978, Schroder 1980b).

Differences in filtering activity and transit time of particles through guts of filter-feeding

species may be due to species, instar, temperature, stream velocity, parasitism, imminent

pupation, and amount and dimensions of available seston (Mulla and Lacey 1976, Chance

1977, Elouard and Elsen 1977, Moore 1977b, Elsen, Quillevere and Hebrard 1978, Wotton

1978, Elsen and Hebrard 1979, Lacey and Mulla 1979, Elsen 1980, Schroder 1980 a, b).

Increased stream velocity and/or additional amount of suspended matter in Stream II was

responsible for the accelerated feeding rate of S. fulvinotum over that observed in Stream I. An

even greater difference in feeding rate might be expected based on disparity of seston

concentration and stream velocity between Streams I and II. Lack of exaggerated differences

might be explained as a funetion of the inherent maximum filtering efficiency of S. fulvinotum.

Kurtak (1978) observed a decrease in filtering efficiency (i.e., the portion of particles ingested

Quaest. Ent., 1983, 19 (1,2)

46

Lacey and Lacey

per larva per second relative to the total number of particles offered per fan area per second)

when concentration of particles and stream velocity increased. Under laboratory conditions,

Lacey and Mulla (1979) and Schroder (1980b) reported plateaus of optimal particle

concentration and stream velocity for maximum ingestion rates. Beyond the optimum range, an

increase in current velocity and particle concentration may result in feeding inhibition (Lacey

and Mulla 1979, Gaugler and Molloy 1980).

A few ultimate instars with dark histoblasts (pharate pupae) were observed without dye

plugs and with partially or completely empty midguts. Feeding had apparently ceased prior to

pupation, and the gut was emptied by peristaltic action. Chance (1977) reported that

Simulium vittatum Zetterstedt larvae may refrain from feeding for periods of 90 min or longer.

In our study, the only individuals which failed to filter dye were pharate pupae.

Choice of larval habitat appears to be related to specific feeding methods for several black

fly species (Carlsson et al. 1977, Kovachev 1979, Wotton 1979, 1980b) or at least choice of a

specific habitat markedly influences type of nutrient the larvae will encounter (Maciolek and

Tunzi 1968, Kurtak 1979). Selection of a habitat with an optimally high stream velocity would

appear to maximize feeding rates where seston levels are as low as those found in the black

water streams inhabited by S. fulvinotum larvae. In streams with waterfalls or other zones of

very swift current, S. fulvinotum larvae are invariably found in the fastest current. Initial

selection of these sites is apparently made by the ovipositing female (Gorayeb 1981).

Other feeding methods that would enhance ingestion would be more efficient use of the

filtering mechanism and behaviour. Simuliid larvae are capable of filtering colloidal sized

particles (Wotton 1976). Capture of these and other particles smaller than the spaces between

the microtrichia of the cephalic fan rays is apparently aided by a mucosubstance which coats

the cephalic fans (Ross and Craig 1980). A single median ray of the primary cephalic fan of S.

fulvinotum is shown in Fig. 2. An enlargement of the middle portion of the ray (Fig. 3) shows

microtrichia used for capture of fine particles. Mictrotrichia of S. fulvinotum are considerably

longer than those of S. vittatum (D. A. Craig, personal communication), a species found in

streams with moderate to high seston concentrations. Accelerated feeding rate of S. fulvinotum

is at least partially influenced by this morphological adaptation to low seston loads. Increased

surface area of the filtration mechanism of S. fulvinotum probably facilitates capture of a

greater number of particles per unit of time.

Time spent by S. fulvinotum larvae with fans open is considerably greater than that

reported by Kurtak (1973) and Craig and Chance (1982) for S. vittatum. Ostensiblly S.

fulvinotum maximizes contact of its cephalic fans with the current in compensation for a low

particulate load. Craig and Chance (1982) hypothesize that larvae with less frequent

mouthpart movements may filter more efficiently than those which clean their fans more often

because the latter have their fans adducted (not exposed to food) for a significantly greater

period of time. Rapid flicking of fans reported by other investigators was only occasionally seen

during our observations. Observations were only made in Stream I because of disturbance of

larvae in Stream II when they were observed from upstream. Observations from downstream

were not possible because of a 2.5 m waterfall.

Gut contents of filter-feeding black fly larvae generally reflect relative abundance of

nutritional materials within a particle size range that they can filter in the stream where they

are located (Maciolek and Tunzi 1968, Moore 1977a, b, Wotton 1977, Kurtak 1979, Wallace

and Merritt 1980). A wide variety of food types has been recorded for simuliid larvae, ranging

from animal matter (Serra-Tosio 1967, Burton 1971, Disney 1971) to algae (Anderson and

Filter feeding of Simulium fulvinotum

A1

Figure 2. Middle ray of primary cephalic fan of Simulium fulvinotum larva.

Figure 3. Median portion of middle ray in Figure 2. Scale = 20 /um.

Quaest. Ent., 1983, 19 (1,2)

48

Lacey and Lacey

Dicke 1960, Maitland and Penney 1967, Burton 1973, Pavlichenko et aL 1977), bacteria

(Fredeen 1964, Pavlichenko et al. 1977, Wotton 1980a) and detritus (Cummins and Klug

1979). Considering the low seston load, it was interesting to find algae making up the major

portion of the diet of S. fulvinotum larvae. Algae in streams are characteristically benthic, so

their contribution to the seston is generally considered to be accidental (Whitton 1975).

Scraping of the substrate by S. fulvinotum larvae may account for the high proportion of algae

in the gut. It is possible that this activity was not observed because of the low number of

observations that were made in situ. Mokry (1975) reported that in an “average” hour,

Simulium venustum larvae scraped for 20 min, filtered for 20 min, and rested for 20 min. Some

investigators have questioned nutritional value of living algae and recently dead plant matter to

larvae which have a relatively brief retention time. Kurtak (1979) and McCullough et al.

(1979), however, reported that more than 50% of ingested diatoms were digested. Similar

findings were reported by Maciolek and Tunzi (1968). In our research, the large number of

empty frustules may be an indication of digestion.

Although bacteria did not appear to be a major source of food in our samples, they may be a

necessary component for detritus use through biochemical alterations of the detrital substrate

(Cummins and Klug 1979). Alternatively, Anderson and Cummins (1979) suggest that since

retention time of food in the gut is so brief in larval simuliids, bacteria stripped from the

surface of refractory detritus particles probably contribute most of the nutritional value. Lotic

food sources of simuliids and their relationship with microbes are summarized by Cummins and

Klug (1979).

Abundant detritus in the form of fine particulate organic matter has been associated with

maintenance of dense populations of simuliid larvae (Carlsson et al. 1977, and other authors

cited by Anderson and Seddell 1979). Detritus may have varying degrees of importance in the

diet of S. fulvinotum larvae depending on season, location and deforestation activity. As was

previously stated, most sites for S. fulvinotum were found under dense canopy. The site where

larvae were collected for analysis of gut content was exposed to sunlight and was undoubtedly

more conducive to a higher proportion of algae in the water and on the substratum. Kurtak

(1979) reported variability in percentage of various food types in streams in which he worked

both as a function of season and location.

The ecological role of black fly larvae, at times the most abundant aquatic insect in small

streams of the Central Amazon, requires further elucidation. Areas for future research could

include production studies on the larvae of S. fulvinotum in a wider variety of locations and

seasons as well as comparison with other species of simuliids in the same general habitat.

Research on the range of particle sizes ingested would be a useful first step in studying the

effects of particlate insecticides on black fly larvae and nontarget organisms in the habitat of S.

fulvinotum.

Due to its ubiquity and accessibility, S. fulvinotum will provide an excellent model for the

study of nutrient utilization and cycling and insecticide use in the nutrient-poor black water

environment.

ACKNOWLEDGEMENTS

The helpful comments and advice of Roger Wotton, Goldsmith’s College, University of

London, Douglas A. Craig, University of Alberta, and Albert Undeen, U.S.D.A., during the

preparation of the manuscript are very much appreciated. We are also grateful to D. A. Craig

Filter feeding of Simulium fulvinotum

49

for making the scanning electron micrographs.

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Simulium fulvinotum sp. n. (Diptera, Nematocera). Amazoniana 1: 205-210.

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temperature de I’eau, de la concentration de particules et de la nature de ces

particules. Ibid 59: 49-58.

Elsen, P., D. Quillevere, and G. Hebrard. 1978. Le transit intestinal chez les larves du

Quaest. Ent., 1983, 19 (1,2)

50

Lacey and Lacey

complexe Simulium damnosum (Diptera, Simuliidae) en Afrique de I’Ouest. 1.

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ser. Ent. Med. et Parasitol 15: 29-39.

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Gorayeb, I.S. and R.R. Pinger. 1978. Deteccao de predadores naturals das larvas de Simulium

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1968. The selective control of Simulium larvae by paraticulate insecticides and its

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Canadian Journal of Zoology 56: 1608-1623.

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60: 585-596.

Morky, J. E. 1975. Studies on the ecology and biology of blackfly larvae utilizing an in situ

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Filter feeding of Simulium fulvinotum

51

Moore, J. W. 1977a. Relative availability and utilization of algae in two subarctic rivers.

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Vereinigung fiir Limnologie 15: 1053-1058.

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Review of Entomology 25: 103-132.

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California Press.

Wotton, R. S. 1976. Evidence that blackfly larvae can feed on particles of colloidal size. Nature

(London) 261:697.

Wotton, R. S. 1977. The size of particles ingested by moorland stream blackfly larvae

(Simuliidae.) Oikos 29: 332-335.

Wotton, R. S. 1978. The feeding-rate of Metacnephia tredecimatum larvae (Diptera:

Simuliidae) in a Swedish lake outlet. Ibid. 32: 121-125.

Wotton, R. S. 1979. The influence of a lake on the distribution of black-fly species (Diptera:

Simuliidae) along a river. Ibid. 32: 368-372.

Wotton, R. S. 1980a. Bacteria as food for blackfly larvae (Diptera: Simuliidae) in a lake-outlet

in Finland. Ann. Zool. Fennici 17: 127-130.

Wotton, R. S. 1980b. Coprophagy as an economic feeding tactic in blackfly larvae. Oikos 34:

282-286.

Quaestiones Entomologicae 1983, 19 (1,2)

TRENDS IN NUMBERS OF AQUATIC INVERTEBRATES IN A LARGE CANADIAN

RIVER' DURING FOUR YEARS OF BLACK FLY LARVICIDING WITH

METHOXYCHLOR (DIPTERA: SIMULIIDAE)

F. J. H. Fredeen

Agriculture Canada Research Station

107 Science Crescent

Saskatoon, Saskatchewan Quaestiones Entomologicae

S7N0X2 19:53-92 1983

ABSTRACT

Methoxychlor was injected at about 0.3 parts a.i per million parts of water, maintained for

15 minutes, into the Saskatchewan River System in Saskatchewan once in 1976, six times in

1977, seven in 1978, 19 in 1979, and five times in 1980. Severe outbreaks o/Simulium lugged

originated from various portions of the river up to 200 km long during the first three years,

but not in 1979 or 1980.

With suprageneric taxa serving as units, trends in numbers of non-simuliids were measured

1977 through 1980. Average densities of combined non-simuliid invertebrate populations

attaching weekly to artificial substrates in mid-river sites in all three branches of the

Saskatchewan River peaked in 1979 (P<0.01) the year of maximum larvicide use, but in 1980

returned to just above the 1977 level. Average numbers of invertebrates in benthic samples

from river margins also generally peaked in 1979 or 1980 in all three river branches. In one or

more of the six locations sampled, however, numbers of certain families of Ephemeroptera

(baetids, heptageniids, caenids, leptophlebiids, and polymitarcyids), of Trichoptera

(hydroptilids, leptocerids, and brachycentrids), and of Diptera (simuliids, tanypodines,

orthocladiines, tanytarsines, and empids) declined after 1977 or 1978; in other locations many

of these taxa peaked in 1979 or 1980.

Some larvae dislodged by methoxychlor treatments apparently reattached in downstream

sites. In 25 to 73 percent of 23 tests there were increases rather than decreases in numbers of

various non-simuliid and simuliid taxa attaching to rope-piece substrates, 25 to 92 km

downstream during the week following an injection.

In summary, significant upward trends in average annual densities of suprageneric taxa

indicated that effects of methoxychlor treatments essentially were neutral when compared

with effects of unidentified extrinsic ecological processes. Furthermore, check lists of benthic

species collected from all three rivers at the conclusion of tests in 1980 proved the survival of

a varied fauna representing apparently complete ranges in feeding habits, activity patterns and

life cycles. Thus, the relatively intensive series of methoxychlor larvicide treatments required

to prevent damaging outbreaks of S. luggeri from the Saskatchewan River was not

permanently harmful to non-simuliid taxa in the river, at least at the suprageneric level.

'The Saskatchewan River in Saskatchewan.

54

Fredeen

RfeUME

Une solution de methoxychlor (0.3 ppm d’ingredients actifs) fut versee pendant 15 minutes dans le reseau de la

riviere Saskatchewan h savoir: en 1976 (une fois), 1977 (six fois), 1978 (sept fois), 1979 (19 fois) et 1980 (cinq fois). Au

cours des trois premieres annees, les populations de Simulium luggeri ont explose h divers endroits de la riviere,

atteignant une et endue de 200 km pres.

En adoptant le niveau supragenerique comme critere de classification, I’auteur a denombre les populations

d’invertebres non-simuliides de 1977 a 1980. La densite hebdomadaire moyenne de I’ensemble des populations

s'attachant aux substrats artificiels places au milieu du cours d’eau, dans la riviere Saskatchewan et ses deux

tributaires, a culmine en 1979 (P<0.01), I’annee d’utilisation maximale du larvicide. En 1980, cependant, la densite

populations a diminue presqu’au niveau enregistre en 1977. Egalement, I’amplitude des moyennes d’invertebres dans les

echantillons preleves du fond le long des rives a generalement culmine en 1979 ou en 1980 dans la riviere et ses deux

tributaires. Toutefois, a I’un ou plusieurs des six sites echantillonnes, les populations de certaines families

d’Ephemeropteres, de Trichopteres et de Dipteres ont diminue apres 1977 ou 1978; aux autres sites, plusieurs de ces

taxons ont culmine en 1979 ou 1980.

Des larves delogees par les traitements au methoxychlor ont paru se reattacher plus bas dans la riviere. Dans 25 h

73% des 23 tests effectues, on a remarque un accroissement inattendu du nombre des taxons simuliides et non-simuliides

s’attachant a des substrats faits de corde, situes de 25 d 92 km en aval durant la semaine suivant I’application du

larvicide.

En resume, un accroissement annuel significatif de la densite moyenne des populations etudiees a indique un effet

neutre des traitements. Les listes des especes recouvrees de la riviere Saskatchewan et de ses deux tributaires en 1980 ont

demontre la survie de toute une gamme d’organismes. L’auteur conclut que les traitements relativement intensifs

necessaires pour contrdler S. luggeri dans la riviere Saskatchewan n’ont pas affecte les taxons non-simuliides de facon

permanente.

TABLE OF CONTENTS

Introduction 54

Experimental 56

Results and Discussion 59

Need for Control of Black Flies at Sites of Breeding 89

Acknowledgements 91

References 91

INTRODUCTION

Single 7.5- to 15-minute injections of methoxychlor [2,2-bis(p-methoxyphenyl)-l,l,l-

trichloroethane] have been used to reduce populations of blaek fly larvae in selected seetions of

the Saskatchewan River in Saskatchewan almost every year, 1968 to the present time. Reports

about effieaey and environmental impaet of 12 injections during the first six years of these tests

have been published (Fredeen, 1974, 1975).

In the final and most eomprehensive test in that series, populations of aquatic insect larvae

were measured weekly in one untreated site and in four treated sites spaced at 40 km intervals

in the North Saskatchewan River, throughout 1 1 consecutive weeks commencing one week

before a single 7.5 minute injection of 0.6 parts of methoxychlor per million parts of water

(p.p.m.). Downstream 161 km from the injection point, 66 percent of larval instars three to six

of Simulium arcticum Malloch and 96 percent of instars one and two disappeared within the

first week. Larger percentages were removed from nearer sites. Larvae of plecopterans were

similarly affected but ephemeropterans, trichopterans, and chironomids were less affected. All

four treated sites were rapidly repopulated. Populations of chironomid larvae larger than one

mm long equalled or exceeded pre-treatment densities within one to three weeks,

ephemeropterans within one to four weeks, trichopterans in one to seven weeks, plecopterans in

Aquatic invertebrates in a large Canadian River

55

four to five weeks, and simuliids within two to ten weeks. Populations of larvae smaller than one

mm long were generally restored more rapidly. Fish were not visibly affected (Fredeen, 1975).

In 1979 this “single-injection” pattern for methoxychlor was registered in Canada for

control of larvae of S. arcticum in large rivers. Before that, methoxychlor was registered for use

in Canada as a black fly larvicide only if applied by air across networks of shallow streams in

200 m wide swaths centered 400 m apart. Dosages of about 25 to 85 g of active ingredient (a.i.)

per swath - ha would have been achieved with this method, equivalent to less than 0.01 p.p.m.,

a.i. sustained for about 0.5 minutes in a deep river such as the Saskatchewan River. Such a

dosage would not have been effective against black fly larvae in this river because tests showed

that exposures to about 0.2 p.p.m., maintained for 15 minutes were only partly effective

(Fredeen 1974). Futhermore, logistics and cost of applying aerial swaths at 400 m intervals

throughout the entire infested portion of the Saskatchewan River (150 to 200 km) would have

been impractical.

It is not surprising that certain large rivers are sources of troublesome, chronic outbreaks of

insects. Nor is it surprising in view of wide differences in habitats between rivers, that varieties

of troublesome species vary widely between rivers. Munroe (1951) and Peterson (1952) listed

34 species of Trichoptera, mainly hydropsychids, emerging in nuisance numbers from the

Niagara River. Corbet et al. (1966) reported that eight species of Trichoptera, again mainly

hydropsychids, dominated nuisance swarms of insects emerging from the St. Lawrence River at

Montreal. These insects created allergic reactions among residents, and navigational problems

for ships in the St. Lawrence Ship Channel and vehicles on nearby highways.

Fremling( 1960(a), (b)) reported that massive flights of two species of mayflies {Hexagenia

bilineata (Say) and H. limbata (Serville)) and one species of caddisfly {Cheumatopsyche

campyla Ross) caused major nuisance and health problems in cities along the upper Mississippi

River. Fredeen (1969) reported that larvae of the black fly S. arcticum Malloch were

widespread in rivers and streams draining the eastern slopes of the Rocky Mountains, and that

massive outbreaks, resulting in livestock losses, originated from certain portions of the

Saskatchewan and Athabasca Rivers. Thus of the five large Nearctic rivers reported to have

produced troublesome numbers of aquatic insects, economically important outbreaks of black

flies have originated only from the Saskatchewan and Athabasca Rivers.

In the early 1970’s, S. luggeri Nicholson and Mickel replaced S. arcticum as the dominant

black fly species breeding in the Saskatchewan River in Saskatchewan, and within a few years

it became a major pest of man and non-hominid animals (Fredeen, 1977). These changes in

black fly populations coincided with major changes in their larval environments in the

Saskatchewan River system in the 1970’s. Summertime monthly water-flow volumes declined

in the South Saskatchewan River to as low as 7.5 percent of the long term monthly averages

(June, 1977) and in the North Saskatchewan River to as low as 34 percent (August, 1975)

(Environment Canada 1977, 1978, 1979, 1980, 1981). This occurred in part because of

completion of three hydroelectric dams in the river system and in part because of widespread

drought in the mid 1970’s. Previously these rivers had been deep, swift and turbid during

summer months. Now they are relatively shallow, slow-flowing and clear, allowing dense

growths of several species of water weeds for the first time. These plants are favored

attachment sites for larvae of S. luggeri and several other black fly species previously found

only in smaller rivers. An increase in tolerance to methoxychlor is not considered to have been

responsible for these changes of black fly species because populations of larvae in treated

sections are continually replenished by downstream drift from extensive untreated sections of

Quaest. Ent. 1983, 19 (1,2)

56

Fredeen

the rivers. However, L.D. 50’s for larvae of these species of black flies have not been

determined.

These changes in black fly communities and river conditions forced changes in abatement

strategies. Whereas S. arcticum required control only in May, S. luggeri cycles continuously

throughout May to September. Thus, in some recent years, larvicide has been injected at two-

to four-week intervals to prevent damaging outbreaks.

Furthermore, it became necessary to space larvicide injection sites closer together

geographically. Vast beds of water weeds developed upon previously barren sand bars when the

river became relatively shallow and clear due to the combined effects of drought and

summertime impoundment of water behind newly-built hydroelectric dams (Fredeen, 1977).

These weed beds not only provided large extensions of larval attachment sites but unfortunately

also rendered larvicide treatments less effective, presumably because of filtering effects. Thus it

is now sometimes necessary to space larvicide injection sites only 20 to 50 km apart to achieve

adequate control.

A four-year environmental impact study reported herein was initiated in 1977 to investigate

long-term effects of this recently intensified larviciding program. An earlier study (Fredeen

1975) had indicated that when a 161 km section of the North Saskatchewan River was treated

with methoxychlor, populations of invertebrates were restored to pre-treatment densities within

a few weeks.

EXPERIMENTAL

Larvicide

The larvicide used throughout was a commercial emulsifiable concentrate containing 0.24

kg methoxychlor per litre. Treatments were performed under federal and provincial permits,

renewed annually.

Injection sites

Locations of sites of larvicide injections are shown in Figure 1 and Table 1. All sites were

located in central Saskatchewan within less than 1 30 km from the confluence of the north and

south branches of the Saskatchewan River. Specific locations within each site that were used

for injections and/or assessments are described in greater detail in the following pages.

Most injections were made from motorized ferries which allowed four continuous swaths

across the entire river during each 15-minute injection. Only three injections were from fixed

points instead of swaths across the river, all from a traffic bridge (Site 2, Fig. 1) spanning the

North Saskatchewan River at Prince Albert in 1978. The main Saskatchewan River just

downstream from the confluence of the north and south branches (Site 4) was injected from a

Sikorsky helicopter with a long tube discharging just beneath the water surface. The helicopter

crossed the river four times during each 15-minute injection.

Tests in 1977 and 1978 showed that injections from a single location (either Fenton Ferry

Site 6 or Birch Hills Site 7) on the South Saskatchewan River were inadequate. Much of that

river remained infested despite treatments, presumably because dense weed beds reduced

effectiveness of the larvicide. Thus, in 1979 treatments sometimes were spaced 20 to 50 km

apart and treatments also were initiated at the confluence (Site 4) when it became evident that

treatments had not travelled effectively beyond that point.

Aquatic invertebrates in a large Canadian River

57

TABLE 1. LIST OF SITES WHERE METHOXYCHLOR BLACK FLY LARVICIDE

WAS INJECTED, AND/OR IMPACT STUDIES CONDUCTED, SASKATCHEWAN

RIVER SYSTEM IN SASKATCHEWAN.

(OSee Fig. 1 to locate these sites on map.

Sampling sites for invertebrate populations

Sampling sites were fixed throughout the four-year study period. Permanently untreated

check sites were not selected at the outset because it was not possible to predict where larvicide

would be injected in each of the four years and also because our research team was not large

enough to cope with additional sites.

Sampling methods

Two methods were used consistently throughout the four summers to measure invertebrate

populations; (a) artificial substrates (rope pieces) anchored in three mid-river locations to

measure weekly increments of drifting populations; and (b) Surber-type net sampling in six

locations along river margins to collect samples of benthic populations.

Artificial substrates. — One-m lengths of 0.5 cm diameter polypropylene rope (Fredeen and

Spurr, 1978) served as artificial substrates. Each rope piece was anchored so that it floated just

under the water surface. Polypropylene has the correct specific gravity for this purpose. Two

anchors, each with one attached rope piece, were placed about mid-channel, about one km

upriver from each of the three selected sites about one week after ice break-up each spring in

water flowing at about 0.5 to 1.0 m/sec depending upon river volum.e. The rope pieces were

collected and replaced with new rope pieces weekly throughout each summer. One pair of

Quaest. Ent. 1983, 19 (1,2)

58

Fredeen

106 105 104

Fig. 1 . Map showing experimental sites on the North, South, and Main Saskatchewan Rivers in Saskatchewan.

anchors was located about one km upstream from Cecil Ferry in the North Saskatchewan

River (Site 3, Fig. 1), another pair was located about one km upstream from Birch Hills Ferry

in the South Saskatchewan (Site 7), and the third pair was located about one km upstream

from Gronlid Ferry in the main Saskatchewan River (Site 9). All three sites were served by “all

weather” roads to ferries which ensured uninterrupted weekly access throughout all four

summers. More than 60 percent of all larvicide injections occurred above these sites and 40

percent downstream (Tables 2, 3, and 4). Rope pieces were exchanged weekly for 17

consecutive weeks in Sites 3 and 7, and for 15 consecutive weeks in Site 9. The samples were

individually preserved in 95 percent ethanol until analysis of attached invertebrates. Generally

only one sample from each pair was analyzed.

These samples from artificial substrates were used for two purposes: (1) To provide weekly

counts of immigrant populations of black fly larvae to estimate need for larvicide treatments.

(A weekly accumulation of 1000 or more black fly larvae per 100 cm of rope indicated that

larvae were arriving in numbers sufficient to cause damaging outbreaks unless controlled.); (2)

To measure long-term trends in numbers of drifting invertebrates in mid-river locations. It was

assumed that populations seen in these samples were related numerically to river bed

populations from whence the drifting invertebrates had originated even though exact sources

were not known (Fredeen and Spurr, 1978).

Benthic samples. — Samples of benthic invertebrates were collected from river margins

under 50 to 60 cm of water with a 645 cm^ Surber-type net with 0.2 mm mesh openings at

Aquatic invertebrates in a large Canadian River

59

weekly intervals for three consecutive weeks each August. Rocky beds precluded sampling with

an automatic dredge. The month of August was selected because larvicide treatments were

completed by that time each year, and because river levels were generally stable or slowly

declining, allowing weekly collections without interruptions, from permanently inhabited

portions of the river bed accessible on foot. In 1979, a one-week interruption (August 21) of the

weekly regime occurred due to a brief rise in the level of the South Saskatchewan River. In

1979, there were no collections from the main Saskatchewan River (Site 9) because of

insufficient staff.

In each of the three selected weeks each year, five 645 cm^ samples (from a total of

approximately 3225 cm^) of river bed material to a depth of more than five cm were collected

from each of six locations: each side of the North Saskatchewan River about one km above

Cecil Ferry (Site 3), the north side of the North Saskatchewan about one km above the

confluence (Site 4), the south side of the South Saskatchewan about two km above the

confluence (Site 4), and each side of the main Saskatchewan River about one km above Gronlid

Ferry (Site 9). Most sites were subjected to one or more larvicide treatments each year, the

notable exception being the two Site 3 locations which were not treated in 1980. Collections

from the North Saskatchewan River, Site 3 north side, were from the effluent path of a pulp

mill, nine km upstream.

Each batch of five samples was combined into a single sample and preserved in 95 percent

ethanol. Specimens were analyzed to families or sub-families. In 1980, keys became available

for identification of species of Ephemeroptera, Plecoptera, Trichoptera, and Diptera inhabiting

the Saskatchewan River in Saskatchewan, allowing preparation of a check list of species found

in benthic samples during that year.

Data analysis. — Extreme variabilities characterized substrate and benthic populations (x,)

and for analysis, logarithmic transformations of the form log (jc, + 1) were applied.

Transformed data were analyzed by the least squares method using the formula: y,^ = /u. + t,

+ Wj + e,y where y,y was the transformed observation in the week (w) of the year (t).

Anti-log conversions of least squares means obtained in analyses provided the average values

shown in tables and text. All tests of significance were based upon transformed data.

RESULTS AND DISCUSSION

Physical and chemical condition of the rivers, 1977-1980

Ice-free conditions on the Saskatchewan Rivers commenced each year during the third week

of April except in 1979 when it commenced May 3 for the north branch and May 12 for the

south. Both rivers refroze during the second or third week of November each year.

Mean monthly water discharge rates for the North Saskatchewan River, 1977, 1978, 1979,

and 1980 for the months of May, June, July, and August were about 365, 490, 410, and 260

m3/sec respectively, and for the south branch about 85, 165, 115, and 85 m^/sec for the same

months (Environment Canada, 1978, 1979, 1980 and 1981; Fisheries and Environment Canada

1977).

Turbidity varied approximately with the discharge rates and ranged from about 10 to 1000

gm/m^ of water in the north branch and from about 10 to 100 gm/m^ in the south branch.

Quaest. Ent. 1983, 19 (1,2)

60

Fredeen

Mean daily water temperatures increased from about 1° to 22°C in May and ranged from

16° to 24°C in June, 17° to 27°C in July, and 14° to 23°C in August.

The pH ranged from about 7.9 to 9.0 in May to about 8.0 to 9.1 in August in both rivers.

Phenolphthalein alkalinity ranged from 0 to about 20 (as p.p.m. CaC03) and total alkalinity

ranged from about 100 to 200 (as p.p.m. CaC03). Hardness ranged from about 130 to 200

p.p.m. (as CaC03) in both rivers.

Larvicide treatments

Complete lists of injections of methoxychlor into the north, south, and main branches of the

Saskatchewan River, 1974 to 1980 inclusive, are shown in Tables 2, 3, and 4. All previous

treatments were reported by Fredeen (1974, 1975).

The larviciding campaign on the Saskatchewan River System was expanded from one

treatment in 1976 to six in 1977, seven in 1978, and 19 in 1979, in attempts to reduce intensity

and duration of outbreaks of S. luggeri that frequently extended 100 to 150 km or even further

from the river in 1976, 1977, and 1978. Initially, the campaign was based upon one developed

for control of S. arcticum, where one or two larvicide injections per year were generally

sufficient to prevent a major outbreak. However, S. luggeri, which became the dominant

species in 1976, cycled continuously all summer, and larvae attached in large numbers in

aquatic weed beds newly developed in shallow sections of the rivers. Injections spaced about

four weeks apart in 1977 and 1978 successfully removed many larvae but the relatively long

intervals between treatments allowed many other larvae to complete growth and produce

abundant females which returned to re-populate the rivers with eggs. Also, injections from a

single location in the South Saskatchewan River in 1977 and 1978 were inadequate. Much of

that river sometimes remained infested, presumably because dense weed beds reduced

effectiveness of the methoxychlor, perhaps by adsorbing it from the water. Furthermore,

treatments did not seem to extend effectively beyond the confluence of the North and South

Saskatchewan Rivers. Increasingly massive outbreaks originated from the main Saskatchewan

River some 70 km or further downstream from the confluence in 1977 and 1978.

Requirement for increased numbers of methoxychlor treatments was believed not due to

development of increased tolerance. Weekly samples collected from artificial substrates above

Site 3 in 1980 and Site 7 in 1977, years in which these sites were not treated, showed that

downriver drift of black fly larvae from untreated sections occurred continuously throughout

each summer.

Thus in 1979, the program was expanded to allow as required (a) injections to be spaced

closer together in time, (b) multiple simultaneous injections at several sites in the weedy South

Saskatchewan River, and (c) injections for the first time into the main Saskatchewan River

below the confluence. Infestations of larvae were treated with methoxychlor larvicide injected

at eight locations that year, three of which were injected only once during the summer, two,

twice, and the others, three to five times. Relatively few larvae were allowed to mature and

produce adults. For the first time in four years, livestock and people along the entire river

experienced major relief from black flies.

In 1980 only one injection was required on the North Saskatchewan River, two at two sites

each on the south branch, and none on the main branch.

Aquatic invertebrates in a large Canadian River

61

TABLE 2. COMPLETE LIST OF METHOXYCHLOR BLACK FLY LARVICIDEO)

TREATMENTS, NORTH SASKATCHEWAN RIVER, 1974 TO 1980, INCLUSIVE.

(OEmulsifiable concentrate containing 0.24 kg active ingredient per litre.

(2)See Fig. 1 to locate sites on map, and Table 1 for distances.;

Quaest. Ent. 1983, 19 (1,2)

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TABLE 3. COMPLETE LIST OF METHOXYCHLOR BLACK FLY LARVICIDEO)

TREATMENTS, SOUTH SASKATCHEWAN RIVER, 1974 TO 1980, INCLUSIVE.

(OEmulsifiable concentrate containing 0.24 kg active ingredient per litre.

(2)See Fig. 1 to locate sites on map, and Table 1 for distances.

Aquatic invertebrates in a large Canadian River

63

TABLE 4. COMPLETE LIST OF METHOXYCHLOR BLACK FLY LARVICIDEO)

TREATMENTS, MAIN SASKATCHEWAN RIVER, 1974 TO 1980, INCLUSIVE.

(1) Emulsifiable concentrate containing 0.24 kg active ingredient per litre.

(2) See Fig. 1 to locate sites on map, and Table 1 for distances.

Quaest. Ent. 1983, 19 (1,2)

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Benthos, River Margins Substrates, Mid-River

Fig. 2. Diagrammatic representations of approximate proportions of invertebrates representing major behaviour and

feeding types in marginal benthos and in mid-river substrate samples collected from the Saskatchewan River, 1977

through 1980.

Habits and trophic relationships of invertebrates collected

Insect larvae. — The varied habits and trophic relationships of larvae of aquatic insects

inhabiting the Saskatchewan River (Table 5, Fig. 2) are important when considering potential

impact of larvicide treatments. In benthos samples from the margins of the Saskatchewan

River most taxa (representing about 80 percent of the total insects collected) were considered

to be dingers or climbers (Merritt and Cummins, 1978). The remainder, mainly Chironomus

spp. and tanytarsines, as well as the rarer polymitarcyids and anisopterans were presumed to be

burrowers or tube builders. The relatively abundant hydropsychids inhabited fixed retreats.

About one percent of the total population were case-building trichopterans.

In samples from artificial substrates from mid-river sites more than 95 percent of the total

insects collected were dingers or climbers.

Regarding feeding habits, according to Cummins (1973), Merritt and Cummins (1978),

and Wiggins (1977), most or all of our ephemeropterans, trichopterans, and dipterans were

considered to be collectors of periphyton, debris, and plankton. Many of these, including

simuliids, hydropsychids, and many species of Ephemeroptera, Chironomini, and Tanytarsini,

were filter feeders. In benthos samples about 50 percent of the larvae were considered to be

filter feeders (Fig. 2) and in samples from mid-river artificial substrates about 90 percent

(mainly simuliids) were filter feeders. About two-thirds of the remainders were assumed to

Aquatic invertebrates in a large Canadian River

65

TABLE 5. HABITS AND TROPHIC RELATIONSHIPSO OF AQUATIC INSECTS

INHABITING THE SASKATCHEWAN RIVER IN SASKATCHEWAN.

Taxa

EPHEMEROPTERA

Siphlonuridae: Isonychia

Baetidae; Baetis

Stenonema

Ephemerellidae:

Ephemerella

Perlidae: Acroneuria

HEMIPTERA: Sigara

COLEOPTERA:

Dytiscidae: Deronectes

Helodidae

TRICHOPTERA

Psychomyiidae:

Psychomyia

Polycentropodidae:

Neureclipsis

(continued on next page)

Habits

Swimmers, dingers

Swimmers, climbers, dingers

Clingers

Clingers, swimmers

Clingers

Swimmers, climbers

Swimmers, climbers

Climbers

Clingers (tube retreats)

Clingers (net builders)

Trophic Relationships

Collectors (filterers); engulfers

(predators)

Collectors (gatherers)

(detritus, diatoms);

scrapers

Scrapers; collectors (gatherers)

Scrapers; collectors (gatherers)

(engulfers)

Collectors (gatherers); scrapers

Collectors (gatherers)

(detritus, algae)

Collectors (gatherers)

Collectors (gatherers); scrapers

Collectors (filterers)

Collectors (gatherers)

Engulfers (predators)

Engulfers (predators)

Shredders (detritovores)

engulfers (predators)

Engulfers (predators)

Piercers (herbivores); collectors

(gatherers)

Piercers (carnivores)

Scrapers; collectors; shredders;

piercers (herbivores)

Collectors (gatherers)

Collectors (filterers)

(herbivores, predators)

Pseudocloeon Swimmers, clingers

Heptageniidae: Heptagenia Clingers, swimmers

Tricorythidae:

Tricorythodes Sprawlers, clingers

Caenidae: Caenis Sprawlers

Leptophlebiidae:

Traverella Clingers

Polymitarcyidae: Ephoron Burrowers

ODONATA

Anisoptera: Ophiogomphus Burrowers

Zygoptera: Ischnura Climbers

PLECOPTERA

Pteronarcidae; Pteronarcys Clingers, sprawlers

Quaest. Ent. 1983, 19 (1,2)

66

Fredeen

Table 5 (continued)

collect food by ‘gathering’ rather than by ‘filtering’ and about one third were assumed to be

either herbivores (piercers or shredders) or carnivores (Table 5).

In general, the invertebrate fauna of the Saskatchewan River was dominated by

filter-feeding black fly larvae (tables 6, 7, 8, 9).

Crustaceans. — Five major taxa of crustaceans were collected. All could be considered

free-swimming although in these rivers they would have lived near or on the substratum (Ward

and Whipple 1959). Ostracods comprised about 80 percent of the total crustacean population

and occurred regularly in all six sites sampled with the Surber-type net. Copepods and

cladocerans were found in the North and South Saskatchewan Rivers but not in the main river

below the confluence. Conchostracans and malacostracans were seldom collected.

Ostracods and copepods are shredders and feed on decaying plant and animal materials.

Cladocerans are filterers (plankton) and malacostracans {Hyalella spp.) are shredders and

filterers.

Acari. — Larvae of Parasitengona, found in all sites, are parasitic on aquatic insects and the

adults are predaceous.

Mollusca. — Pelecypods (Sphaeridae) were relatively abundant and widely distributed in

the Saskatchewan River. They are filterers, subsisting on detritus and plankton. Gastropods

(mainly Ancylidae) were scarcer and less widely distributed. They are browsers, feeding on

algae and detritus.

Identification of Invertebrates

It was within our expertise to identify Simuliidae to species from the outset. However, keys

to identify many species in other major taxa were not available until after we had completed

analyzing substrate and benthic samples in 1980. All samples have been retained at our

Research Station in the event that identification of additional taxa to species levels would

eventually prove productive.

Aquatic invertebrates in a large Canadian River

67

Trends in numbers of invertebrates attached to artificial substrates

Population trends for taxa sampled with artificial substrates weekly during the summers of

1977 to 1980, inclusive, are given for the North Saskatchewan River (Site 3) in Table 6, for the

South Saskatchewan River (Site 7) in Table 7, and for the main Saskatchewan River about 70

km downriver from the confluence of the two branches (Site 9) in Table 8.

Note that Site 3 received only one treatment in 1977, three in 1978, five in 1979 and none in

1980. Site 7 received no treatments in 1977, four in 1978, five in 1979 and two in 1980. The

main Saskatchewan River below the confluence was not treated in 1977, 1978 or 1980 so that

Site 9 received four treatments in 1979 but none in the other three years unless one assumes

that effects from some or all of the six, seven, 14, and five injections into the two branches

above the confluence may have affected populations at Site 9. Permanently untreated check

sites were not available for reasons stated earlier in this paper.

Mites peaked in 1979 at Sites 3 and 7 (P<0.01 at Site 7) and in 1980 at Site 9 (P<0.01).

This suggests that there were parallel increases in numbers of certain invertebrate taxa upon

which these animals preyed. Alternatively, as discussed in a following section, displaced larvae

may have reattached further downriver.

Mean annual numbers of plecopterans attaching weekly to artificial substrates declined

after 1978 at Site 3 (P<0.01) but remained relatively unchanged at Sites 7 and 9. Larvae of

ephemeropterans remained relatively abundant and unchanged in numbers at Sites 3 and 7 but

peaked in 1980 at Site 9 (P<0.01). Larvae of trichopterans remained relatively abundant but

unchanged in numbers at all three sites. Larvae of chironomids, the most abundant of all

non-simuliid taxa, peaked in 1979 in all three sites with highly significant differences between

years in Sites 7 and 9 (P<0.01).

Numbers of larvae of S. luggeri declined annually after 1977 in Site 3 (n.s.) and after 1978

in Sites 7 and 9 (P<0.01 at both sites) to four-year lows in all three sites. Numbers of S.

arcticum and S. meridionale Riley were relatively small compared to numbers of S. luggeri

and remained unchanged in Sites 3 and 9, but declined after 1977 in Site 7 (P<0.05).

Numbers of S. vittatum Zetterstedt peaked in 1977 in Site 7 (P<0.01) and in 1979 in Site 9

(P<0.01).

Larvae of these four species of Simulium were considered to be relatively susceptible to

methoxychlor, not only because they inhabited sites that would have ensured direct contact

with the larvicide but also because they were filter feeders and thus would have ingested

suspended particles containing adsorbed methoxychlor (Fredeen et al., 1975). Despite this, no

single species of Simulium was eliminated during the four years of treatment. In fact, S.

vittatum (not a pest species) actually attained maximum abundance at two sites in 1979, the

year of maximum use of methoxychlor.

These data help to explain why prominent non-simuliid taxa remained abundant, and of

greater importance, suggest that trends in numbers of each non-simuliid order or family as

shown in Tables 6, 7, 8 and 9 are actually representative of parallel trends in numbers of most

or all of the species comprising those suprageneric taxa.

Densities of populations of invertebrates on rope-piece substrates were believed related to

densities of the benthic populations from whence the drifting populations had originated.

Substrates offered convenient means of obtaining uninterrupted series of weekly samples from

otherwise inaccessible mid-river sites throughout the four-year study. However, it was not

possible to determine how far larvae had drifted in the rivers before attaching to the substrates.

Presumably many (most?) larvae originated from treated sections of the rivers because

Quaest. Ent. 1983, 19 (1,2)

68

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TABLE 6. MEAN NUMBERSO) OF AQUATIC INVERTEBRATES ATTACHED TO

ROPE PIECES(2) ANCHORED FOR ONE-WEEK PERIODS THROUGHOUT 17

WEEKS EACH YEAR, MAY TO SEPTEMBER, ONE KM UPRIVER FROM SITE 3(3)

(CECIL FERRY), NORTH SASKATCHEWAN RIVER, SASKATCHEWAN, 1977 TO

1980 INCLUSIVE.

(OGeometric means calculated from logjo (x + 1) values.

(2) Polypropylene rope: length = 100 cm; diam. = 0.5 cm.

(3) See Fig. 1 to locate sites on map, and Table 1 for distances.

(4) These statistics were calculated from log,o (x + 1) values. Means followed by

different letters, differ significantly (P<0.05) as indicated by Duncan’s

New Multiple Range tests.

Aquatic invertebrates in a large Canadian River

69

TABLE 7. MEAN NUMBERSU) OF AQUATIC INVERTEBRATES ATTACHED TO

ROPE PIECES(2) ANCHORED FOR ONE-WEEK PERIODS THROUGHOUT 17

WEEKS EACH YEAR, MAY TO SEPTEMBER, ONE KM UPRIVER FROM SITE 7(3)

(BIRCH HILLS FERRY), SOUTH SASKATCHEWAN RIVER, SASKATCHEWAN,

1977 TO 1980 INCLUSIVE

(OGeometric means calculated from logjo (x + 1) values.

(2) Polypropylene rope: length = 100 cm; diam. = 0.5 cm.

(3) See Fig. 1 to locate sites on map, and Table 1 for distances.

C^)These statistics were calculated from logjo (x + 1) values. Means followed by

different letters, differ significantly (P<0.05) as indicated by Duncan’s

New Multiple Range tests.

Quaest. Ent. 1983, 19 (1,2)

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TABLE 8. MEAN NUMBERSd) OF AQUATIC INVERTEBRATES ATTACHED TO

ROPE PIECES(2) ANCHORED FOR ONE-WEEK PERIODS THROUGHOUT 15

WEEKS EACH YEAR, MAY TO SEPTEMBER, ONE KM UPRIVER FROM SITE 9(3)

(GRONLID FERRY), MAIN SASKATCHEWAN RIVER, SASKATCHEWAN, 1977 TO

1980 INCLUSIVE

(^Geometric means calculated from logio (x + 1) values.

(2) Polypropylene rope: length =100 cm; diam. = 0.5 cm.

(3) See Fig. 1 to locate sites on map, and Table 1 for distances.

(4) Numbers in brackets indicate numbers of treatments in the entire Saskatchewan

River system above the sampling site. Unbracketed numbers indicate numbers

of treatments in the Main Saskatchewan River alone.

(3)These statistics were calculated from logio + 1) values. Means followed by

different letters, differ significantly (P<0.05) as indicated by Duncan’s

New Multiple Range tests.

Aquatic invertebrates in a large Canadian River

71

TABLE 9. MEAN NUMBERSO) OF AQUATIC INVERTEBRATES ATTACHED TO

ROPE PIECES(2) ANCHORED FOR ONE- WEEK PERIODS THROUGHOUT 15

WEEKS EACH YEAR, MAY TO SEPTEMBER, 1977 TO 1980 INCLUSIVE.

COMBINED DATA FROM THREE SITES(3), SASKATCHEWAN RIVER,

SASKATCHEWAN.

(OGeometric means calculated from logjo (jc + 1) values.

(2) Polypropylene rope: length =100 cm; diam. = 0.5 cm.

(3) One km upriver from each of sites 3, 7, and 9 (Fig. 1), in the North, South, and

Main Saskatchewan Rivers respectively.

(4) These statistics were calculated from logjo (x + 1) values. Means followed by

different letters, differ significantly (P<0.05) as indicated by Duncan’s

New Multiple Range tests.

Quaest. Ent. 1983, 19 (1,2)

72

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TABLE 10. PERCENTAGES OF METHOXYCHLOR LARVICIDE TESTS IN WHICH

SAMPLES OF AQUATIC INVERTEBRATES FROM ROPE-PIECE SUBSTRATESO)

WERE LARGER THAN THOSE OF THE PREVIOUS WEEK, SASKATCHEWAN

RIVER SYSTEM, 1977 TO 1980, INCLUSIVE.

Percentages of tests showing increased densities (%)

(OOne-metre lengths of 0.5 cm diameter polypropylene rope anchored for one week

at sites 3, 7, and 9 (Fig. 1).

(2)Data from many weeks during the four summers could not be included because:

a. ) the river was reinjected within less than two weeks before post-treatment

samples could be collected,

b. ) injections were repeated on one date in more than one site in a single river,

c. ) samples from site 9 in the Main Saskatchewan River were not included in

this summation if either of the two branches above the confluence had been

injected within the two previous weeks.

Aquatic invertebrates in a large Canadian River

73

sampling sites were located many km downriver from many or all larvicide injection sites. The

most distant injection sites upstream from the three sampling sites were 92 km for Site 3, 84

km for Site 7 and 198 km for Site 9.

In summary, combined data from numbers of invertebrates attaching weekly to artificial

substrates anchored mid-river in three sites (Table 9) show that mean densities of non-simuliid

taxa in treated sections of these rivers peaked in 1979, but in 1980 returned to about 1977

densities. Differences between years were highly significant (P<0.01). Mean annual densities

of simuliids (excluding S. vittatum) peaked in 1978 and then declined rapidly during the final

two years of the program to a four-year low in 1980 (about one-fifth the density of 1978) and

differences between years were highly significant. When numbers of S. vittatum larvae are

included in these means, total populations peaked in 1979 and then also declined in 1980 to

about one-fifth of the peak value. S. vittatum larvae were totalled seperately because some

females of this species laid eggs on floats supporting rope-piece substrates and some larvae

found attached to the rope pieces may have hatched from those eggs rather than arriving as

drifting larvae.

Increases in numbers of invertebrates following larvicide injections

Many collections of invertebrates from mid-river rope-piece substrates were larger

immediately after larvicide injections than before (Table 10). This was particularly true for

rope pieces anchored at relatively great distances downstream from injection sites. Thus

collections from rope pieces anchored for one week 71 to 92 km downstream from an injection

site contained larger numbers of mites, ephemeropterans, trichopterans, chironomids, and

simuliids in 64 to 73 percent of the samples collected immediately after an injection than

before. In 37 percent of those same post-treatment collections, numbers of plecopterans were

larger than in pre-treatment collections.

From substrates anchored nearer the injection sites (i.e. 25 to 32 km downstream) numbers

of these same taxa in post-treatment samples exceeded those in pre-treatment collections in

only 25 to 58 percent of the collections.

In comparison, in the absence of larvicide treatments, consecutive weekly pairs of collections

showed increases in the second week in 36 to 61 percent of the samples (Table 10).

These data provided new evidence that certain detached larvae survived and successfully

reattached in sites further downriver as reported earlier by Fredeen (1974, 1975). The

relatively substantial recolonization following methoxychlor treatments would also have been

aided by regular drift of larvae newly hatched from eggs unaffected by methoxychlor, and

larvae from untreated upstream sections and from protected niches in treated sections. Fredeen

(1975) showed that larvae in sand beds in the Saskatchewan River were entirely unaffected by

passage of methoxychlor-treated water. Pupae in general appeared to be resistant to

methoxychlor and as well, pupae of certain chironomids and hydroptilids drifted into treated

sections, attached to fragments of water weeds.

A prerequisite to successful reattachment by larvae would have been the reduction of

methoxychlor concentrations to tolerable levels. The main Saskatchewan River downstream

from the confluence would generally have received diluted larvicide from the branches.

Although both branches above the confluence often were injected with methoxychlor on the

same day (Tables 2 and 3) it was very unlikely that treated masses of water arrived at the same

time at the confluence.

Quaest. Ent. 1983, 19 (1,2)

74

Fredeen

Also Fredeen et al. (1975) showed that as an injected mass of water travelled downstream it

became progressively attenuated due to friction with the river bed. Furthermore adsorption to

river bed sand was demonstrated. Alternatively, adsorption of methoxychlor to particles

suspended in the water, especially in the relatively turbid North Saskatchewan River, would

have aided long distance transport of methoxychlor.

Adsorption to water weeds, and filamentous algae, especially in extensive beds of several

species in the South Saskatchewan River, was not proved but may be assumed. Edwards and

Glass (1971), Butler et al. (1975), Paris and Lewis (1976) and others demonstrated that

methoxychlor was rapidly adsorbed to grass, many species of algae, fungi and bacteria.

Data from limited laboratory tests indicate that larvae of certain aquatic invertebrates are

less susceptible to methoxychlor than are black fly larvae. Sebastien and Lockhart (1981)

showed that 100 percent of stonefly nymphs {Pteronarcys dorsata Say) were moribund after 24

hours of exposure to 0.3 mg/1 of methoxychlor formulated as an emulsifiable concentrate, in

recirculated, dechlorinated water at 17°C. Forty-eight hours of exposure of chironomid larvae

{Chironomus tentans Fabricius) to 0.3 mg/1 at 20°C followed by 48 hours in fresh water

produced 99 percent mortality. In comparison, fifth and sixth instar black fly larvae {S.

decorum Walker) suffered 100 percent detachment during 16-minute exposure to 0.3 mg/1 of

methoxychlor at 19°C, and 100 percent mortality when returned to fresh water for about 20

hours. Fredeen (1972) reported that the L.C. 50 for relatively full-grown larvae of

Hydropsyche morosa Hagen following a six-hour exposure at 10°C to methoxychlor and 18

hours in fresh water was 0.04 mg/1 of methoxychlor. Anderson and DeFoe (1980) reported that

following continuous exposure to methoxychlor in flowing water throughout 28 days the L.C.

50 for the isopod Asellus communis Say was 0.42 m/ug/l, and for Hydropsyche sp. larvae was

1.3 m/ug/1. Stonefly larvae P. dorsata Say and a snail Physa Integra did not die at the highest

concentration tested, 4.2 m)ug/l.

Muirhead-Thomson (1973) suggests several reasons why one cannot use data from

laboratory bioassays of toxicants to accurately predict events in field tests. Furthermore, the

above bioassays were based upon relatively long exposures (6 hours to 28 days) whereas in our

field tests with methoxychlor a treated pulse of water lasted only 15 minutes at the point of

origin. Nevertheless, data quoted above from these few bioassays, and especially those from

Sebastien and Lockhart (1981) suggest that certain non-simuliid species are less sensitive than

simuliid larvae to small concentrations of methoxychlor.

Long-term effects of downstream displacements of larvae

As to long-term effects of downstream displacements of larvae due to larvicide injections

and other causes, comparisons of data presented in Tables 6, 7, and 8 show that numbers of

mites, plecopterans and ephemeropterans attaching weekly to artificial substrates anchored in

Site 9 (about 70 km below the confluence) peaked in 1980, one to three years after peaks were

observed in the tributaries (Sites 3 and 7). Numbers of trichopterans did not trend significantly

at any of the three sites but remained relatively abundant throughout the four years. Larvae of

chironomids also remained abundant throughout but peaked in all three sites in the same year

(1979) and differences between years were highly significant. Identification to the species level

may have revealed differences in responses between species in each of these major taxa but this

was not investigated. For reasons previously explained, changes in species complexes are

believed not to have occurred.

Aquatic invertebrates in a large Canadian River

75

Downstream displacements of black fly larvae presumably were responsible for increasingly

dense accumulations of larvae of S. luggeri in the main Saskatchewan River and in the

downstream end of the South Saskatchewan which resulted in major outbreaks from those

regions in 1977 and 1978. Average weekly numbers of larvae of S. luggeri near Site 7, South

Saskatchewan River (Table 7) increased from 1045 in 1977 (the year that that river was

injected below the collecting site) to 1551 in 1978 when the river was injected some 32 km

upstream (Tables 1 and 3). But when this weedy river was injected at four sites in 1979,

average number of larvae was reduced to 271 per week and there were no destructive outbreaks

that year from the South Saskatchewan River.

In the North Saskatchewan River treatments carried longer distances, presumably because

that river was deeper, more turbid, and contained sparser weed beds. Injections at single sites

(Table 2) were sufficient to steadily reduce populations of black fly larvae from a weekly mean

of 3236 larvae in 1977 to 921 in 1980 (Table 6).

In the main Saskatchewan River at Site 9, mean weekly populations of black fly larvae

attaching to artificial substrates increased from 158 in 1977 to 604 in 1978 (Table 8)

apparently due to downstream drift of larvae from the North and South Saskatchewan Rivers.

There were numerous severe outbreaks along the entire main Saskatchewan River in 1978.

Some outbreaks inflicted losses 100 or more km from the river. Outbreaks were reduced to

tolerable levels in 1979 following initiation of methoxychlor treatments that year at the

confluence (Table 4).

Trends in numbers of invertebrates in benthic samples

Quantitative samples of benthic organisms collected with Surber-type nets each August

provided additional substantial evidence of increases in populations of many taxa and of

non-significant trends in others between 1977 and 1979 or 1977 and 1980 (Tables 11 to 17).

Details of living habits and trophic relationships are shown in Table 5.

Populations of crustaceans in four of the six locations (all three rivers) attained peak

abundance in either 1979 or 1980 (P<0.01 in three sites). In only one location (Site 3, north

side, in the effluent path of a pulp mill located nine km upstream from the sampling site) did

crustaceans decline in abundance after 1977 (P<0.05). Ostracods comprised about 80 percent

of the total populations; copepods and cladocerans about 20 percent. Conchostracans and

malacostracans {Hyalella sp.) were rare throughout. All could be considered free-swimming

although in lotic waters they would have lived on or in the substratum which may have offered

them some protection from passage of the treated masses of water.

Mean August numbers of larvae of all taxa of Ephemeroptera combined, peaked in all six

locations in 1980. Differences between years were significant in all three locations in the North

Saskatchewan River, highly significant in the South Saskatchewan, but not significant in the

main Saskatchewan River.

Examining data for various families of Ephemeroptera individually, baetids (about 98

percent Baetis nr. pluto) were relatively abundant and peaked in 1980 in at least one location

in each of the three river branches. Differences in means between years were significant only in

the South Saskatchewan River. Heptageniids (about 80 percent Heptagenia spp.), also

relatively abundant in most locations, peaked in 1979 in the North Saskatchewan River (Site 3,

south side) with significant differences between years in Site 3, north side in 1980 and the

North Saskatchewan near the confluence in 1978. In the South Saskatchewan near the

confluence they peaked in 1979, and in the main Saskatchewan River (Site 9) in 1977 (no

Quaest. Ent. 1983, 19 (1,2)

76

Fredeen

TABLE 11. MEAN NUMBERSd) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED ABOUT ONE KM UPRIVER FROM SITE 3 (CECIL

FERRY), NORTH SASKATCHEWAN RIVER, SOUTH SIDE.

<’^Geometric means calculated from log,o (x + 1) values.

(2)In each year a set of five, 645 cm^ samples were collected each week for three consecutive weeks.

(^>Calculated from log,Q (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Aquatic invertebrates in a large Canadian River

77

TABLE 12. MEAN NUMBERSO) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED ABOUT ONE KM UPRIVER FROM SITE 3 (CECIL

FERRY), NORTH SASKATCHEWAN RIVER, NORTH SIDE.

(')Geometric means calculated from log|o (x + 1) values.

each year a set of five, 645 cm^ samples were collected each week for three consecutive weeks.

^^^Calculated from log,Q (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Quaest. Ent. 1983, 19 (1,2)

78

Fredeen

TABLE 13. MEAN NUMBERSU) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED FROM THE NORTH SASKATCHEWAN RIVER NORTH

SIDE, ABOUT ONE KM UPRIVER FROM SITE 4 (THE CONFLUENCE)

('^Geometric means calculated from log,Q (x + 1) values.

<2)In each year a set of five, 645 cm^ samples were collected each week for three consecutive weeks.

<^lCalculated from log,o (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Aquatic invertebrates in a large Canadian River

79

TABLE 14. MEAN NUMBERSd) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED FROM THE SOUTH SASKATCHEWAN RIVER, SOUTH

SIDE, ABOUT TWO KM UPRIVER FROM SITE 4 (THE CONFLUENCE).

^'^Geometric means calculated from log,Q (x + 1) values.

each year a set of five, 645 cm^ samples were collected each week for three consecutive weeks.

^^^Calculated from log,Q (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Quaest. Ent. 1983, 19 (1,2)

80

Fredeen

TABLE 15. MEAN NUMBERSd) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED ABOUT ONE KM UPRIVER FROM SITE 9 (GRONLID

FERRY), MAIN SASKATCHEWAN RIVER, SOUTH SIDE.

<')Geometric means calculated from log,o (x + 1) values.

<2)In each year a set of five, 645 cm^ samples were collected each week for 3 consecutive weeks.

<^)Numbers in brackets indicate numbers of treatments in entire Saskatchewan River System above sampling

site. Unbracketed numbers indicate numbers of treatments in Main Saskatchewan River alone.

(“^^Calculated from log,o (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Aquatic invertebrates in a large Canadian River

81

TABLE 16. MEAN NUMBERS^) OF AQUATIC INVERTEBRATES IN SURBER-NET

SAMPLES(2) COLLECTED ABOUT ONE KM UPRIVER FROM SITE 9 (GRONLID

FERRY), MAIN SASKATCHEWAN RIVER, NORTH SIDE.

^'^Geometric means calculated from log,Q (x + 1) values.

each week a set of five, 645 cm^ samples were collected each week for 3 consecutive weeks.

^^^Numbers in brackets indicate numbers of treatments in entire Saskatchewan River System above

sampling site. Unbracketed numbers indicate numbers of treatments in Main Saskatchewan River alone.

<'*)Calculated from log,o (x + 1) values. Means followed by different letters, differ significantly (P<0.05)

as indicated by Duncan’s New Multiple Range tests.

Quaest. Ent. 1983, 19 (1,2)

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Fredeen

TABLE 17. SUMMARY OF TABLES 11 TO 16, INCLUSIVE, IDENTIFYING THOSE

YEARS WHICH PROVIDED MAXIMUM DENSITIES OF BENTHIC

INVERTEBRATES IN EACH OF THE SIX LOCATIONS IN THE NORTH, SOUTH,

AND MAIN BRANCHES OF THE SASKATCHEWAN RIVERO)

1979 DATA MISSING(3)

Aquatic invertebrates in a large Canadian River

83

Table 17 (continued)

(S) or highly significant (HS).

<2)Samples from this location were collected nine km downstream from a pulp mill (newsprint), directly

in the effluent path.

(^*No samples were collected from Site Nine in 1979.

Quaest. Ent. 1983, 19 (1,2)

84

Fredeen

samples collected there in 1979). Differences between years were not significant in either site.

Tricorythids (almost 100 percent Tricorythodes minutus Traver), the third of three

ephemeropteran families most abundantly represented, peaked in 1980 in all six locations.

Differences between years were highly significant in the South Saskatchewan River and

significant in one location in the main Saskatchewan River.

Larvae of all three of these families are considered to be swimmers, climbers, and dingers,

and feed by collecting and scraping detritus, diatoms, etc. from substrates (Table 5). Fredeen et

al. (1975) showed that methoxychlor was rapidly adsorbed to solids in river water. Thus these

larvae could have been exposed to methoxychlor either or both as a contact or stomach poison.

Despite this they appeared to resist harmful effects.

The less abundant caenids (about 98 percent Caenis tardata McDunnough) and

leptophlebiids (about 100 percent Traverella albert ana (McDunnough)) which possessed

similar living and feeding habits, and filter-feeding siphlonurids (about 100 percent Isonychia

sicca) also peaked in the North Saskatchewan River in one or more locations in 1980, in the

South Saskatchewan in 1978 (leptophlebiids) and 1979 (caenids) with highly significant

differences between years, and in the main Saskatchewan River in 1977 (caenids) and 1980

(leptophlebiids), with differences between years not significant.

Burrowing polymitarcyids (100 percent Ephoron album (Say)) peaked in the North

Saskatchewan River in two locations in 1980, with highly significant differences between years

in one location. They also peaked in the South Saskatchewan River in 1979 and in the main

Saskatchewan River (south side) in 1980.

Mean August populations of Plecoptera, all taxa combined, peaked in five of the six

locations in 1980 (P<0.01 in four locations), and peaked in the sixth location (the South

Saskatchewan River), in 1979. Larvae were scarce however, peaking at 38 or fewer larvae per

m2 of river bed (Table 17). Larvae of Isoperla would not have hatched until after the August

collection dates each year.

Larvae of Odonata and Coleoptera were generally present although rare (absent from some

sites) and significant trends in numbers were not evident.

Immature hemipterans (corixids) also were relatively scarce in most sites but numbers

peaked in two rivers in 1980 and in the main Saskatchewan River in 1977. Samples were not

collected there in 1979. Corixids migrate from surrounding regions to overwinter in the rivers.

Mean numbers of trichopteran larvae, all taxa combined, peaked in one or more locations in

all three rivers in 1980. In two of the three locations in the North Saskatchewan River,

however, peaks occurred in 1979 with differences between years highly significant.

Ninety-eight percent of the trichopterans were hydropsychids which, because of their size and

number, easily comprised the most important portion of the total invertebrate biomass of the

river. Sixty to 90 percent of the hydropsychids were Cheumatopsyche spp. and the the

remainder were mainly Hydropsyche alternans (Walker) and H. confusa. It had been thought

that filter-feeding hydropsychid larvae (Table 5) might have been relatively sensitive to

methoxychlor which is known to readily adsorb to particles suspended in the water (Fredeen et

al., 1975). liowever, population numbers showed no indication of this.

Larvae of three other families of Trichoptera, the Hydroptilidae (Mayatrichia ayama

Mosely), Leptoceridae {Nectopsyche diarina (Ross)), and Brachycentridae (Brachycentrus

ocidentalis Banks), all considered to be herbivores (Table 5), were relatively scarce. In the

North and South Saskatchewan Rivers their populations peaked in either 1979 or 1980. The

one exception was Brachycentridae in the South Saskatchewan, peaking in 1978. In the main

Aquatic invertebrates in a large Canadian River

85

Saskatchewan River they peaked in 1977 or 1978. However, no samples were collected there in

1979, which could have been their peak year as in the other rivers.

Larvae of Diptera, of which about 98 percent were chironomids, peaked in all three benthic

sampling locations in the North Saskatchewan River in 1979, with differences between years

highly significant in two locations. Numbers peaked in the South Saskatchewan in 1980 and in

the main Saskatchewan in 1977 (samples were not collected in 1979) and differences between

years were significant in one North Saskatchewan location.

Of the Chironomidae about 50 to 75 percent of the specimens were species of Chironomini,

a subfamily with many filter-feeders, but otherwise with very diverse living and feeding habits

(Table 5). In the North Saskatchewan River numbers of Chironomini peaked in 1979, and in

the South Saskatchewan in 1980 with significant or highly significant differences between

years in three of the four sampling locations.

Larvae of the less abundant predaceous Tanypodini peaked in the three North

Saskatchewan River locations in 1979, with differences between years significant in one

location, and peaked in the South and main Saskatchewan Rivers in 1977 with non-significant

differences between years.

Larvae of the herbivorous Orthocladiinae peaked in 1979 and 1980 in the North and South

Saskatchewan Rivers at Site 4 with significant differences between years but elsewhere showed

non-significant trends. Larvae of Tanytarsini peaked in 1979 in the North Saskatchewan River

(Site 4) with highly significant differences between years and in the South Saskatchewan (Site

4) in 1980. Elsewhere populations did not trend significantly.

Trends in the relatively small populations of Empididae and Simuliidae were generally

inconclusive but peaked in the North Saskatchewan River in 1977 or 1980, in the South

Saskatchewan River in 1979 or 1980 and in the main Saskatchewan River in 1977 or 1978

(where no benthic samples were collected in 1979).

Mites, whose larvae are parasitic, and adults are predators of various aquatic invertebrates,

peaked in all three locations in the North Saskatchewan River in 1979 (with significant or

highly significant differences between years in two locations). They peaked in the South

Saskatchewan River in 1980 (with significant differences between years), and in the main

Saskatchewan River in 1980 (with non-significant differences between years). These trends

provide indirect evidence that numbers of their prey species also may have increased in

abundance during these four years.

Molluscs were scarce in most locations and trends in numbers were not significant.

Thus in general, data from benthic samples collected each August from all three branches of

the Saskatchewan River System indicated that with few exceptions populations of non-simuliid

taxa increased rather than decreased throughout three years of increasingly intensive use of

methoxychlor black fly larvicide followed by a fourth year of less intensive use.

Check-list of taxa in benthic samples

A detailed list of taxa (Table 18) was prepared from benthic samples collected in 1980.

(Samples collected in earlier years were not analyzed in such detail because of inadequate

systematic keys. However, samples from previous years are being retained in the event that

re-examinations in greater detail are required.)

These benthic samples would not have contained species which would have been in the egg

stage in August, particularly some heptageniid mayflies and some plecopterans. But data in

Table 18 show that sufficient other taxa were present in some or all locations sampled to

Quaest. Ent. 1983, 19 (1,2)

86

Fredeen

TABLE 18. LIST OF INVERTEBRATE TAXA COLLECTED IN SURBER-TYPE NETS

FROM SIX SITES IN THE NORTH, SOUTH, AND MAIN BRANCHES OF THE

SASKATCHEWAN RIVER IN SASKATCHEWAN, AUGUST 5, 12, AND 19, 1980(0 (O.

(continued on next page)

Aquatic invertebrates in a large Canadian River

87

Table 18 (continued)

(continued on next page)

Quaest. Ent. 1983, 19 (1,2)

88

Fredeen

Table 18 (continued)

(1) On each of the three dates, five 645 cm2 river bed samples were collected from

each of the six sites. The diameter of the mesh openings in the net was 0.2 mm.

(2) “0” = absent in samples; “X” = present

(3) See Fig. 1 to locate sites on map, and Table 1 for distances.

Aquatic invertebrates in a large Canadian River

89

indicate that a complex invertebrate fauna still existed following four years of relatively

intensive use of methoxychlor black fly larvicide. Included were representatives of widely varied

life cycles, larval activity patterns, and feeding habits (Table 5). Particularly abundant and/or

widespread in 1980 were mayfly larvae (especially the clinging, herbivorous heptageniids,

tricorythids, and leptophlebiids); swimming herbivores (baetids); carnivorous odonatids;

herbivorous stoneflies; filter-feeding, herbivorous or detrivorous hydropsychids; herbivorous

hydroptilids, brachycentrids, and leptocerids; filter-feeding simuliids and chironomids (many

taxa); herbivorous and predatory chironomids (tanypodines and orthocladiines); parasitic mites

and various molluscs.

Included also were species with life cycles as short as four to five weeks (especially S.

luggeri or others with larvae continuously abundant throughout the summer (heptageniids and

other mayfly taxa, hydropsychids and many chironomids)), and species with life cycles lasting

one or more years (pteronarcid and perlid stoneflies). With repeated insecticide treatments

throughout four consecutive summers one might have expected that species with relatively

short life cycles such as simuliids would have become relatively more abundant in comparison

with species which spent longer periods as larvae. However, there was no evidence of this

occurring and in fact simuliid larvae, excluding S. vittatum, became less abundant in successive

years and differences between years were highly significant, at least in mid-river sites (Table

9), whereas species which spent much of each summer as larvae, such as many

ephemeropterans, trichopterans, and chironomids, either peaked in the third or fourth years of

the tests or trended insignificantly between years (Tables 6, 7, and 8).

Need for Control of Black Flies at Sites of Breeding

Throughout four consecutive years of tests with methoxychlor as a black fly larvicide in the

Saskatchewan River system in Saskatchewan, populations of larvae of the problem species of

black fly, S. luggeri declined progressively to their lowest levels (P<0.01) in the final year

(Table 9). Residents in east-central Saskatchewan experienced major outbreaks in 1976, 1977

and 1978 but almost complete relief from black fly outbreaks during 1979 and 1980. Maximum

use of larvicide occurred in 1979, both as regards numbers of sites and numbers of injections.

Also, in 1979, the main Saskatchewan River below the confluence of the two branches was

injected for the first time. Greatly reduced populations of S. luggeri larvae throughout 1980

allowed reductions in numbers of treatments that year (Tables 2, 3, 4).

Provincial and Federal Departments of the Environment are rightfully concerned about the

invertebrate fauna of the Saskatchewan River when renewals of permits for larviciding are

requested each year. Data presented in this paper should help to rationalize those concerns. At

the same time there has to be concern for human environments in some one to three million ha

of land sometimes blanketed by widespread, prolonged outbreaks of S. luggeri. Not only were

people unable to work out-of-doors during outbreaks in 1976 through 1978 unless well

protected, but also livestock owners incurred considerable financial losses (Fredeen, 1981).

Despite many years of research towards protecting livestock individually from black fly

attacks, both in this country and elsewhere, there still are no practical or economical means of

protection available, especially for animals in large pastures. Within the region affected by

outbreaks of S. luggeri in east-central Saskatchewan (up to three million ha in 1978) there are

estimated to be more than 250,000 beef and dairy cattle in community and private pastures

each summer. Most of these animals could not be regularly rounded up for insecticide

treatments, even if durable, registered insecticides were available, because management and

Quaest. Ent. 1983, 19 (1,2)

90

Fredeen

chemical costs would be prohibitive. Labour and chemical costs are more efficiently used if

infestations of black fly larvae are reduced when in their relatively limited river environments,

rather than by attempting to reduce numbers of adults that have dispersed widely into the

countryside. Furthermore, dispersals of adult black flies are affected by varying weather

conditions which frequently catch livestock producers unprepared.

Thus at this time, black fly larviciding with methoxychlor remains the most practical

method for minimizing damage from widespread outbreaks of S. luggeri from the

Saskatchewan River, and this study shows that larviciding throughout a four-year period was

relatively harmless to the natural environment of that river in the long term.

Alternatively one might consider reducing numbers of larvae of S. luggeri by reversing

whatever ecological trends in the river system may have encouraged invasion by populations of

this species in the early 1970’s, and proliferation within a few years to the extent that

widespread outbreaks of economic proportions were possible. Other haematophilic species of

black flies such as S. meridionale, S. venustum, and S. tuberosum recently have become

established in this river and may also require control in future years (Fredeen, 1981). Thus one

might consider attempting to reverse recent ecological trends but such a “biological control”

method would also affect the present day environments of non-simuliid taxa.

CONCLUSIONS

Trends in numbers of non-simuliid invertebrates throughout 1977 to 1980 in the North,

South, and main branches of the Saskatchewan River in Saskatchewan were generally directly

related to the numbers of methoxychlor larvicide injections each year. Thus when total

numbers of larvicide injections were increased from one in 1976, to six in 1977, seven in 1978,

19 in 1979, and five in 1980, densities of most non-simuliid taxa in benthic samples also peaked

either in 1979 or 1980, many significantly so (Table 17). Mean numbers of total non-simuliids

in all mid-river sites combined, also peaked in 1979 (P<0.01) (Table 9). Thus numbers of

non-simuliid aquatic invertebrates showed similar upward trends during these years whether

collected with artificial substrates from mid-river, or with Surber-type nets from benthic sites

along the river margins.

Repopulation of treated sections was assured by drift of invertebrates downstream from

extensive untreated sections, as well as by hatching of eggs and by movement of larvae from

protected niches within each treated section. The abundant and complex benthic fauna

surviving along the river margins presumably served as a rich source of emigrants for

depopulated areas.

Some observed increases appear also to have been due to downstream displacements of

larvae which had been stimulated to release during larvicide treatments.

Qualitatively also, the non-simuliid fauna appeared healthy at the conclusion of tests in

1980 (Table 18). Benthic samples collected from all three river branches throughout August

that year contained a variety of species representing normal ranges of activity patterns, feeding

habits, and life cycles. Although data in Tables 6 through 17 relate mainly to suprageneric

taxa, qualitative analyses showed each of these taxa to be represented by relatively few

dominant species throughout the four years of tests. Thus upward trends in numbers

throughout two or more of the four years of treatments were caused mainly by increases in the

numbers of relatively few species.

Aquatic invertebrates in a large Canadian River

91

The fact that significant upward trends in numbers of most suprageneric taxa occurred

during three years of increasingly intensive use of methoxychlor and a fourth year of less

intensive use suggests that long-term effects of this larvicide on numbers of most non-simuliids

were essentially neutral when compared with effects of natural ecological processes.

In general, this study of trends in quantities and qualities of invertebrate taxa inhabiting the

three branches of the Saskatchewan River throughout four years of injections of methoxychlor

larvicide indicates that S. luggeri may be successfully controlled in a limited portion of the

Saskatchewan River without permanently harming major non-simuliid taxa. The results

support data from an earlier study (Fredeen, 1975) which showed relatively rapid repopulation

of a 160-km section of the Saskatchewan River following a single injection of methoxychlor.

ACKNOWLEDGEMENTS

I am greatly indebted to Richard and Lesley Moffatt whose steady persistence and careful

work throughout the past five years helped to bring this huge project to completion. I am also

indebted to D. Lehmkuhl, Peter Mason, Eric Whiting, and Douglas Smith for providing

systematic expertise whenever required. Frequent consultations with Dr. J. R. Jowsey

throughout the project were also very helpful. David Spurr prepared computer programs for the

statistical analyses and Drs. Alfred Arthur and FI. T. Fredeen provided suggestions for

preparation of the manuscript. I thank Dr. G. Seguin-Swartz for the French version of the

abstract.

REFERENCES

Anderson, R.L, and D.L. DeFoe. 1980. Toxicity and bioaccumulation of endrin and

methoxychlor in aquatic invertebrates and fish. Environmental Pollution 22:1 1 1-121.

Butler, G.L., T.R. Deason, and J.C. O’Kelley. 1975. Loss of five pesticides from cultures of 21

planktonic algae. Bulletin of Environmental Contamination and Toxicology.

13:149-152.

Corbet, P.S., F. Schmid, and C.L. Augustin. 1966. The Trichoptera of St. Helen’s Island,

Montreal. I: The species present and their relative abundance at light. The Canadian

Entomologist 98:1284-1298.

Cummins, K.W. 1973. Trophic relations of aquatic insects. Annual Review of Entomology.

18:183-206.

Cummins, K.W. and G.F. Edmunds. 1978. Summary of ecological and distributional data for

Ephemoroptera (mayflies);77-80. In An introduction to aquatic insects of North

America. Kendall/Hunt Publishing Company, Dubuque.

Cummins, K.W. and P.P. Harper. 1978. Summary of ecological and distributional data for

Plecoptera (stoneflies);! 15-1 18. Ibid.

Cummins, K.W. and G.B. Wiggins. 1978. Summary of ecological and distributional data for

Trichoptera (caddisflies); 175-1 85. Ibid.

Cummins, K.W. and W.P. Coffman. 1978. Summary of ecological and distributional data for

Chironomidae (Diptera);370-376. Ibid.

Edwards, W.M. and B.L. Glass. 1971. Methoxychlor and 2,4,5-T in lysimeter percolation and

runoff water. Bulletin of Environmental Contamination and Toxicology. 6:81-84.

Environment Canada. 1978. Surface water data, Saskatchewan, 1977. Minister of Supply and

Services, Canada; 188 pages.

Environment Canada. 1979. Surface water data, Saskatchewan, 1978. Ibid. 190 pages.

Quaest. Ent. 1983, 19 (1,2)

92

Fredeen

Environment Canada. 1980. (a) Surface water data, Saskatchewan, 1979. Ibid. 192 pages.

Environment Canada. 1980. (b) Historical streamflow summary, Saskatchewan, to 1979. Ibid.

339 pages.

Environment Canada. 1981. Surface water data, Saskatchewan, 1980. Ibid. 188 pages.

Fredeen, F.J.H. 1969. Outbreaks of the black fly Simulium arcticum Malloch in Alberta.

Quaestiones Entomologicae 5:341-372.

Fredeen, F.J.H. 1972. Reactions of the larvae of three rheophilic species of Trichoptera to

selected insecticides. Canadian Entomologist 104:945-953.

Fredeen, F.J.H. 1974. Tests with single injections of methoxychlor black fly (Diptera:

Simuliidae) larvicides in large rivers. Ibid. 106:285-305.

Fredeen, F.J.H. 1975. Effects of a single injection of methoxychlor black fly larvicide on insect

larvae in a 161-km (100 mile) section of the North Saskatchewan River. Ibid.

107:807-817.

Fredeen, F.J.H. 1977. Some recent changes in black fly populations in the Saskatchewan River

system in Western Canada coinciding with the development of reservoirs. Canadian

Water Resources Journal 2:90-102.

Fredeen, F.J.H. 1981. Keys to the black flies (Simuliidae) of the Saskatchewan River in

Saskatchewan. Quaestiones Entomologicae 17:189-210.

Fredeen, F.J.H., J.G. Saha and M.H. Baiba. 1975. Residues of methoxychlor and other

chlorinated hydrocarbons in water, sand and selected fauna following injections of

methoxychlor black fly larvicide into the Saskatchewan River, 1972. Pesticides

Monitoring Journal 8:241-246.

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artificial substrates. Quaestiones Entomologicae 14:41 1-431.

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Mississippi River. Iowa State University, Research Bulletin 482, Ames, Iowa.

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Mississippi River. Iowa State University, Research Bulletin 483, Ames, Iowa.

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America. 441 pp. Kendall/Hunt Publishing Company, Dubuque, Iowa.

Muirhead-Thomson, R.C. 1973. Laboratory evaluation of pesticide impact on stream

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Munroe, E.G. 1951. Pest Trichoptera at Fort Erie, Ontario. Canadian Entomologist 83:69-72.

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Ontario. Canadian Entomologist 84:103-107.

Sebastien, R.J., and W.L. Lockhart. 1981. The influence of formulation of toxicity and

availability of a pesticide (methoxychlor) to black fly larvae (Diptera:Simuliidae),

some non-target aquatic insects and fish. Ibid. 1 13:281-293.

Ward, H.B. and G.C. WI e. (Editor, W.T. Edmondson). 1959. Fresh-water biology. 1248

pp. John Wiley and Sons, New York, U.S.A.

Wiggins, G.B. 1977. Larvae of North American caddisfly Genera (Trichoptera). 401 pp.

University of Toronto Press, Toronto, Canada.

CYMINDINE LEBIINI OF AUTHORS: REDEFINITION AND RECLASSIFICATION OF

GENERA (CLEOPTERA: CARABIDAE)

George E. Ball

Department of Entomology

University of Alberta

Edmonton, Alberta

Canada T6G 2E3

Quaestiones Entomologicae

19:93-216 1983

Gerald J. Hilchie

Department of Entomology

University of Alberta

Edmonton, Alberta

Canada T6G 2E3

ABSTRACT

Based on examination of character states of adults (in particular, sclerites of the

ovipositor) of a limited sample of taxa, heretofore included in the lebiine subtribe Cymindina

(= Tribes Cymindina and Pseudomasoreini, or Subfamily Cyminditae of authors), the

following tribes and subtribes were found to be represented: Tribe Pterostichini, subtribe

Platynina; Tribe Lachnophorini; Tribe Lebiini, Subtribes Pericalina, Apenina, Cymindina,

Calleidina, and Dromiina; and Tribe Zuphiini. The South African Anarmosta Peringuey,

1896 (= Euplynes Schmidt-Goebel, 1846) is confirmed as a platynine. The New World

tropical and subtropical Eucaerus LeConte, 1853 and Lachnaces Bates, 1872, are included in

the Eucaerus complex, and transferred to the Lachnophorini. Eucaerus and Lachnaces are

regarded as congeneric subgenera (new rank). Also included in the Eucaerus complex are the

Neotropical genera Asklepia Liebke, 1938, and Phaedrusium Liebke, 1951. Transferred to the

subtribe Pericalina are the Afrotropical (South African) Leptosarcus Peringuey, 1896, and

(East African montane) Selenoritus Alluaud, 1917, the latter included as a subgenus of

Thyreopterus Dejean, 1831 (new rank). Transferred to the subtribe Apenina are three genera:

the New World Apenes LeConte, 1851, with subgenera Apenes sensu stricto Malisus

Motschulsky, 1864), and Sphalera Chaudoir, 1875 (= Didymochaeta Chaudoir, 1875, new

synonymy); Palaearctic Trymosternus Chaudoir, 1873; and the Old World Tropical

Cymindoidea Castelnau, 1832. The latter genus includes as subgenera Cymindoidea (sensu

stricto), Platytarus Fairmaire, 1850 (new rank), and Habutarus new subgenus (generitype

Nototarus papua Darlington, 1968). The subtribe Cymindina includes the new Oriental genus

Ceylonitarus (generitype C. ceylonicus, new species, with type locality vicinity of Mannar, Sri

Lanka), the Megagean Cymindis Latreille, 1806, and the Afrotropical-western Palaearctic

Hystrichopus Boheman, 1848. The genus Cymindis includes four subgenera (new rank):

Oriental Taridius Chaudoir, 1875; Nearctic-Neotropical Pinacodera Schaum, 1857;

Afrotropical-Oriental Afrotarus Jeannel, 1949; and Holarctic Cymindis sensu stricto.

Hystrichopus includes four subgenera (new rank): Madagascan Assadecma Basilewsky, 1982;

Afrotropical-Palaearctic Pseudomasoreus Desbrochers des Loges, 1904; Afrotropical

Hystrichopus sensu stricto; and Afrotropical Plagiopyga Boheman, 1848. Transferred to the

94

Ball and Hilchie

subtribe Calleidina are the Palaearctic- Old World Tropical- Australian Anomotarus

Chaudoir, 1875, and the Australian Trigonothops MacLeay, 1864. Transfer of Anomotarus

renders the names Calleidina and Anomotarina synonyms; the latter name is junior.

Anomotarus includes three subgenera: Palaearctic- Old World Tropical- Australian

Anomotarus sensu stricto; Australian Nototarus Chaudoir, 1875, new rank f= Lithostrotus

Blackburn, 1894, new synonymy); and Afrotropical Dromiotes Jeannel, 1949 {= Cephalotarus

Mateu, 1973). Trigonothops includes five subgenera (new rank): Trigonothops sensu stricto;

Phloeocarabus MacLeay, 1871; Diabaticus Bates, 1878; Abaditicus new subgenus (generitype

Diabaticus collaris Blackburn, 1901); and Speotarus Moore, 1964. Transferred to the

Dromiina is the Afrotropical (South African) genus Metaxymorphus Chaudoir, 1873,

including as subgenera (new rank): Metaxymorphus sensu stricto; Periphobus Peringuey, 1896;

and Callidomorphus Peringuey, 1896. Inclusion of the South African Syndetus Peringuey,

1896 (= Coptoptera Chaudoir, 1837) in the Dromiina is comfirmed. The Old World Agastus

Schmidt-Goebel, 1846 is transferred to the tribe Zuphiini. Also included in the Dromiina are

the tribes Lichnasthenini and Singilini.

The Madagascan genera Thysanotus Chaudoir, 1837, Antimerina Alluaud, 1897, and

Madecassina Jeannel, 1949 (formally tribe Thysanotini, subfamily Calleiditae) are placed in

the subtribe Pericalina, with the name Thysanotini becoming thereby a junior synonym of the

name Pericalina.

The name Lachnaces sericeus Bates, 1872 is changed to Eucaerus fLachnacesj sericeus,

thereby becoming a junior secondary homonym of Eucaerus fsensu strictoj sericeus Bates,

1871. Eucaerus sericatus is proposed as a name for the junior homonym.

The nominal species Cymindis (Taridius) stevensi (Andrewes, 1923) is expanded to include

as subspecies C. s. nilgirica (Andrewes, 1935), C. s. andrewesi (van Emden, 1937), and C. s.

stevensi sensu stricto. Taridius niger Andrewes, 1935 is transferred to subgenus Afrotarus

Jeannel. New species of Hystrichopus (subgenus Pseudomasoreus) are described, based on

material from the Union of South Africa: H. (P) reticulatus (type locality- Cape Province,

Clanwilliam District, Sederburg); H. (P.) basilewskyi (type locality- Cape Province,

Swellendam Distr., Grootvaderbos); H. (P.) thoracicus (type locality- Grahamstown); and H.

(P.) mateui (type locality Natal, Malvern). A new species of Trigonothops is described: T.

(Abaditicus) meyeri (type locality- AUSTRALIA, Victoria, Nunniong Plateau, Woodhouse

Creek).

RESUME

L’examen des caracteres des adultes (en particulier des sclerites de I’ovipositeur), realise sur un echantillon limite de

taxons jusqu’ici inclus dans la sous-tribu lebiienne des Cymindina (= tribus des Cymindina et des Pseudomasoreini, ou

sous-famille des Cyminditae de certains auteurs), revele que les tribus et sous-tribus suivantes y sont representees: tribu

des Platynina; tribu des Lachnophorini; tribu des Lebiini, sous-tribus des Pericalina, Apenina, Cynindina, Calleidina et

Dromiina: et tribu des Zuphiini. Cette etude confirme en outre que le genre sud-african Anarmosta Peringuey, 1896 (=

Euplynes Schmidt-Goebel, 1846) est bien platyninien. Les genres Eucaerus LeConte, 1853 et Lachnaces Bates, 1872, des

tegions tropicales et subtropicales du Nouveau Monde, sont inclus dans le complexe des Eucaerus, et transferes dans les

Lachnophorini. Eucaerus et Lachnaces sont consideres comme des sous-genres congeneriques (nouveau rang). Les genres

neotropicaux Asklepia Liebke, 1938 et Phaedrusium Liebke, 1951 sont egalement inclus dans le complexe Eucaerus.

Leptosarcus Peringuey, 1896, du sud de I’Afrique, et Selenoritus Alluaud, 1917, des montagnes est-africaines [ce dernier

etant considere comme un sous-genre de Thyreopterus Dejean, 1831 (nouveau rang)], sont transferes dans la sous-tribu

des Pericalina. Trois genres sont transferes dans la sous tribu des Apenina: Apenes LeConte, 1851, du Nouveau Monde,

comprenant les sous-genres Apenes sensu stricto ('= Malisus Motschulsky, 1864), et Sphalera Chaudoir, 1875 (=

Didymochaeta Chaudoir, 1875, synonyme nouveau); Trymosternus Chaudoir, 1873, de ;’Eurasie; et Cymindoidea

Castelnau, 1832, des tropiques de I’Ancien Monde. Ce dernier genre comprend les souis-genres Cymindoides fsensu

Cymindine Lebiini of Authors

95

stricto, Platytarus Fairmaire, 1850 (nouveau rang), et Habutarus, nouveau genre (genotype Nototarus papua

Darlington, 1968). La sous-tribu des Cymindina comprend un nouveau genre de la region orientale, Ceylonitarus

(genotype C. ceylonicus, nouvelle espece, localite du type situee dans les environs de Mannar, Sri Lanka), Cymindis

Latrielle, 1806, reparti en Amerique du Nord, Eurasie et Afrique, et Hysteichopus Boheman, 1848, de I’Afrique

tropicale et de la partie occidentale de I'Eurasie. Le genre Cymindis inclut quatre sous-genres (nouveau rang): Taridius

Chaudoir, 1875, de la region orientale; Pinacodera Schaum, 1857, des regions nearctique et neotropicale; Afrotarus

Jeannel, 1949, des regions orientale et afrotropicale; et Cymindis sensu stricto de la region holarctique. Hystrichopus

comprend quatre sous-genres (nouveau rang): Assadecma Basilewsky, 1982, de Madagascar; Pseudomasoreus

Desbrochers des Loges, 1904, des regions palearctique et afrootropicale; Hystrichopus sensu stricto, de la region

afrotropicale; et Plagiopyga Boheman, 1848, aussi de I’Afrique tropicale. Anomotarus Chaudoir, 1875, des regions

palearctique et australienne ainsi que des tropiques de I’Ancien Monde, et Trignothops MacLeay, 1864, d’Australie,

sont transferes dans la sous-tribu des Calleidina. Calleidina et Anomotarina deviennent synonymes d la suite du

transfert ^/’Anomotarus, Anomotarina etant le plus recent des deux. Anomotarus sensu stricto, reparti en Eirasie, dans

les tropiques de I’Ancien Monde et dans la region australienne; Nototarus Chaudoir, 1875, nouveau rang (=

Lithostrotus Blackburn, 1894, nouveau synonyme), d’Australie; et Dromiotes Jeannel, 1949 (= Cephalotarus Mateu,

1973), de lAfrique tropicale. Trigonothops comprend cinq sous-genres (nouveau rang); Trigonothops sensu stricto;

Phloeocarabus MacLeay, 1871; Diabaticus Bates, 1878; Abaditicus nouveau genre (genotype Diabaticus collaris

Blackburn, 1901); et Speotarus Moore, 1964. Metaxymorphus Chaudoir, 1873, de I’Afrique tropicale (sud de I’Afrique)

est transfere dans les Dromiina et inclut les sous-genres (nouveau rang) Metaxymorphus sensu stricto, Periphobus

Peringuey, 1896, et Callidomorphys Peringuey, 1896. Cette etude confirme en outre I’inclusion du genre sud-africain

Syndetus Peringuey, 1896 (= Coptoptera Chaudoir, 1837) dans les Dromiina. Agastus Schmidt-Goebel, 1846, de

I’Ancien Monde, est transfere dans la tribu des Zuphiini. Les tribus des Lichnasthenini et des Singilini sont aussi

incluses dans les Dromiina.

Les genres malgaches Thysanotus Chaudoir, 1837, Antimerina Alluaud, 1897, et Madecassina Jeannel, 1949

(formellement, de la tribu des Thysanotini, sous-famille des Calleiditae) sont inclus dans la sous-tribu des Pericalina,

rendant ainsi le nom Thysanotini synonyme recent du mom Pericalina.

Le binome Lachnaces sericeus Bates, 1872 est change en Eucaerus (Lachnaces) sericeus, et devient ainsi homonyme

secondaire recent ^y’Eucaerus (sensu stricto) sericeus Bates, 1871. L’auteur propose Eucaerus sericatus comme

remplacement de Thomonyme recent.

La signification de I’espece nominale Cymindis (Taridius) stevensi Andrewes, 1923) est elargie piur inclure les

sous-especes C. s. nilgirica (Andrewes, 1935), C. s. andrewesi van Emden, 1937), et C. s. stevensi sensu stricto. Taridius

niger Andrewew, 1935 est transfere dans le sous-genre Afrotarus Jeannel. De nouvelles especes ’Hystrichopus

(sous-genre Pseudomasoreus) sont decrites d partir de specimens provenant de TUnion Sud-Africaine; ce sont: H. (P.)

reticulatus (localite du type: province du Cap, district de Clanwilliam, Sederburg); H. (P.) basilewskyi (localite dy

type: province du Cap, district de Swellendam, Grootvaderbos); H. (P.) thoracicus (localite du type: Grahamstown; et

H. (P.) mateui (localite du type: Natal, Malvern). Une nouvelle espece de Trigonothops est decrite; il s’agit de T.

(Abaditicus) meyeri (localite du type: Australie, Victoria, plateau du Nunniong, Woodhouse Creek).

TABLE OF CONTENTS

Introduction 96

Materials and Methods 96

Materials 96

Methods 97

Structures used in Classification 98

Classification 98

Tribe Pterostichini, Subtribe Platynina 99

Tribe Lachnophorini 101

Tribe Lebiini 108

Subtribe Pericalina 116

Subtribe Apenina 120

Subtribe Cymindina 129

Subtribe Calleidina 173

Subtribe Dromiina 196

Tribe Zuphiini 201

Concluding Remarks 202

Quaest. Ent., 1983, 19 (1,2)

96

Ball and Hilchie

Acknowledgements 203

References 204

Index 213

INTRODUCTION

During preparation of a revision of the species of the New World taxon Pinacodera Schaum,

we wished to identify its sister group, and so undertook what was hoped to be a brief review of

the genera that Rene Jeannel and other previous workers had included in the subtribe

Cymindina. That outstanding Japanese student of Carabidae, Akinobu Habu (1967), showed

that details of the ovipositor of adult lebiines were of substantial value in classification. We also

knew that mandibles offered useful and previously unused character states.

Preliminary examination of these structures of adults of a few supposedly cymindine genera

showed striking heterogeneity, so much so that it became evident that the cymindine

assemblage was very likely to be unsatisfactory, at least from a phylogenetic viewpoint. This

realization left us with three choices: to abandon the original goal, and to proceed with an

analysis of Pinacodera without knowing the sister group; or to attempt to locate close relatives

of Pinacodera and leave the rest of the cymindines for another time; or to attempt to sort out

the group by assigning all genera to their proper subtribes, and at the same time, to identify the

sister group of Pinacodera. We chose the last course, and this paper is the result.

At first, we thought that reclassification of the cymindine Lebiini would form the

introductory part of a treatment of Pinacodera, but the introduction grew in volume and

complexity, until it became obvious that inclusion of a detailed treatment of that genus would

appear almost as an appendage. Therefore, revision of the species of Pinacodera will be

published separately.

In the present paper, genera of the Cymindina of authors are briefly characterized on the

basis of features of adults, and assigned to their proper groups. Several subtribes of Lebiini are

characterized. Most genera are treated in cursory fashion, but for some, material was available

for partial revision, and we took advantange of the opportunities thus offered.

This paper is not a revision of the higher classification of the Lebiini. It is more a collection

of notes that ought to be useful for such a revision. Habu (1967) provided the basis for such a

treatment, but structures of many more taxa must be examined in detail, to assess character

systems thought to be of value, and to identify evolutionary trends.

MATERIALS AND METHODS

Materials

Several hundred lebiine adults were examined, representing described cymindine genera. A

few taxa were represented in the Strickland Museum, University of Alberta (UASM), but most

specimens were borrowed. Listed below, with abbreviations used in the text, are names and

addresses of the lending institutions.

BMNH Department of Entomology, British Museum (Natural History), Cromwell

Road, London, England, SW7 5BD.

CAS Department of Entomology, California Academy of Sciences, Golden Gate

Park, San Francisco, California U.S.A., 94118.

Cymindine Lebiini of Authors

97

CSIRO Commonwealth Scientific and Industrial Research Organization, Division of

Entomology, Black Mountain, Canberra City, ACT 2601, Australia

IRSB Section d’Entomologie, Institut Royal des Sciences Naturelles du Belgique,

Bruxelles 4, Rue Vautier 31, Belgium.

MACT Musee Royal de I’Afrique Centrale, B- 1980, Tervuren, Belgique.

MCZ Museum of Comparative Zoology, Harvard University Cambridge,

Massachusetts, U.S.A. 02138.

MNHP Entomologie, Museum National d’Histoire Naturelle, Paris (Ve), France.

SAMC South African Museum, P.O. Box 61, Cape Town, South Africa.

USNM Department of Entomology, United States National Museum of Natural

History, Smithsonian Institution, Washington, D.C., U.S.A. 20560.

ZSIC Zoological Survey of India, 34 Chittaranjan Avenue, Calcutta, 700 012 India.

Methods

Because of the nature of this study, most taxa were represented by few specimens.

Therefore, no attempt was made to assess range of variation of character states studied, and

few specimens of each taxon were dissected or measured. In general, however, characters used

tend to be stable intraspecifically.

Taxonomic principles, criteria for ranking taxa, and general working methods were the same

as those previously described (Ball, 1975 and 1978, and Allen and Ball, 1980), and are not

repeated here. However, if we have erred in taxonomic judgement, it is in the direction of

lumping rather than splitting, by emphasis of similarities that we felt are likely to represent

close phylogenetic relationship, rather than emphasis of differences that, although they might

be numerous, seem the sort of features that might change rapidly.

Genitalia and other small structures were preserved in glycerine, in microvials, pinned

beneath the specimens from which they were removed.

Measurements made with a Wild M5 stereobinocular microscope, at 25X or 50X, are as

follows, and are expressed in the text by these abbreviations:

HI- length of head, measured on left side, from base of left mandible to posterior

margin of compound eye;

Hw- maximum transverse distance across head, including eyes;

Vwm- minimum transverse distance across vertex (used for specimens with markedly

constricted head, posteriorly);

PI- length of pronotum, measured along mid-line, from base to apex;

PwB- width of pronotum, at base;

Pwm- maximum width of pronotum;

MESl

(and w)- length of metepisternum, measured along lateral margin; (width of

metepisternum, measured along basal margin);

El- length of longer elytron (if elytra of a single specimen were unequal) from basal

ridge to apex.

Size was expressed in the text as the sum of HI, PI, and El, and referred to as Standardized

Body Length, or SBL. Other measurements were used to form ratios which seemed to provide

adequate diagnostic features for differentiation among members of some taxa.

Quaest. Ent., 1983, 19 (1,2)

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Ball and Hilchie

For photographs of some structures, a Stereo Electron Microsope was used, Cambridge

Model SI 50. Specimens were cleaned, using a sonicator, and were gold-coated.

STRUCTURES USED IN CLASSIFICATION

All of the features used are standard for carabids, especially lebiines. Nonetheless, attention

is drawn here to terms that have yet to be stabilized in the carabid literature for various

structures.

For micro-units of surface sculpture bounded by lines of microsculpture, we use “sculpticell”

(Allen and Ball, 1980: 486); for elytral stria, “interneur” (Erwin, 1974: 3-5). For abdominal

sterna, Roman numerals are used, with first visible sternum being II, and the last one that is

not normally retracted, VII.

The median lobe of the male genitalia is classified depending upon position of the apical

orifice: anopic, if dorsal; catopic, if ventral (Jeannel, 1949: 878). For a discussion of the

significance of catopy see Jeannel (1955: 82-86). The word “hemiopic” is used for median lobes

in which the apical orifice is more lateral than it is dorsal or ventral (Ball and Shpeley, in

press).

Sclerites of the ovipositor are named according to Tanner (1927), with modifications

proposed by Noonan (1973), and Ball and Shpeley (in press). Thus, the terminal sclerite of the

ovipositor is “stylomere 2”, abbreviated S2. Terms used for surfaces are those proposed by Ball

and Shpeley (in press), based on orientation of surfaces in the extended position.

CLASSIFICATION

The cymindine genera of authors represent one subtribe of Pterostichini, the Lachnophorini,

five subtribes of Lebiini, and the Zuphiini. As a guide to the text, we list by name these

supraspecific taxa, as well as two that are new, and several not included in the Cymindina of

authors, but related more or less directly to the general subject matter of this study.

Tribe PTEROSTICHINI

Subtribe PLATYNINA

Anarmosta Peringuey, 1896 (junior subjective synonym of Euplynes

Schmidt-Goebel, 1846)

Tribe LACHNOPHORINI

Eucaerus LeConte, 1853

Lachnaces Bates, 1872

Asklepia Liebke, 1938

Phaedrusium Liebke, 1951

Tribe LEBIINI

Subtribe PERICALINA (including THYSANOTINI)

Thysanotus Chaudoir, 1837

Antimerina Alluaud, 1897

Madecassina Jeannel, 1949

Selenoritus Alluaud, 1917

Thyreopterinus Alluaud, 1932

Thyreopterus Dejean, 1831

Cymindine Lebiini of Authors

99

Leptosarcus Peringuey, 1896

Subtribe APENINA

Apenes LeConte, 1851

Malisus Motschulsky, 1864

Sphalera Chaudoir, 1875

Didymochaeta Chaudoir, 1875

Trymosternus Chaudoir, 1873

Cymindoidea Castelanu, 1832

Platytarus Fairmaire, 1850

Habutarus, new subgenus

Subtribe CYMINDINA

Ceylonitarus, new genus

Taridius Chaudoir, 1875

Pinacodera Schaum, 1857

Afrotarus Jeannel, 1949

Cymindis \.2iirQ\\\Q, 1806

Assadecma Basilewsky, 1982

Pseudomasoreus Desbrochers des Loges, 1 904

Hystrichopus Boheman, 1848

Plagiopyga Boheman, 1848

Subtribe CALLEIDINA

Trigonothops Macleay, 1864

Phloeocarabus Macleay, 1871

Diabaticus Bates, 1878

Abaditicus, new subgenus

Speotarus Moore, 1964

Subtribe DROMIINA (including LICHNASTHENINI and SINGILINI

Metaxymorphus Chaudoir, 1873

Periphobus Peringuey, 1896

Callidomorphus Peringuey, 1896

Syndetus Peringuey, 1896 (junior subjective synonym of Coptoptera Chaudoir,

1837)

Tribe ZUPHIINI

Agastus Schmidt-Goebel, 1846

Details about these subtribes and genus-group taxa are provided below.

TRIBE PTEROSTICHINI, SUBTRIBE PLATYNINA

Genus Euplynes Schmidt-Goebel

Figs. 1 and 2

Euplynes Schmidt-Goebel, 1846: 52. GENERITYPE: Euplynes cyanipennis Schmidt-Goebel, 1846: 52 (monotypy).-

Burgeon, 1937: 397.- Jeannel, 1949: 611.- Mateu, 1974: 487-506.- Habu, 1978: 292.

Euplenes Darlington, 1952: 122.

Xatis Fairmaire, 1901: 125. GENERITYPE: Xatis nigripes Fairmaire, 1901: 125 (monotypy).- Jeannel, 1949: 611.-

Habu, 1978: 294.

Anarmosta Peringuey, 1896: 221. GENERITYPE: Anarmosta dispar Peringuey, 1896: 222. { = Euplynes callidoides

Chaudoir, 1878) (monotypy).- Mateu, 1974: 487.

Quaest. Ent., 1983, 19 (1,2)

Figs. 1 and 2. Photographs of Platynina, Euplynes callidoides Chaudoir (= Anarmosta dispar Peringuey).— Fig. 1:

habitus, dorsal aspect (SBL -9.79 mm). Fig. 2: SEM photograph of ovipositor, right stylomeres— A, lateral aspect; B,

medial aspect; C, apico-ventral aspect. Scale bars = 50 jam. Legend: a, lateral ensiform seta; b, medial ensiform seta; c,

sensory furrow peg; d, nematoid seta; SI, stylomere 1; S2, stylomere 2.

Cymindine Lebiini of Authors

101

Notes about types and synonymy. — Although we have not seen type material, we have

studied three specimens from the Peringuey collection (SAMC) from the type locality of

Salisbury, and labelled as follows: male, Salisbury, Rhodesia, 17.1.11, J.A. O’Neill; female,

Salisbury, 11.2.18; female, Salisbury, 3.11.1914, J. O’Neil. Additionally, each specimen bears:

two determination labels (Anarmosta dispar Per.; and Euplynes dispar Pering. det. Ball, ’80);

and a museum label (SAMC). Mr. V. Whitehead, of the South African Museum, advised us

that these were the only specimens available of this species in the Peringuey collection. The

features of these specimens fit those provided in the original description. Fig. 1 illustrates the

habitus of E. callidoides Chaudoir.

It seems difficult to believe that Peringuey would have placed a typical platynine among the

Lebiini. However, there are some clues about how such an error could be made. First, his

diagnosis of the “Lebiides” does not exclude specimens with approximately normal elytral

apices (“...or very deeply sinuate behind...”). Second, in the key to genera of “Cymindidae”

(included in the “Lebiides”), Peringuey gave the name ""Haplopeza'" following the singlet in

which Anarmosta runs out, and the former name is not listed again. It seems likely that he

originally regarded the specimen of A. dispar as belonging to Haplopeza, realizing at a later

date (possibly when the manuscript was in press) that this was incorrect. Haplopeza, however,

is a platynine. From this, we infer that A. dispar, although not a species of Haplopeza, is a

platynine. We feel confident that the specimens labelled Anarmosta dispar Peringuey are

indeed members of that nominal species. This is the same conclusion that Straneo (1943: 58)

reached.

The above comments are not made to criticize Peringuey. Rather, they illustrate the

difficulties that our predecessors had in distinguishing among lebiines and platynines, and

especially some of the tropical members of these groups. As a further example of the problem.

Bates (1883:158) suggested that Euplynes might be related to Leptotrachelus .

Figs. 2A-C illustrate the highly distinctive stylomere 2 of the ovipositor of E. dispar, with its

dorso-lateral row of thick spines, and the well developed basal lobe. We think that it might be a

generic character state for Euplynes. Habu’s illustrations (1978: 293-295, Figs. 590-592a) of

Oriental- eastern Palaearctic females are about the same as our Fig. 2. Jeannel (1949: 611)

suggests that the African genus Haplopeza Boheman is related to Euplynes.

Mateu (1974) revised the African species of Euplynes.

Tribe LACHNOPHORINI

To this tribe, four genus-group taxa are assigned: Eucaerus LeConte, Lachnaces Bates,

Asklepia Liebke, and Phaedrusium Liebke. We have seen representatives of only the first two

groups. T. L. Erwin (personal communcation) suggested that the latter two groups should be

included, also. Figures provided by Reichardt (1974: 178, Figs. 1, and 3-7) confirm that

Asklepia is indeed like Eucaerus, and the original description of Phaedrusium (Liebke, 1951:

240-241) includes mention of character states that are Eucaerus- like.

The marked similarity of adults of Lachnaces and Eucaerus in several features is taken as

evidence of very close relationship of these taxa. Therefore, we combine them as subgenera of a

single genus. We believe that re-examination of specimens of Asklepia and Phaedrusium will

show that these groups should be included in Eucaerus, as well.

Reichardt (1974: 178) transferred Asklepia Liebke from the Colliurini to the

Lachnophorini, and Phaedrusium was compared with lachnophorines {Lachnophorus and

Quaest. Ent., 1983, 19 (1,2)

102

Ball and Hilchie

Calybe by Liebke (1951: 241), though he included the genus in the Lebiini. Bates (1871: 77)

noted both lachnophorine and lebiine affinities of Eucaerus. Horn (1881: 155) commented

about the lachnophorine affinities of Eucaerus, referring to it as “an osculant form” between

that group and the Lebiini. He decided, nonetheless, that Eucaerus was a lebiine, a view that

was accepted by subsequent cataloguers and American workers (see Ball, 1960: 162, and

Reichardt, 1977:444).

Terry L. Erwin (personal communication) suggested that this complex belonged in the

Lachnophorini, and we place it there on the basis of: terminal palpomeres with acuminate tips

(Figs. 10 and 12); mandibles of same form (details provided in description of Eucaerus); elytral

apices subtruncate; wings with oblongum cell reduced (stalked), wedge cell absent; stylomere 1

of ovipositor with terminal row of setae, stylomere 2 of plesiotypic form and setation (Figs. 17

and 18). Form of palpomeres is autapotypic. Details of wing venation are also apotypic, but

could have been independently acquired by reduction. Mandibles are probably a mixture of

symplesiotypic and autapotypic features. We cannot sort out the details at this time. All

antennomeres of Eucaerus {sensu stricto) and Asklepia adults have a vestiture of short setae,

like antennomeres 4-11. Antennomeres 1-3 of Phaedrusium adults and antennomere 1 of

subgenus Lachnaces adults are without such vestiture, contrasting with antennomeres 4-11.

According to Reichardt (1977: 413), the Lachnophorini (excluding Anchonoderus Reiche)

is “A weakly characterized tribe of still uncertain position and constitution”. He provided an

account of the taxonomic history of the group (1977: 406 and 413), which has been treated as

an independent tribe near the Perigonini (with or without Anchonoderus), or as a subtribe of

the Pterostichini. Liebherr (MS) presents evidence based on structural features of larvae and

adults, showing clear lebiomorph affinities of lachnophorines, and this is our basis for ranking

this group (including Anchonoderus) as a tribe apart from the Pterostichini, and placing it in

the lebiomorph assemblage. Further work might require including in a single tribe the

lachnophorines and lebiines, but this possibility remains to be investigated.

Geographical distribution. — This complex is confined to the tropics and warm temperate

areas of the New World: all four genera are known from South America, but only Eucaerus

ranges north to Middle America and to southeastern Unites States.

Description of the Eucaerus complex. — The following describes range of variation of

selected features useful in recognizing lachnophorine taxa, and for determining their

relationships.

Color. Various, from somber to pale; dorsum all black to combinations of rufous and testaceous, elytra spotted or not;

legs and palpi testaceous; antennae uniformly testaceous, or tricolored, antennomeres 1-3 rufous or piceous, 4-6 black, and

7-1 1 white.

Microsculpture. Various, but generally transverse; some members of Eucaerus with dorsum of head and/or pronotum

with isodiametric meshes and sculpticells convex, surface thus beaded.

Luster. Generally iridescent, or dorsum of head and pronotum dull.

Macrosculpture. Dorsum generally smooth, without constant depressions or swellings, but frontal impressions with

transverse rugulae; ventral surface rather coarsely but sparsely punctate.

Vestiture. Dorsal surface generally glabrous; all antennomeres setose; or antennomeres 1 or antennomeres 1-3 glabrous

except for normal preapical setae; terminal palpomeres densely setose; maxillary palpomere 3 densely setose, palpomere 2

sparsely setose; ventral surface sparsely setose.

Fixed setae. Average for lachnophorine adults: labrum with six long apical setae; head and pronotum with two pairs;

elytron with three setae on interval 3, or in Asklepia strandi adults, with two rows of setae on disc; umbilical series of

about 10-12 setigerous punctures laterally, broadly interrupted medially, penultimate lateral seta in straight line with

antepenultimate and ultimate setae.

Head. Clypeus transverse, anterior margin truncate. Frontal impressions broad and shallow or deep and linear.

Sub-antennal ridge average. Eyes Orbicular, convex, prominent. Antennae average for lachnophorine adults: filiform,

flagellar antennomeres sub-cylindrical distinctly longer than wide; antennomere 2 short, 3 longer than 4.

Cymindine Lebiini of Authors

103

Mouthparts. Labrum with anterior margin truncate. Left and right mandibles about same in overall shape. Scrobes

less than 0.50 total length of mandibles, ventral edge of scrobe curved upward. Left mandible (Figs. 3A, C, 4A, C, 5A,

C) with terebral ridge distinct, extended more than half length of terebra; terebral tooth absent; retinacular ridge

cutting edge; retinacular tooth prominent, cleft, ventral ridge well developed; premolar tooth blunt, small, set off from

posterior part of retinacular ridge by indentation; ventral premolar ridge not developed, ^ight mandible (Figs. 3B, D,

4B, D, 5B, D) cutting edge terebral ridge anteriorly, retinacular ridge posteriorly: terebral tooth blunt; retinacular ridge

well developed; anterior retinacular tooth prominent in Eucaerus, small in Asklepia\ (Reichardt, 1974: Fig. 6); premolar

tooth blunt, small, continuous with retinacular ridge; ventral premolar ridge indistinct. Ventral grooves long, setose,

extended more than 0.5 length of mandibles. Maxilla (Figs. 6-7) with sclerites generally elongate; lacinia with long

setae on dorsal surface; galeomere 2 distinctly shorter than 1; palpomere 4 slightly swollen, subulate apically. Labium

with mentum bisetose, median tooth developed (some members of Lachnaces), or not, or very slightly developed; lateral

lobes pointed apically; epilobes expanded apically; glossal sclerite narrow, bisetose, keeled ventrally; paraglossae

membranous, glabrous either shorter (Fig. 10) or longer (Fig. 12) than glossal sclerite; palpus with palpomeres 1 and 2

slender, 3 swollen, subulate apically.

Thorax. Pronotum various: subcordate (Fig 13) to pronouncedly transverse; base lobed or not; anterior angles

broadly rounded, posterior angles sharp or rounded; disc slightly convex, median longitudinal impression sharp, well

developed; anterior and posterior lateral impressions well developed. Metepisternum distinctly longer than wide.

Elytra. Average in form; humeri broadly rounded; basal ridge marginal, prominent, extended to scutellum; apical

margin obliquely subtruncte. Interneurs average or effaced, impunctate.

Wings. Well developed; wedge cell absent, oblongum cell stalked, well developed. Venation otherwise normal for

carabids.

Legs. Generally average for Lachnophorini. Tarsomere 4 with apical margin sub-truncate, tarsomere 5 with row of

ventro-lateral setae, each side. Male anterior tarsus ventrally (Figs. 14-16) with reduced adhesive vestiture, on

tarsomeres 2 and 3; present or not on tarsomere 1; tarsomere 4 with pair of flattened, expanded sense organs

apicoventrally (Fig.l4B).

Abdominal sterna. Average for Carabidae, in form; surface generally setose, or glabrous.

Male genitalia. Median lobe relatively broad in cross section, dorsal surface mostly membranous; apical orifice dorsal.

Internal sac with microtrichial fields only, or with latter and varied number and groups of spines. Parameres average for

Lachnophorini.

Ovipositor and associated abdominal sclerites. Tergum VIII completely sclerotized basally, not divided into two parts

by median membranous area; apodemes with apices curved laterad. Sternum VIII extensively unsclerotized medially.

Tergum X transverse, narrow. Valvifers average. Stylomeres 1 and 2 subequal in length, stylomere I with row of setae

apically, stylomere 2 (Figs.l7A, 18 A) falcate, blade slender, with preapical sensory furrow and long nematoid setae on

ventral surface, with two or three long spines on dorso-lateral margin, one on dorso-medial margin; row of sensory pits on

lateral and ventral surfaces.

Key to Genera of the Eucaerine Complex

1 (0 ) Pronotum with base truncate, not lobed medially. Disc of elytron with two rows

of setigerous punctures; interneurs effaced, intervals flat; bicolored;

microsulpture not evident at ordinary magnifications (to SOX)

Asklepia Liebke.

r Pronotum with base lobed medially (Fig. 13). Elytral disc with single row of

setigerous punctures (on interval 3); interneurs effaced or evident; concolorous

or bicolored; microsculpture not evident, or meshes transverse 2.

2 (1') Antennomeres 1-3 without vestiture of short setae, glabrous except for few,

normal (long) preapical setae. Male anterior tarsomeres ventrally without

adhesive vestiture Phaedrusium Liebke.

1' Antennomeres 1-3 (or 2-3) with vestiture of short setae, like antennomeres 4-11.

Male anterior tarsomeres 2 and 3, or 2-4 ventrally with adhesive vestiture (Figs.

14A, 15, and 16) Eucaerus LeConte, p. 107

The genus Asklepia includes the single species A. strandi Liebke, 1938. Phaedrusium

Liebke, 1951 includes P. suturalis Liebke, 1951 (generitype), and P. titschacki Liebke, 1951.

We have nothing further to add about these genera.

Quaest. Ent., 1983, 19 (1,2)

104

Ball and Hilchie

Figs. 3-7. SEM photographs of structures of Lachnophorini.— Figs. 3-5: mandibles, A and C, left, dorsal and ventral

aspects, respectively, B and D, right, dorsal and ventral aspects, respectively, of: 3, Lachnophorus guttulatus Bates; 4,

Eucaerus (sensu stricto) species; 5, E. (Lachnaces) olisthopoides (Bates). Eigs. 6-7: right maxilla of~-6, Eucaerus (sensu

stricto) species, ventral aspect; 7, E. (Lachnaces) olisthopoides (Bates), A, entire structure, ventral aspect, B, lacinia and

galea, ventral aspect, C, galea and lacinia, dorsal aspect. Scale bars = 50 yam. Legend, mandibles: art, anterior retinacular

tooth; m, molar; pm, premolar; prt, posterior retinacular tooth; rr, retinacular ridge; tm, terebral margin; vg, ventral

groove. Legend, maxilla: gl, galea; lac, lacinia

Cymindine Lebiini of Authors

105

Figs. 8-12. SEM photographs of Lachnophorini.— Structures of the labium. Figs. 8 and 9, palpomeres, microsculpture, A,

palpomere 1, B, palpomere 2, and C, palpomere 3, of: 8, Eucaerus (sensu stricto) species; 9, E. (Lachnaces) olisthopoides

(Bates). Scale bars = 5 um, Fig. 10: labium, ventral aspect, of Eucaerus (sensu stricto) species. Fig. 11: mentum and

palpigers, ventral aspect, of Eucaerus (sensu stricto) species. Fig. 12: labium, ventral aspect, of E. (Lachnaces)

olisthopoides (Bates). Scale bars = 50 Mm. Legend: el, epilobe; li, glossal (or ligular) sclerite; lp3, labial palpomere 3; m,

mentum; pg, paraglossae; pgr, palpiger.

Quaest. Ent., 1983, 19 (1,2)

106

Ball and Hilchie

Figs. 13-18. SEM photographs of structures of Lachnophorini. — Fig. 13: bases of head and elytra, and pronotum, dorsal

aspect, of Eucaerus (sensu stricto) hilaris Bates. Figs. 14-16, front tarsomeres of males, ventral aspect, showing adhesive

vestiture; 14, Eucaerus (sensu stricto) hilaris Bates, A-tarsomeres 1-4, B-tarsomere 4; 15, Eucaerus (sensu stricto)

species, tarsomeres 1-4; 16 E. (Lachnaces) olisthopoides (Bates), tarsomeres 1-5. Figs. 17-18: ovipositor, left stylomeres,

A-medial aspect, B-apico-ventral aspect, of: 17, Eucaerus (sensu stricto) species; 18, E. (Lachnaces) olisthopoides. Scale

bars. Figs. 13, 15, 16 = 50 Figs. 14, 17, 18 = 10 mhi. Legend, for tarsi: fs-foliose seta. Legend, for stylomeres: a,

lateral ensiform seta; b, medial ensiform seta; c, furrow pegs; d, nematoid seta.

Cymindine Lebiini of Authors

107

Eucaerus LeConte

Figs. 4-18

Eucaerus LeConte, 1853: 386. GENERITYPE: E. varicornis LeConte, 1853 (monotypy).- 1862: 22.- Chaudoir,

1871: 285.- Horn, 1881: 157, 159.- 1882: 158.- LeConte and Horn, 1883: 45.- Csiki, 1932: 1497.- Leng, 1920: 67.-

Blackwelder, 1944: 63.- Ball, 1960: 162.- Erwin et al, 1977: 4: 60.

Lachnaces Bates, 1872: 201. GENERITYPE: L. sericeus Bates, 1872: 201 (here designated). Csiki, 1932: 1497.-

Blackwelder, 1944: 63. NEW SYNONYMY.

Note about nomenclature. — The name Lachnaces sericeus, 1872 becomes Eucaerus

sericeus by virtue of combining Eucaerus and Lachnaces. However, in 1871, Bates had already

proposed the name E. sericeus for another species. Thus, the Bates name of 1872 becomes a

junior secondary homonym. For the species to which that name applied, we propose E.

sericatus, new name.

Classification. — The species of Eucaerus are arranged in two subgenera and two species

groups, as indicated in the following key.

Key to Subgenera and Species Groups of Eucaerus LeConte

1 (0 ) Antennomere 1 without vestiture of short setae. Pronotum subquadrate, sides

rounded, not sinuate; disc smooth, without pair of shallow depressions; surface

iridescent, microsculpture meshes transverse, in form of diffraction grating.

Elytron with inteneurs average, intervals convex. Maxillary palpomere 3 longer

than antennal scape. Labium with mentum as long as wide; paraglossa (Fig. 12)

narrow apically, longer than glossal sclerite. Male front tarsomere 1 without

adhesive vestiture ventrally, tarsomeres 2 and 3 with single row, only (Fig. 16).

Median lobe of male genitalia with apical portion very short and broad; internal

sac without spines subgenus Lachnaces Bates.

V Antennomere 1 with vestiture. Pronotum (Fig. 13) cordate, sides markedly

sinuate posteriorly, posterior angles sharp; disc with pair of paramedian shallow

depressions; pronotum with surface iridescent, microsculpture meshes grated,

not visible at 50X, or surface dull, meshes isodiametric, microlines visible at

50X. Elytron with interneurs average or effaced, intervals convex or flat.

Maxillary palpomere 3 shorter than antennal scape. Labium (Fig. 10) with

mentum wider than long; paraglossa broad apically, shorter than glossal sclerite.

Male front tarsomere 1 with or without adhesive vestiture; tarsomeres 2 and 3

with vestiture uniseriate (Fig. 14A) or biseriate (Fig. 15). Median lobe of male

genitalia with apical portion very short, or longer; internal sac with or without

spines Eucaerus (sensu stricto) 2

2 (L) Pronotum with sides narrow, proepisternum visible from dorsal aspect. Elytra

bicolored. Head and pronotum smooth, microlines absent. Elytra with

interneurs impressed or not, meshes transverse, surface iridescent, or microlines

obsolete, surface shining. Male front tarsomere 1 without adhesive vestiture,

tarsomere 2 and 3 with vestiture uniseriate. Median lobe with apical portion

short E. hilaris Group.

2' Pronotum (Fig. 13) with sides average, proepisternum not visible from dorsal

aspect. Elytra concolorous. Head and pronotum with surface dull,

microsculpture meshes isodiametric; elytra with surface iridescent, microlines in

form of diffraction grating. Elytra with interneurs normally developed. Male

Quaest. Ent., 1983, 19 (1,2)

108

Ball and Hilchie

front tarsomeres 1-3 with biseriate adhesive vestiture. Median lobe with apical

portion larger E. varicornis Group.

List of species. — The senior author has studied representatives of all described species of

Eucaerus . Names are listed here, and the species are assigned to their respective groups.

Subgenus Eucaerus

E. varicornis Group

E. sulcatus Bates

E. striatus Bates

E. sericeus Bates

E. opacicollis Bates

E. insularis Darlington

E. haitianus Darlington

(additionally, three undescribed species from Mexico).

E. hilaris Group

E. geminatus Bates

E. hilaris Bates

E. lebioides Bates

E. pulchripennis Bates

Subgenus Lachnaces Bates

E. sericatus, new name (=E. sericeus Bates, 1872, not 1871).

E. badestrinus Bates

E. olisthopoides Bates

Notes about habitat. — Members of this genus live in leaf litter, in swamp forest, or in flood

zones along tropical rivers. Adults of the E. hilaris Group are in litter in areas with more light,

close to river edges, whereas adults of the E. varicornis Group and Lachnaces are in more

densely shaded places. On the Rio Negro, in northern Brazil, adults of the latter two groups are

microsympatric.

Geographical distribution. — Species of subgenus Lachnaces and of the E. hilaris Group

are known only from the Amazon Basin, in Brazil. Range of the E. varicornis Group extends

from the Amazon Basin northward to southeastern United States, and eastward to the Greater

Antilles. However, no species are shared between South America and areas further north, nor

between the West Indies and the adjoining continents.

Tribe LEBIINI

As background for more detailed consideration of cymindines, we need to comment about

the tribe Lebiini, which includes the subtribe Cymindina. Collectively, lebiine adults are

strikingly divergent in form, color, and in more detailed external features, making it difficult to

provide a simple diagnosis for recognition of the tribe. Some adults (cymindines) look much like

platynines, others (Nemotarsus members) have the long pectinate tibial spurs of masoreines,

others (some Lebia members) are hardly different from pentagonicines in form and color, and

still others (members of Agra) are colliurine- like. Internal features and mouthparts offer a

similar range of attributes. While it seems unlikely that the Lebiini is a polyphyletic taxon, it

could very well be paraphyletic. It is polythetic, for most character states used for recognition

of the group are not shared by all member taxa, and those states that seem to be almost

universal (biperforate anterior coxal cavities, two pairs of supraorbital setae, for example) are

shared with members of non-lebiine taxa.

We are not, however, prepared to pursue this subject further. These comments are words of

caution for those who use the following list of features for identification of adults, or those who

wish to pursue phylogentic studies of carabids.

Cymindine Lebiini of Authors

109

Recognition. — Most lebiine adults exhibit most of these character states: apical margins of

elytra truncate or subtruncate; tergum VIII more or less extensively membranous medially,

laterally exposed, each side posteriorly with a projection that bears the openings of ducts of

defensive glands; head with two pairs of supraorbital setigerous punctures; tibial spurs of

middle and posterior legs of equal length, smooth, not serrate (if unequal and serrate, head

sharply constricted posteriorly); terminal palpomeres more or less pubescent, apical margins

subtruncate or truncate (not swollen medially and tapered to narrow apex); antennomeres 4-11

setose; front tarsomeres 1-3 of males with biseriate adhesive vestiture; anterior coxal cavities

biperforate; abdomen with sternum X principally membranous; median lobe of male genitalia

with dorsal surface extensively sclerotized, membranous area relatively small; right paramere

smaller than left paramere; ovipositor with stylomere 1 setose or spinose.

Some pericaline and gallerucoid calleidine adults have virtually complete elytra, with apices

extended to the apex of tergum VII. However, pericalines are recognized by a combination of

well developed suborbital setae, displaced penultimate umbilical setigerous puncture, and long,

slender labrum. Gallerucoid calleidines are chrysomelid-like in appearance, with well developed

suborbital setae on the head.

Notes about classification.— The tribe Lebiini, as generally accepted by carabid specialists

(for example, LeConte and Horn (1883), Sloane (1923), Andrewes (1929), Ball (1960),

Lindroth (1969), and Erwin (1979)) was assembled by Horn (1881: 154), who combined the

Lebiides and Pericalides of Lacordaire (1854), but excluded the genera Mormolyce Hagenbach

and Agra Fabricius. Subsequently, these genera were returned to the Lebiini {Mormolyce, by

Ball, 1975: 147, and Agra, by Erwin, 1978: 263). Erwin (1979: 590) also returned the

eucheiline genera Eucheila Dejean and Inna Putzeys to the Lebiini.

Grouping the numerous lebiine genera has been a problem since it was first attempted by

Lacordaire (1854: 102). In addition to the Pericalides, he recognized three basic forms

centering on Cymindis Latreille, Dromius Bonelli, and Lebia Latreille. Lacordaire wrote that

he was unable to find diagnostic characters for such groups.

Chaudoir gave tribal ranking to these groups, as well as to several others, based on slight

differences in structure of the labium, as well as on other features. Horn (1881) undertook a

detailed study of maxillae and labia of carabids, and one of his conclusions was that the

differences among lebiine tribes were too slight and inconstant to be valid as taxonomic

characters at the tribal level. Horn’s lead was followed by European workers of the late 19th

and early 20th centuries. For example, Csiki (1932: 1305-1500) included in the Lebiini most of

the groups that Horn had included. He recognized seven subtribes, four of which were groups

proposed by Lacordaire: Lebii, Catascopi (equivalent to Pericalides), Dromii, and Cymindina.

Three other subtribes were established for genera included by Lacordaire in one or the other of

his groups of Lebiides: Physoderi, Lebidii, and Callidi. Nemotarsines, agrines, and masoreines

were excluded, each being assigned to a tribe of its own.

Jeannel (1949: 876-1039) used a system similar to that of Lacordaire, for organizing the

lebiine fauna of Madagascar, but he excluded nemotarsines and masoreines. He recognized

three families (Lebiidae, Thyreopteridae, and Lionychidae), the second including many of the

genera that Lacordaire included in the Pericalides. Jeannel included physoderines and

lebidiines in the Lebiidae. Genera of Lebiidae were arranged in five subfamilies: Cyminditae,

Lebiitae (including also physoderines), Coptoderitae, Calleiditae (including Lebidii), and

Dromiitae. Genera of Thyreopteridae were arranged in two subfamilies: Thyreopteritae and

Pericalitae. Lionychidae, a new family, included four genera regarded as dromiines by most

Quaest. Ent., 1983, L9 (1,2)

110

Ball and Hilchie

authors.

Jedlicka (1963: 295-464) recognized the same seven subtribes into which Csiki arranged the

genera of Lebiini.

Habu (1967: 60) recognized eight subtribes: Cymindina, Catascopina, Pericalina,

Anomotarina, Calleidina, Lebiina, Demetriina, and Dromiina. The Cymindina and Lebiina are

each about the same as proposed by Csiki; catascopines and pericalines are the equivalent of

Catascopi; demetriines (proposed first by Bates (1886: 207)) and dromiines are the equivalent

of Dromii; and calleidines (including Lebidiina and Physoderina) and anomotarines (new

subtribe) are the equivalent of Callidi.

It is evident that central to these more or less divergent arrangements is the system proposed

by Lacordaire, with various assemblages of his four basic groups (three of Lebiides plus

Pericalides) shifted about on the basis of detailed study and weighting of various character

systems. Authors previous to Habu relied principally on details of structure of: labium

(particularly of the ligula); pronotum; and tarsi, particularly form of tarsomere 4 and

pectination of the claws. Habu used these features, and also form of mandibles and details of

structure and armature of the ovipositor sclerites.

Although Habu’s treatment is restricted to the fauna of Japan and adjacent islands, most of

the major groups of lebiines are represented there. His illustrations of structures are profuse,

well-chosen, and well executed, his descriptions are detailed and accurate, and he has exhibited

a good sense of proportion in ranking. It seems to us that Habu has provided a firm basis for

resolving the long-standing problem of recognition of natural (i.e., phylogenetically valid)

groups of lebiines.

To work out a phylogentically valid classification, it is necesary to reconstruct the phylogeny

of the Lebiini. Clues are provided by association of many groups of lebiines with vegetation,

and at least some character states of adults (particularly those of the tarsi) seem to be

associated with life above the surface of the ground (Erwin, 1979: 552). Which way has

evolution of lebiines proceeded: from occupation of terrestrial to arboreal habitats; or from

arboreal to terrestrial; or from terrestrial to arboreal and back to terrestrial? The same sorts of

questions are applicable to arboreal habitats. Some lebiines live principally on tree trunks,

others hunt on small branches and twigs, still others on leaf surfaces (Erwin, 1979: 559-560,

Table 1). What has been the direction of evolution within arboreal habitats?

If these questions could be answered for all comparisons of taxa thought to be related, it

would be possible to work out a classification consonant with direction of habitat change.

Probably the arboreal zone has been invaded by terrestrial- based ancestors (Erwin, 1979: 509,

Fig. 13), but it also seems likely that some ancestral stocks have given rise to terrestrial

inhabitants, as well. Movements in both directions may have taken place several times.

Structure of the ovipositor may be associated with different modes of egg-laying, and if

these modes were known they might offer another basis for inferring evolutionary sequences.

Mode of oviposition is known for some terrestrial calleidines: females of Tecnophilus and

Philophuga climb on low plants, carrying on the stylomeres of the ovipositor a small ball of

mud. An egg is laid in the mud ball, and the latter is suspended from a twig by a silken thread

produced by the female (Larson, 1969: 64).

Females of most groups of carabids are believed to oviposit in the ground, in chambers

scooped out by the ovipositor. Compared to the latter, calleidines seem to be apotypic in

oviposition. The ovipositor of Tecnophilus and many other calleidines is characterized by

absence of ensiform setae from stylomere 2 and narrow form, whereas females of

Cymindine Lebiini of Authors

111

ground-ovipositing carabids have broader second stylomeres and ensiform setae.

Erwin (1982: 40), referring to the remarkable telescopic ovipositor that charcterizes females

of the genus Agra, inferred that such structures are used to lay eggs “deep in existing burrows

in wood or in other deep fissures”. Stylomeres of Agra females also seem apotypic in their

elongate form and reduced number of spine-like ensiform setae.

Among lebiines, pericalines (most taxa are arboreal) and apenines (all known taxa are

terrestrial) have the more plesiotypic form of ovipositor. However, almost nothing is known

about where eggs are laid or how they are laid by members of these groups. (An exception is

the genus Eurycoleus, females of one species of which lay eggs on the surfaces of wood, near

endomychid pupae which the developing Eurycoleus larvae eat [Erwin and Erwin, 1976]). We

are satisfied that evidence from structure of the ovipositor offers sufficient grounds to infer that

apenines and pericalines are relatively primitive lebiines, that cymindines, with moderately

modified ovipositors, occupy an evolutionarily intermediate position, and that the other

subtribes whose females have highly modified ovipositors, represent more highly evolved

groups. Details of relationships among genera and subtribes remain to be worked out.

In lieu of a definitive treatment of classification of the Lebiini, we offer a key to the

subtribes, based on features of adults.

Key to Subtribes of Lebiini

1 (0 ) Head ventrally with at least one pair of suborbital setigerous punctures 2.

V Head ventrally without suborbital setigerous punctures 4.

2 (1 ) Labrum narrow, as long or longer than wide. Penultimate setigerous puncture of

umbilical series of elytron displaced laterally (as in Fig. 27 B)

Subtribe Pericalina, p. 1 16

1' Labrum normal, wider than long. Penultimate setigerous punctures of elytra not

displaced laterally 3.

3 (2' ) Elytron smooth, without striae. Pronotum with sides curved, widest near base,

narrowed evenly anteriorly, apical margin much narrower than basal margin.

Head sharply constricted posteriorly, pedunculate. Stylomere 2 of ovipositor

with broad apex, without ensiform setae

gallerucoid Calleidina.‘

3' Elytron striate. Pronotum with sides sinuate posteriorly, widest at or anterior to

middle. Head gradually constricted posteriorly. Stylomere 2 of ovipositor with

narrowed apex, ensiform setae two, one dorsal, one ventral

genus Euproctinus Leng and Mutchler, 1927.^ p.

4 (L) Penultimate setigerous puncture of elytron displaced laterally. Stylomere 2 of

ovipositor with ensiform setae, and stylomere 1 with prominent ventral

projection extended beyond base of stylomere 2 (Figs. 38 and 39)

^Adults of Lebidia Morawitz and Gallerucidia Chaudoir (Lebidii or Gallerucidiini, of authors)

key out here although in all other respects they seem to be calleidine.

^This Neotropical and southern Nearctic genus seems to be of uncertain position. It has been

included with calleidines, based on general appearance and structure of tarsi, but Larson (1969:

23) suggested Euproctinus should be placed in a subtribe of its own.

Quaest. Ent., 1983, 19 (1,2)

112

Ball and Hilchie

Subtribe Apenina, p. 120

4' Penultimate setigerous puncture of elytron not displaced laterally, thus in line

with rest of series, or displaced toward stria 8. Stylomere 1 of ovipositor without

projection; stylomere 2 with (Fig. 62 A) or without (Fig. 96B) ensiform setae . 5.

5 (4') Posterior tibial spurs markedly unequal, margins serrate, inner spur almost as

long as tarsomere 1 . Head sharply constricted posteriorly, pedunculate

Subtribe Nemotarsina.

5' Posterior tibial spurs subequal, margins smooth, not markedly serrate. Head

sharply constricted or not 6.

6 (50 Mandible widened near base, scrobe wide, lateral margins markedly rounded . 7.

6' Mandible not conspicuously widened basally, scrobe narrowed, lateral margins

not markedly rounded 8.

7 (6 ) Head markedly narrowed and prolonged behind eyes. Pronotum longer than

wide, markedly narrowed anteriorly, without lateral flange. Ovipositor

strikingly telescopic, stylomere 2 elongate Subtribe Agrina.

7' Head average, not markedly prolonged behind eyes (Fig. 101). Pronotum wider

than long, or as wide as long, not markedly narrowed anteriorly, basal and

apical margins subequal in width, with lateral flange. Ovipositor not strikingly

telescopic, stylomere 2 not especially lengthened

Subtribe Calleidina, p. 173

8 (60 Tarsomeres broad, tarsomere 4 with apex subtruncate, not bilobed. Female with

stylomere 2 with one or two ensiform setae (Fig. 55A)

Subtribe Cymindina, p. 129

8' Tarsomeres broad, with tarsomere 4 bilobed, OR tarsomeres slender and

tarsomere 4 with apical margin sub-truncate. Stylomere 2 of ovipositor without

ensiform setae 9.

9 (80 Tarsomeres slender, tarsomere 4 with apical margin sub-truncate. Stylomere 2

of ovipositor glabrous or setose apically Subtribe Dromiina^ p. 196

9' Tarsomeres stout, dilated, tarsomere 4 bilobed. Ovipositor with stylomere 2

glabrous 10.

10 (90 Tarsomere 4 with lobes almost half length of tarsomere 5. Ovipositor with

stylomere 1 fully sclerotized, stylomere 2 narrow, tapered apically

Subtribe Demetriina.

10' Tarsomere 4 with lobes short, less than half length of tarsomere 5. Stylomere 1

partially desclerotized, stylomere 2 broad, short, broadly rounded apically ....

Subtribe Lebiina.

^Habu (1967: 250) expressed doubt about including Celaenephes Schmidt-Goebel in the

Dromiina because of the setose stylomeres 1 and 2 of its females. Thus it would not key out

above. Bates (1892: 156) included this genus among the cymindines, along with several other

genera that were subsequently assigned to the Dromiina (Csiki, 1932). Celaenephes is clearly

not a dromiine, and we believe that the stylomeres of its females are too plesiotypic for the

genus to be included in the Cymindina. It may be a platynine, or it may represent a separate

lineage of Lebiini that will require establishment of another subtribe.

Cymindine Lebiini of Authors

113

Figs. 19-21. Photographs of Pericalina, genus Thyreopterus.—H^h'xins, dorsal aspect. 19, T. (Thyreopterinus) species?

(SBL = 5.38 mm); 20, T. (sensu stricto) kivuanus Basilewsky. (SBL = 6.30 mm); 21, (Selenoritus) ptolemaei (Alluaud)

(SBL = 5.32 mm).

Quaest. Ent., 1983, 19 (1,2)

114

Ball and Hilchie

Cymindine Lebiini of Authors

115

Figs. 22-24. Line drawings of structures of Pericalina, genus Thyreopterus.—Msi\c genitalia. Fig. 22: T. (Thyreopterinus)

species. A, B, C— median lobe, left lateral, ventral, and right lateral aspects, respectively, D and E, parameres, left and

right, respectively, ventral aspect. Fig. 23; T. (sensu stricto) kivuanus Basilewsky -A and B, median lobe, left lateral, and

ventral aspects, respectively; C and D, parameres, left and right, respectively, ventral aspect. Fig. 24; T. (Selenoritus)

ptolemaei (Alluaud)-A and B, median lobe, left lateral and ventral aspects, respectively; C and D parameres, left and

right, respectively, ventral aspect. Fig. 25. Line drawings of structures of Apenina.— Wing cells and surrounding veins of

Cymindoidea (sensu stricto) indica Schmidt-Goebel, left wing: A, oblongum cell; B, wedge cell. Legend: cells-O,

oblongum, W, wedge; veins— A, Anal; Cu, Cubital; M, Median; R, Radial.

Quaest. Ent., 1983, 19 (1,2)

116

Ball and Hilchie

Tribe LEBIINI, Subtribe PERICALINA

Two genera {Selenoritus Alluaud, 1917, and Leptosarcus Peringuey, 1896), described

originally as cymindines, are more appropriately assigned to the Pericalina because adults of

each genus exhibit the diagnostic features of this subtribe: extended mouthparts (including

elongate labrum), pair of suborbital setae, laterally displaced penultimate umbilical setigerous

puncture of an elytron (Fig. 27B), stylomere 2 relatively small, falcate, with three large dorsal

setae, and without a ventral preapical sensory furrow or nematoid setae (Figs. 28A-C).

Within the Pericalina, we place both of these genera in the thyreopteroid assemblage:

Selenoritus, because it is actually a member of Thyreopterus; and Leptosarcus because

stylomere 2 of the ovipositor lacks nematoid setae.

Jeannel (1949: 975) included Selenoritus in the tribe Thysanotini, subfamily Calleiditae,

along with the Madagascan endemic genera Antimerina Alluaud, Thysanotus Chaudoir, and

Madecassina Jeannel. External features of adults of these genera (seen in the MCZ) confirm

that they are pericalines, and absence of nematoid setae from stylomere 2 of females of

Antimerina elegans Alluaud, and Thysanotus alluaudi (Jeannel) provide the basis for

assigning this geographical complex of genera to the thyreopteroid assemblage. Basilewsky

(1953a: 10) suggested that Thysanotini should be included in the Thyreopteridae, but Ball

(1975:147), on the basis of study of descriptions and illustrations, suggested that such a

grouping would be incorrect. This group could be near the base of the stock that gave rise to the

thyreopteroid radiation on Madagascar.

Selenoritus Alluaud, 1917

Figs. 21-22

Selenoritus Alluaud, 1917: 103. GENERITYPE: Selenoritus ptolemaei Alluaud, 1917: 104 (monotypy).

LECTOTYPE male (here selected), labelled: MUSEUM PARIS MONTS ROUWENZORI zone des forets Makitawa

(2660 m) Ch. Alluaud 1909 [blue paper]; TYPE [red paper]; Museum Paris coll. Ch. Alluaud [blue paper]; Selenoritus

ptolemaei Alluaud Type [white paper, with blue strip across top]. [MNHP]. PARALECTOTYPE male, similarly labelled

in Musee d’Afrique Centrale, Tervuren.- Burgeon, 1937: 356.

Selenorites (misspelling) Jeannel, 1949: 975.- Basilewsky, 1962: 300 and 321.

Notes about type material. — The type locality of S. ptolemaei is more fully specified, as

follows: ZAIRE, Mount Ruwenzori, east versant, in forest above the shelter, beneath peak of

Makitawa, between 2600 and 2800 meters (Alluaud, 1917). Alluaud (1917: 103-104) provided

a detailed description of external features of type specimens. His basis for claiming a

relationship of this species to the cymindines is a combination of these features: truncate elytra,

not covering apex of abdomen; broad paraglossae, clearly extended beyond apex of ligula; and

denticulte tarsal claws.

Alluaud lists the following features as diagnostic of Selenoritus: disc of elytra more covex;

elytra more ovoid with humeri more rounded, and basal groove not sinuate between humeri and

scutellum; posterior pair of supraorbital setigerous punctures far removed posteriorly on

occiput; antennomere 3 with more than apical setae; lateral margins of pronotum without

setigerous punctures; and posterior tarsi with tarsomeres 1-5 filiform, not dilated nor grooved

dorsally, elongate and subequal to one another. Most of these character states, however, appear

in the pericaline genus Thyreopterus (sensu lato) as pointed out in conversation by Dr. P.

Basilewsky, who had previously recognized the similarities between members of these two taxa.

Cymindine Lebiini of Authors

117

Pectinate tarsal claws and small size place Selenoritus near the subgenus Thyreopterinus

Alluaud.

Character states that distinguish adults of Selenoritus from those of Thyreopterinus are;

small eyes (Fig. 21; cf. Fig. 19); posterior pair of supraorbital setigerous punctures clearly

behind posterior margins of compound eyes; pronotum without posterior pair of setigerous

punctures (members of both groups lack the anterior pair); basal ridge of elytron not extended

to sutural margin, but terminated near base of interneur 4; metathorax and hind wings reduced.

Small eyes, loss of setae, and reduced metathorax and hind wings seem to be adaptations

associated with life in montane environments, and the position of the posterior pair of

supraorbital setigerous punctures is probably the result of reduction of eyes, rather than

posterior migration of the setigerous punctures. These differences might have evolved relatively

recently, and thus do not constitute evidence that S. ptolemaei is phylogenetically old. Instead,

this species may be only a moderately specialized member of Thyreopterinus.

On the other hand, many montane-adapted stocks seem to be relics of older stocks that have

been replaced in the lowlands by later evolving relatives. Until the relationships of

Thyreopterinus and Selenoritus can be more fully resolved, it seems as well to treat the two

groups as separate subgenera of Thyreopterus. Evidence supporting this decision is provided by

details of stylomere 2 of the ovipositor, for a combination of number and length of ensiform seta

and form of these sclerites themselves distinguish females of these groups from one another. See

Table 1 for details.

Male genitalia of Selenoritus ptolemaei are also markedly different from those of the one

species of Thyreopterus examined (Fig. 24; cf. Fig. 23). In males of both S. ptolemaei and T.

TABLE 1.

COMPARISON :OF FEATURES OF STYLOMERE 2 OF :THE OVIPOSITOR OF

FEMALES OF SUBGENERA OF THYREOPTERUS DEJEAN

STYLOMERE 2

Ensiform Setae Apical Portion

L. dorso-medial

Quaest. Ent., 1983, 19 (1,2)

118

Ball and Hilchie

(sensu stricto) kivuanus, the apical orifice of the median lobe is slightly left of the mid-line; in

males of Thyreopterinus species, it is to the right. However, such differences are common

among pericalines, and their evaluation must be made in terms of additional species of

subgenus Thyreopterinus.

Thyreopterus (Selenoritus) ptolemaei Alluaud, 1917, new combination

Figs. 2 1-22 A, C

Description. — Habitus as in Fig 21. Standarized body length 6.20 mm. (lectotype; other specimens of similar

size). Form pterostichoid or agonoid, slender.

Color generally rufo-piceous dorsally, more rufous ventrally, palpi, antennae and legs flavous. Elytra each with three

groups of rufo-flavous marks: one group in basal 0.20 on intervals 2, 3, 6, 7, and 8; one group medially on intervals 7 and 8;

and one group in apical 0.80 on intervals 2-8.

Microsculpture of dorsum. Head and elytra, with meshes isodiametric, those of elytra slightly shingled; pronotum with

meshes transverse.

Luster. Surface generally shining.

Head. Clypeus longer than average; anterior margin concave; bipunctate, each puncture in longitudinal groove

extended to posterior margin. Frons with impressions broad and shallow, each side with single longitudinal ridge; vertex

slightly convex. Posterior pair of supraorbital setigerous punctures well posterad of posterior margin of compound eyes.

Temples not extended.

Eyes. Reduced. Paragenae at narrowest less than width of antennal scape.

Antennae. Length average: scape slightly longer than antennomere 3, and slightly broader; outer antennomeres longer

than wide (ant. 9 1/w- 3.00). Scape with single seta; pedicel with terminal ring of setae; antennomere 3 generally sparsely

setose; remaining antennomeres setose.

Mouthparts. Labrum longer than average, tapered anteriorly. Mandibles elongate, slender (not studied in detail).

Maxilla: stipes with several setae; palpus slender, palpomere 4 distinctly longer than 3; apical margin truncate, narrow.

Labium: mentum with well developed tooth; ligula narrow, bisetose apically; paraglossae broad, extended clearly beyond

apex of ligula; palpi slender, palpomere 2 bisetose; palpomere 3 with apical margin truncate.

Pronotum. Without lateral setae. Dorsal surface generally sparsely setose, setae short. Form slender, elongate, sides

markedly sinuate posteriorly; anterior margin concave, angles short but distinctly set off; basal margin truncate. Sides

moderately elevated, lateral grooves narrow, indistinctly isolated from posterior lateral impressions by convexity; median

longitudinal impression shallow; anterior and posterior transverse impressions evident, but broad.

Prosternum. With few setae at apex of intercoxal projection.

Metepisternum. Short, almost quadrate.

Elytra. Slightly explanate, widest point evidently behind middle; humerus broadly rounded; basal ridge terminated

near base of interneur 4, not extended to suture; apical margin sinuately truncate. Surface sparsely punctate, setae short.

Parascutellar setigerous punctures present. Interneurs teminated before apex, shallow; intervals slightly convex.

Umbilicate punctures 16, penultimate puncture slightly displaced laterally. (Lectotype with right elytron broken and

detached).

Hind wings. Markedly reduced.

Legs. Average, generally. Tibial spines reduced, as usual for pericalines. Anterior femur with numerous setae

ventrally. Anterior tibia with terminal spur thickened. Claws long, each with 4-5 pectinations. Anterior tarsomeres without

adhesive vestiture.

Abdomen. Sterna average, sternum 6 with four setae near posterior margin.

Male genitalia. Median lobe (Figs. 22A-C) short, broad; apical portion in ventral aspect short, rounded; dorsal surface

extensively sclerotized; apical orifice long, inclined to left. Internal sac with narrow sclerotized rim apically, otherwise

unarmored. Left paramere with apical margin sinuate- truncate. Right paramere with apex acute. (Cf. Figs. 23A-C and

24A-C)

Ovipositor. Stylomeres 1 and 2 subequal in length. Stylomere 2 elongate, hardly curved, dorsally with two broad

ensiform setae; without nematoid setae.

Geographical distribution and habitat. — This species is known from the higher slopes of

Mt. Ruwenzori, Zaire. Two specimens were collected in dead bamboo.

Material examined. — We have seen the types and three additional specimens, as follows:

Two males- Kilindera, north face of Ruwenzori, 2750 m., VII- VIII. 1974 R. P. M. Lejeune (MACT). Female.- Vallee

Mont Mulungu, Massif Ruwenzori, 2600 m., 29.1 1.1957, P. Vanschuytbroeck (MACT).

We also examined superficially material representing five additional species of

Thyreopterus {sensu stricto) and four additional species of subgenus Thyreopterinus, from

Cymindine Lebiini of Authors

119

Figs. 26-28. Photographs of Pericalina, genus Leptosarcus.—¥'\%. 26: L. hessei Basilewsky, habitus, dorsal aspect (SBL =

12.86mm). Figs. 27-28. SEM photographs of elytra and stylomeres of L. porrectus Peringuey. Fig. 27: Left elytron,

microsculpture, dorsal aspect-A, discal area; B, preapical area. Fig. 28 ovipositor, left stylomeres; A, stylomeres 1 and 2,

medial aspect; B, stylomere 2, lateral aspect; C, stylomere 2, apico-ventral aspect. Scale bars. Figs. 27-28 = 50 ^m.

Legend, elytra: dp, penultimate umbilical puncture, displaced toward lateral margin. Legend, stylomeres: a, lateral

ensiform seta; b-medial ensiform seta; SI, stylomere 1; S2, stylomere 2.

Quaest. Ent., 1983,19 (1,2)

120

Ball and Hilchie

collections of CAS. Fig. 20 illustrates the habitus of T. {sensu stricto) kivuanus Basilewsky.

Leptosarcus Peringuey, 1 896

Leptosarcus Peringuey, 1896: 218. GENERITYPE: Leptosarcus porrectus Peringuey, 1896: 219 (monotypy).-

Basilewsky, 1954a: 83.

Basilewsky (1954a) studied the few specimens of Leptosarcus that were available, including

type material of L. porrectus Peringuey (type locality- Vonstantia, Cape Province, South

Africa). He provided a description of adult features generally satisfactory for recognition of

specimens, and figured heads, labra, labia, and male genitalia. He also described a second

species, L. hessei (type locality- Zululand). To Basilewsky’s characterizations, we add the

following observations.

Microsculpture of the elytra is shingled (Figs. 27 A, B), like that of some of the more highly

derived members of the New World genus Phloeoxena (see Ball, 1975), and is quite unlike the

smoother microsculpture characteristic of the elytra of cymindine adults. The penultinate

umbilical setigerous punctures of the elytra are displaced laterally (Fig. 27B). Stylomeres 1 and

2 (Figs. 28A-C) are typical of the thyreopteroid Pericalina. Probably adults of Leptosarcus

should be sought in the types of habitats occupied by Phloeoxena adults; i.e., in association

with fallen logs, or standing trees with loose or scaly bark, in wet forests.

We conclude that general similarity in form and size between adults of Leptosarcus and of

Hystrichopus {sensu stricto) is convergent. Males of Leptosarcus have anopic median lobes as

have males of Cymindis, but this feature is plesiotypic, and is thus not of use in establishing

relationships.

This genus seems to be relict for several reasons: low diversity; seemingly without close

relatives among pericalines; adults brachypterous, and metathorax reduced; and geographical

distribution peripheral to the main area (tropics) of the Pericalina.

Material examined. — We have seen seven specimens representing both known species, all

from the collections of the South African Museum, and all collected at localities in the Union of

South Africa, as follows.

Leptosarcus porrectus Peringuey

Figs. 27-28

Male, holotype, labelled: C.T. 8.26 type; HOLOTYPUS [red paper]; Leptosarcus porrectus P; Leptosarcus porrectus

Per Basilewsky vid 1953. Male, paratype, from same locality as holotype, and also seen by Basilewsky. Female, same

locality, det. by Basilewsky, 1953. Female, Hott- Holl Mts. 4000 f., Caledon C.C., Bernard 1916; det. by Basilewsky,

1953.

Leptosarcus hessei Basilewsky

Fig. 26

Female HOLOTYPUS [red paper]; Mt. Kendhla forest Zululand; Leptosarcus hessei n.sp. P. Basilewsky det. 1953.

Tribe LEBIINI, Subtribe APENINA

This subtribe was erected by Ball (1982). Diagnostic character states are: head without

suborbital setigerous punctures; elytron with penultimate umbilical puncture laterad of

antepenultimate and ultimate umbilical punctures; tibiae and tarsi relatively slender;

Cymindine Lebiini of Authors

121

ovipositor, (Figs. 38, 39A, B) with stylomere 1 much longer than 2, asetose; stylomere 2

markedly curved, apex of blade pointed; two large ensiform setae on dorsal margins; preapical

sensory furrow and associated setae absent.

Description. — The following statements indicate range of variation of selected features

useful for recognizing apenine adults, and for determining relationships of taxa.

Color. Various, but mostly somber: dorsum dark rufous to black, elytra with or without paler spots; venter piceous to

testaceous; legs and palpi of most specimens pale- rufous to testaceous, though femora of some specimens as dark as

ventral surface.

Microsculpture. Labrum and clypeus with meshes isodiametric. Dorsum of head with meshes isodiametric, or

microlines effaced; venter with meshes transverse. Pronotum with meshes isodiametric, or transverse, or microlines

effaced. Lateral and ventral thoracic sclerites with meshes transverse (characteristic of most groups) or isodiametric.

Scutellum with meshes isodiametric (characteristic of most groups) or transverse. Elytra with meshes isodiametric,

transverse, or effaced. Abdominal sterna with meshes transverse, or transverse medially, and isodiametric laterally.

Luster. Surface of head and thorax shining to dull; surface of elytra and abdominal sterna iridescent, shining, or dull.

Macrosculpture. Surface generally smooth, except as noted. Head (frons and vertex), pronotum, and elytral intervals

smooth, or variously transversely ridged and grooved. Surface impunctate, or covered with coarse punctures.

Vestiture. Surfaces of adults of most taxa glabrous, but Trymosternus adults generally setose. Antennomeres 1 and 2

with setae confined to apex, or generally setose; antennomere 3 with setae confined to apical 0.50, or generally setose;

antennomeres 4-1 1 generally setose. Tarsomeres dorsally setose.

Fixed setae. Average for lebiine adults: labrum with six long marginal setae, clypeus with one pair; head and pronotum

each with two pairs; elytra each with two discal setae in interval 3, parascutellar and preapical setae, and 15 umbilical

setae along lateral margin; penultimate umbilical seta displaced laterally of an imaginary line extended between

antepenultimate and ultimate umbilical punctures. Legs (anterior, middle, and posterior) with number of setae as follows:

coxae- 0-1, 2-5, 2; trochanters- 1, 1, 1; femora- 2 (posterior face), 3-5 (anterior face), 2 (anterior face). Sternum VII

with two setae in males, and two or four setae in females.

Head. Clypeus transverse, anterior margin of each truncate or slightly concave. Frontal impressions shallow, indistinct.

Sub-antennal ridge average or prominent. Eyes: orbicular, convex, visible in ventral aspect; or reduced, longer than wide,

flattened, ventral margin obliquely truncate, and not visible in ventral aspect. Insertion of antennal scape close to or remote

from anterior margin of adjacent eye.

Antennae. Average for lebiine adu'ts: filiform, flagellar antennomeres sub-cylindrical, distinctly longer than wide,

antennomere 2 short, antennomere 3 longer than 4.

Mouthparts. Labrum transverse, anterior margin truncate or slightly concave. Mandibles. Left and right mandible

about same shape, overall. Left mandible (Figs. 29A, C,- 30A, C) with terebral margin reduced, no terebral tooth. Cutting

edge retinacular ridge; posterior retinacular tooth small, not divided; ventral retinacular ridge blunt; premolar triangular;

premolar ridge well developed, sharp. Right mandible (Figs. 29B, D - 30B, D) with terebral margin cutting edge, terebral

tooth blunt, large; retinacular ridge well developed, anterior and posterior teeth blunt; ventral ridge not developed;

premolar tooth triangular, sharp at apex; ventral groove extended basad, to premolar area. Maxillae, average for lebiine

adults: lacinia with (Fig. 32), or without (Fig. 31) apico-lateral setae; palpomeres slender, 4 with apical margin truncate,

slightly longer than 3, markedly longer than 2. Labium: mentum (Figs. 33-36) bisetose, with lateral lobes pointed apically

(Fig 33) or broadly rounded (Fig. 34), tooth well developed, pointed apically (Figs. 33-35), or absent (Fig. 36); glossal

sclerite with apical margin broad, sub-truncate, bisetose (or quadrisetose, median two setae close together, much shorter

than lateral pair); paraglossae fused to glossal sclerite, apical margins finely setose, hardly extended beyond apex of glossal

sclerite; palpomeres 1 and 2 slender, palpomere 3 more (Fig. 33) or less (Fig. 36) broadly securiform, more so in males

than in females.

Thorax. Pronotum with sides rounded, more constricted basally than apically (or markedly cordate, constricted

basally, sides strikingly sinuate basally); base lobate medially (or almost truncate); anterior angles broadly rounded,

posterior angles sharp, prominent; disc slightly convex, median longitudinal impression sharply defined, anterior and

posterior transverse impressions hardly evident; posterior lateral impressions shallow, indistinct. Prosternum with

intercoxal process immarginate. Metepisternum distinctly longer than wide, lateral margin 1.5 times longer than anterior

margin (or almost as long as wide, anterior and lateral margins subequal).

Elytra. Average in form; humeri prominent, extended slightly forward, basal ridge marginal, extended to edge of

scutellum. Apical margin obliquely subtruncate. Interneurs average for carabids (or broader than average), punctate;

scutellar interneur well developed. Intervals slightly convex (or alternate odd-numbered intervals sub-carinate to carinate,

raised above even-numbered intervals).

Wings. Well developed (or short stubs); wedge cell absent (Fig. 25B), oblongum cell average (Fig. 25A) (or reduced,

or absent). Venation otherwise normal for lebiines.

Legs. Average for carabids. Middle tibia with spines of outer row numerous, extended length of margin (or spines few,

located in apical 0.25). Tarsal claws pectinate. Tarsomere 4 notched, but not bilobed. Male with front tarsomeres 1-3

ventrally with biseriate adhesive vestiture.

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122

Ball and Hilchie

Figs. 29-37. SEM photographs of structures of Apenina. — Figs. 29-30, mandibles-A and C, left, dorsal and ventral

aspects, respectively, B and D-right, dorsal and ventral aspects, respectively, of: 29, Cymindoidea (sensu stricto) indica

Schmidt-Goebel; 30, Apenes (sensu stricto) lucidula Dejean. Figs. 31 and 32, right maxilla, ventral aspect of: 31, C.

indica-, 32, A. lucidula. Figs. 33-36, labium, ventral aspect of: 33, C. indica-, 34, A. lucidula-, 35, A. (Sphalera) species; and

36, A. (Sphalera) postica (Dejean). Fig. 37, C. indica-. head, microsculpture, dorsal aspect. Scale bars = 100 nm. Legend,

mandibles: m, molar; pm, premolar; prt, posterior retinacular tooth; rr, retinacular ridge; tm, terebral margin; tt, terebral

tooth; vg, ventral groove.

Cymindine Lebiini of Authors

123

Figs. 38-40. Photographs of Apenina.— Figs. 38-39: SEM photographs of ovipositor, left stylomeres. Fig. 38: Cymindoidea

(sensu stricto) indica Schmidt-Goebel, stylomeres 1 and 2, lateral aspect, Fig. 39: Apenes (sensu stricto) lucidula Dejean:

A and B, lateral and apico-ventral aspects, respectively,. Scale bars = 50 Fig. 40: Cymindoidea (Habutarus) papua

(Darlington), habitus, dorsal aspect (SBL = 4.81 mm.). Legend, stylomeres: a, lateral ensiform seta; SI, stylomere 1; S2,

stylomere 2; x, projection of stylomere 1 .

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124

Ball and Hilchie

Abdomen. Sterna II-VII average. Female: tergum VIII broadly membranous medially; sternum VIII broadly

membranous medially, lateral apodemes short; tergum X completely sclerotized.

Male genitalia. Median lobe cylindrical, elongate, slightly curved ventrally. Apical portion slender, without

projections, quite short, but varied in length; anopic, orifice either dorsal, or dorso-lateral, toward left side. Internal sac

with vestiture of smaller (or larger) microtrichia; with or without long, coiled, flagellum. Left paramere average for

lebiomorph males; right paramere, though reduced, large for lebiomorphs.

Ovipositor (Figs. 38 - 39A, B). Valvifer markedly transverse, narrow. Stylomere 1 about twice length of stylomere

2, ventral apical angle markedly produced beyond base of stylomere 2, asetose; stylomere 2 with base extended dorsally

as lobe; apical portion sword-like, apex pointed; two very large ensiform setae dorsally; ventral surface with two rows of

sensory pits; without ventral preapical sensory furrow and associated setae.

Classification. — Included in the Apenina are three genera: Apenes LeConte (subgenus

Apenes and Sphalera Chaudoir); Cymindoidea Castelnau (subgenus Cymindoidea, Platy tarns

Fairmaire, and Habutarus, new subgenus); and Trymosternus Chaudoir. Reduction of the

oblongum cell of the hind wing is an autapotypic feature establishing monophyly of the New

World genus Apenes. Monophyly for the Old World assemblage of Trymosternus and

Cymindoidea sensu lato is established by an autapotypic feature of the interal sae of male

genitalia: possession of a moderately to very long and coiled flagellum. In the Old World

assemblage, monophyly of Trymosternus is established by a eombination of: integument

generally setose, and labial palpomere 2 plurisetose.

We have not been able to establish monophyly of Cymindoidea, for we have not identified

synapotypic features for all three subgenera. Cymindoidea and Platytarus are linked by a

quadripunctate glossal sclerite, broadened pronotum, and rugose dorsum. We could make

Cymindoidea monophyletic by including in it Trymosternus, but we suspect this decision would

not be acceptable to our European colleagues, who seem generally to prefer retention of

traditionally recognized taxa, in spite of phylogenetic considerations. We could also achieve the

desired result by exluding Habutarus, but this would require establishment of a monobasic

genus, and we are reluctant to do this. The compromise (whieh yields a cladisitcally

unacceptable genus) is to include Habutarus in Cymindoidea on the basis of a symplesiotypic

feature: the glabrous integument.

Geographical distribution. — This subtribe has a Gondwanian distribution pattern, with a

sister group on each side of the Atlantic Ocean, mainly in the Southern Hemisphere and

tropies.

Key to Genera and Subgenera of Subtribe Apenina

1 (0 ) Dorsum setose. Eyes reduced, not visible in ventral aspeet. Antennal fossa

remote from anterior margin of eye. Elytron with humerus sloped.

Metepisternum quadrate, wings represented by short stubs. Metasternum with

deep pit near middle coxae Trymosternus Chaudoir, p. 128

V Dorsum glabrous, except for normal fixed setae. Eyes various. Antennal fossa

close to or remote from anterior margin of eye. Elytron with humerus broadly

rounded. Metepisternum and wings various. Metasternum without pit near

middle coxae 2.

2 (E) Glossal selerite with four setae. Dorsal surface modified, either coarsely and

irregularly sculptured and punetate, or mieroseulpture with lines deep,

sculpticells convex, luster dull, and diseal elytral intervals keeled 3.

2' Glossal selerite with two setae. Dorsal surface unmodified, smooth, elytral

intervals more or less flat 4.

3 (2 ) Microsculpture of thoraeie pleura and sterna with meshes isodiametric.

Cymindine Lebiini of Authors

125

Antennomeres 1 and 2 generally setose. Elytron with odd-numbered intervals

carinate. Fossa of antenna and anterior margin of adjacent eye separated by

wide gap C. (Platytarus) Fairmaire.

3' Microsculpture meshes of thoracic pleura and sterna transverse. Antennomeres

1 and 2 with setae near apices, only. All elytral intervals non-carinate. Antennal

fossa and anterior margin of adjacent eye close together

C. (Cymindoidea) Castelnau p. 126

4 (2') Metepisternum quadrate, wing represented by short stub. Dorsum of head with

irregular shallow grooves and irregular ridges. Range- New Guinea

C. (Habutarus) new subgenus, p. 127

4' Metepisternum elongate, wing long, normally developed. Head with dorsum

smooth or ridged. Range- Neotropical and southern Nearctic Regions 5.

5 (40 Head with dorsum ridged or coarsely punctate

A. (Apenes) LeConte, p. 125

5' Head with dorsum smooth, not punctate or ridged

A. (Sphalera) Chaudoir.

Apenes LeConte

Figs. 30, 32, 34-36, and 39

Apenes LeConte, 1851: 174. GENERITYPE: Cymindis lucidula Dejean, 1831:320 (subsequent designation, by

Motschulsky, 1 864: 240, table).- LeConte, 1 861: 24.- Chaudoir, 1 875: 21, 35.- Horn, 1881:1 56.- 1 882: 1 56.- Bates, 1883:

188.-Blatchley, 1910: 147, 154.- Ball, 1960: 161.- Lindroth, 1969a: 1087.- Reichardt, 1977:443

Sphenopalpus Blanchard, 1853: 32. GENERITYPE: Sphenopalpus parallelus Blanchard, 1853: 32 { = Cymindis

aenea DeJean, 1831: 319) (monotypy):- Chaudoir, 1871: 385.

Sphenopselaphus Gemminger and Harold, 1868: 299. Unjustified emendation of Sphenopalpus.

Nominus Motschulsky, 1864: 240 (table). GENERITYPE: Cymindis punctulata Dejean, 1831: 316 { = Cymindis

sinuata Say, 1823: 8) (original designation by Motschulsky, 1864: 240, table).- Chaudoir, 1875: 42.

Malisus Motschulsky, 1864: 240 (table). GENERITYPE: Cymindis variegata Dejean, 1825: 217 (original

designation).

Didymochaeta Chaudoir, 1875: 50. GENERITYPE: Didymochaeta hamigera Chaudoir, 1875: 53 (here designated).

Sphalera Chaudoir, 1875: 54. GENERITYPE: Cymindis postica Dejean, 1831:317 (monotypy). NEW

SYNONYMY.

Notes about synonymy.— Chaudoir (1875) recognized four genus-group taxa that we

include in Apenes: Apenes (sensu stricto); A. {Malisus Motschulsky); Didymochaeta Chaudoir,

1875; and Sphalera Chaudoir, 1875. Bates (1883: 189) synonymized the first three names

because the taxa were based on “slight characters {Malisus) on general form and facies,

{Didymochaeta) on the narrow ligula and tooth of mentum”. To these names, we add Sphalera

Chaudoir, this taxon being based on absence of a mental tooth (Fig. 36). This feature involves a

minor desclerotization. Otherwise, adults are strikingly like those included in Didymochaeta.

For the atypical subgenus, we choose the name Sphalera (rather than Didymochaeta)

because the former has fewer letters, and is thus easier to write, if not to remember.

Recognition. — Adults of this genus are distinguished from other apenines by the following

combination of character states: glossal sclerite with a single pair of setae, dorsum glabrous,

metepisternum longer than wide, hind wings normally developed, metasternum smooth, without

a pit near the middle coxae. Additionally, males are distinguished by lack of a flagellum of the

internal sac.

Description. — Character states mostly as for subtribe, with restrictions or exceptions as follows.

Microsculpture. Dorsum of head with meshes isodiametric. Pronotum and elytra with meshes isodiametric or

transverse.

Quaest. Ent., 1983, 19 (1,2)

126

Ball and Hilchie

Vestiture. Surface generally glabrous. Antennomeres 1 and 2 with setae confined to apex, antennomere 3 with setae

confined to apical 0.50.

Head. Sub-antennal ridge average. Eyes orbicular, prominent, ventral margin rounded. Antennal fossa close to

antero-ventral margin of eye. Flagellar antennomeres distally longer than wide or length and width subequal, and

antenna short.

Mouthparts. Labium: mentum with lateral lobes broadly rounded or pointed apically; tooth absent or present and

bluntly or sharply pointed; glossal sclerite bisetose; palpomere 2 bisetose; palpomere 3 slightly to markedly seeuriform.

Thorax. Pronotum with sides rounded, or sinuate posteriorly; base lobate medially. Metepisternum distinctly longer

than wide.

Elytra. Interneurs average. Intervals slightly convex.

Wings. Well developed: oblongum cell shortened (stalked) or absent.

Male genitalia. Internal sac without coiled flagellum.

Classification. — The species of Apenes are here grouped into two subgenera: Apenes {sensu

stricto), including the species of Malisus), adults larger, body thicker, more terete, with head

grooved or coarsely punctate, and oblongum cell of wing stalked; and Sphalera (including

Didymochaeta), adults smaller, flatter, with head smooth (frontal impressions extended

diagonally to anterior supraorbital setigerous punctures), and wings without oblongum cell.

Phylogenetic considerations. — External features of adults of subgenus Sphalera seem more

plesiotypic, but absence of the oblongum cell from the wing, and absence of a mental tooth are

apotypic features. Conversely, adults of Apenes (sensu stricto) seem more derived in body form,

but retain the oblongum cell. The more sculptured integument characteristic of Apenes (sensu

stricto) adults is shared with adults of the Old World Cymindoidea (sensu stricto) and

subgenus Platytarus. This similarity is probably convergent.

Geographical distribution. — The range of Apenes extends from northern Argentina in

South America, to southern Ontario in eastern North America.

Cymindoidea Castelnau

Figs. 29,31,33, 37, 38, and 40

Cymindoidea Castelnau, 1832: 390. GENERITYPE: Cymindis bisignata Dejean, 1831: 322 (monotypy).- Andrewes,

1930: 140-141.- Basilewsky, 1961a: 154.-Csiki, 1932: 1490.- Jedlicka, 1963:462.

Philotecnus Mannerheim, 1837: 42. GENERITYPE: Philotecnus stigma Mannerheim, 1837: 42 ( = Cymindis

bisignata Dejean) (monotypy).

Platytarus Fairmaire, 1850, XVII (Bull.), XVII. GENERITYPE: Cymindis famini Dejean 1826: 447. (original

designation).- Basilewsky, 1961a: 165.- Antoine, 1962: 554.- Jedlicka, 1963:463.

Notes about synonymy. — Basilewsky (1961a: 154 and 165-166) provided relatively recent

listings of references to the above genus-group names. Reasons for including Cymindoidea

(sensu stricto) and Platytarus in the same genus are given under “Classification”.

Recognition. — Adults of this genus are distinguished from those of Trymosternus by the

glabrous dorsum and unmodified metasternum. Additionally, adults of subgenus Platytarus

(the only group partially sympatric with Trymosternus) have four glossal setae, and flatter

eyes. Adults of the Papuan subgenus Habutarus are like those of the New World subgenus

Apenes, but the two groups are distinguished not only on the basis of wing development (see

key) and geographical distribution, but males of Habutarus have a long flagellum in the

internal sac that is characteristic of Cymindoidea.

Description. — Character states mostly as described for subtribe, with restrictions and exceptions as follows.

Head. Frons and vertex with longitudinal ridges and grooves, irregularly rugose (Fig. 37); with or without prominent

supraocular ridges. Subantennal ridge prominent. Eyes orbicular or flattened, and longitudinally oriented; ventral margin

straight or curved. Temples well developed. Antennal fossa close to or remote from anteroventral margin of eye. Flagellar

antennomeres longer than wide.

Mouthparts. Maxilla: lacinia (Fig. 31) without setae on lateral preapical margin, few setae on ventral surface; mentum

(Fig. 33) with lateral lobes pointed apically, tooth well developed, pointed apically. Glossal sclerite (Fig 33) with two or

four setae, for latter condition, median pair very close together basally; palpomere 3 markedly securiform, maximally so in

Cymindine Lebiini of Authors

127

males.

Thorax. Metathorax normal, or reduced, with metepisternum quadrate.

Wings. Well developed, with oblongum cell not reduced (Figs. 25A, B), or brachypterous.

Legs. Spines of tibiae reduced.

Male genitalia. Internal sac with long coiled flagellum.

Classification. — Although Jeannel (1949: 947) included Platytarus in the subfamily

Calleiditae on the basis of reduced tibial spines of adults, other character states show that the

group is correctly placed near Cymindoidea - where it was placed by previous authors. In fact,

the only character states separating the two groups seem neither sufficiently numerous nor

sufficiently important (they involve form and surface sculpture only) to accord generic rank to

these groups. On the other hand, with antennae shifted forward, eyes flatter and seemingly

more protected by the rest of the head, the body generally narrower and deeper, we believe that

the species of Platytarus occupy an ecological zone rather different from that occupied by the

species of Cymindoidea {sensu stricto). On this basis, we accord subgeneric rank to these

groups.

Adults of the new taxon Habutarus, described below, are superficially strikingly different

from those of Cymindoidea and Platytarus. Nonetheless, they have the basic attributes of

Cymindoidea, and we prefer to emphasize similarities rather than differences. We do this by

including Habutarus in Cymindoidea (sensu lato).

Identification of species. — Andrewes (1935: 202-204) provides keys to adults of the species

of Cymindoidea {sensu stricto) and the subgenus Platytarus. Basilewsky (1961a) provides keys

to adults of the African species of Cymindoidea (pp. 155-158) and Platytarus (pp. 166-168).

Material examined. — We have seen adults of the following: Cymindoidea (sensu sthcto}- 19

specimens (two dissected; CAS), representing four Afrotropical and four Oriental species; Platytarus - 41 specimens (two

dissected, CAS), representing four species; and Habutarus - 17 specimens (three dissected, MCZ), representing C. papua

(Darlington), all paratypes, from Dobodura, Papua, New Guinea.

Geographical distribution.— The range of this genus is discontinuous: Cymindoidea (sensu

stricto) and Platytarus are widespread in Africa and the Oriental Region, with the range of

Platytarus extended eastward to Indo-China and northward to Hong Kong, and that of

Cymindoidea only as far as Burma (Jedlicka, 1963: 462-463); Habutarus is known only from

New Guinea, that is, the northern part of the Australian Region. Species of Cymindoidea

(sensu lato) have not previously been recorded from the Indo-Australian Archipelago.

Habutarus, new subgenus

Fig. 40

GENERITYPE: Nototarus papua Darlington, 1968: 186 (monotypy; here designated).

Derivation of name. — From the surname of Dr. Akinobu Habu; and "‘'tarus'', one of the

junior synonyms of Cymindis, and a name used in various combinations for cymindine-like

forms. Features of the ovipositor provide the principal clue to determining the correct location

of this taxon. Dr. Habu emphasized the importance of features of this structure in classification

of Lebiini, and so we are pleased to dedicate this subgenus to him, in recognition of his

contribution.

Recognition. — Adults of the single species included here resemble those of the Australian

calleidine subgenus Nototarus Chaudoir (see below), but as indicated above, they have the

basic attributes of the Apenina in general, and of Cymindoidea in particular.

Description. — Darlington (1968: 185-186) provides a good description of the type species of Habutarus. We draw

attention here to certain features that are useful in comparing this group with other members of Cymindoidea sensu lato.

Quaest. Ent., 1983, 19 (1,2)

128

Ball and Hilchie

Habitus as in Fig. 40. Body size small (SBL ca. 5. 5-6.0 mm.). Dorsal surface shining, lines of microsculpture fine,

meshes of elytra irregular, from isodiametric to slightly transverse. Eyes and temples like those of Cymindoidea (sensu

stricto), antennal fossae near anterior margins of eyes. Pronotum with base markedly narrower than maximum width,

hind angles acute; median longitudinal impression rather wide and deep. Metathorax reduced, metepisternum

subquadrate; brachypterous. Male genitalia and ovipositor average for Cymindoidea sensu lato.

Habitat. — Darlington (1968: 186) stated that adults of C papua were collected from flood

debris on rain forest floor.

Phylogenetic relationships. — Because of its plesiotypic character states (relatively

unmodified dorsal integument, glossal sclerite with single pair of setae, and pronotum cordate),

we believe that Habutarus must be closely related to the ancestral stock of Cymindoidea sensu

lato, and thus remote from the other extant species of this genus. Geographical remoteness

from the main range of the genus and reduced hind wings are also features suggesting a relict

status for this subgenus.

Trymosternus Chaudoir

Trymosternus Chaudoir, 1873: 106. GENERITYPE: Cymindis onychina Dejean, 1825: 217 (subsequent designation,

by Antoine, 1962: 559). Seidlitz, 1887: 8.- 1888: 8.- Bedel, 1906: 242.- lakobson, 1907: 396.- Csiki, 1932: 1486.-

Jeannel, 1942a: 1057.- 1949: 396.-Mateu, 1952: 109-141. 1958: 1-6.- Antoine, 1962: 559.

Recognition. — Adults of this genus are distinguished from other apenines by combination

of a markedly cordate pronotum, metasternum with a deep pit near middle coxae, short

(reduced) metepisternum, and generally setose integument.

Description. — Character states mostly as for subtribe, with restrictions and exceptions as follows. See Mateu

(1952: 1 1 1-113) or Antoine (1962: 559) for a more detailed description.

Color. Body piceous to rufo-piceous; elytra concolorous.

Vestiture. Surface generally coarsely punctate, setose, including mandibular scrobes and antennomeres 1-3.

Head. Frons laterally with pronounced ridge each side. Sub-antennal ridge prominent. Eyes oblong, flattened. Temples

prominent. Antennal fossa well in front of antero-ventral margin of eye. Flagellar antennomeres longer than wide.

Mouthparts. Labium: mentum with lateral lobes pointed apically; tooth acute at apex; glossal sclerite bisetose;

palpomere 3 distinctly securiform.

Thorax. Pronotum cordate, sides markedly sinuate posteriorly; base subtruncate, not lobed medially. Metepisternum

short. Metasternum with deep pit anteriorly, near middle coxae.

Elytra. Humeri distinctly narrowed. Interneurs average, though coarsely punctate. Intervals slightly convex.

Wings. Reduced to short stubs.

Legs. Middle and posterior tibiae with reduced spines, latter absent from lateral margins.

Male genitalia. Internal sac with coiled flagellum.

Notes about identification of species. — See Mateu (1952).

Material examined. — Three specimens (CAS): Trymosternus onychinus (Dejean), male; and T.

cordatus Rambur, male and female.

Geographical distribution. — The range of the 10 species of this genus is confined to the

mountains of the Iberian Peninsula and to North Africa north of Morocco and Oran (Mateu,

1952, 1958; Antoine, 1962). Only one polytypic species (T. truncatus Rambur) occurs in North

Africa, and in that part of Spain immediately adjacent to Gibraltar. The other nine species are

on the mainland, most of them in southern Spain, and most with markedly restricted

geographical ranges. Trymosternus onychinus is wide-ranging (see Mateu, 1952: Fig. 4).

Phylogenetic considerations. — Antoine (1962: 560) regards this genus as highly evolved

and isolated. Certainly, body form resulting in part from wing loss and in part from the striking

lateral lobes of the pronotum exhibited by adults of some species, give this impression.

However, the bisetose glossal sclerite, relatively unmodified elytral intervals, and restricted

geographical range suggest that this genus is the survivor of an old stock. It was probably

isolated for an extended period on the Miocene betico-rifian massif (Antoine, 1962: 560),

where it differentiated. In post-Miocene time, it dispersed northward, attaining its present

Cymindine Lebiini of Authors

129

range (Mateu, 1952: 117).

Evolution of the Apenina: preliminary considerations. — We are not in position to address

this topic in detail, but some aspects of a general pattern seem clear enough to formulate a

preliminary hypothesis in the form of a scenario.

The ancestral stock of the extant taxa, whose adults were like those of Sphalera and

Habutarus, was widespread in Gondwana. Following the split which led to formation of South

America and Africa, and thus to division of the ancestral stock of Apenina, the New World

group differentiated as Apenes. In the Tertiary, various stocks dispersed northward,

differentiating to produce the complex of extant species that presently inhabit Middle and

North America and the West Indies.

In the Old World, events seem to have been more complex, for the distribution of extant

taxa seems to suggest at least two major episodes of evolution: an early one, represented by taxa

with limited ranges- Trymosternus (centered in the Iberian Peninsula), and Habutarus

(known only from New Guinea); and a later episode, represented by centrant groups

Cymindoidea and Platytarus. We believe that the present centrant groups overran the ranges

of the early-evolved taxa, displacing the latter from the central areas, and leaving only

peripheral remnants. This does not explain absence of species of Cymindoidea (sensu lato) from

the Indo-Australian Archipelago, but we expect that the group is represented there, though

specimens have not yet been collected.

If our hypothesis is correct, the main islands of the Indo-Australian Archipelago will be

populated by stocks of Cymindoidea (sensu stricto) or Platytarus. and the peripheral islands

(near New Guinea) by Habutarus. We also anticipate that the pattern we presently perceive

will not be altered by subsequent discoveries. However, if it is altered by discovery of additional

remnants of early-evolved groups in Africa or on the mainland of southeastern Asia, they will

be residents of high altitude forests, and their adults will be brachypterous.

Subtribe CYMINDINA

We have seen specimens representing seven taxa of this group that are currently ranked as

genera: Cymindis Latreille, 1806; Hystrichopus Boheman, 1848; Plagiopyga Boheman, 1848;

Pinacodera Schaum, 1857; Taridius Chaudoir, 1875; Pseudomasoreus Desbrochers des Loges,

1904; and Afrotarus Jeannel, 1949. We have not seen material of Assadecma Basilewsky,

1982, so our comments about it are based on study of the description and illustrations. In spite

of the rank accorded them, these taxa are not easily characterized on the basis of adult

features. In our opinion, they are over-ranked. Accordingly, we make in the following pages

adjustments in ranking that seem required by the evidence available.

We add to this subtribe a new monobasic genus, Ceylonitarus. Reasons for assigning this

rank are presented below.

Recognition. — Diagnostic features of the subtribe are: head without suborbital setigerous

punctures; elytron with penultimate umbilical puncture not laterad of antepenultimate and

ultimate punctures; scutellar interneur separate from interneur 1, base of latter evident; tibiae

average, spined laterally; tarsomeres slender, glabrous or setose dorsally, male front tarsomeres

moderately expanded, articles 1-3 with biseriate adhesive vestiture ventrally; tarsal claws

pectinate; ovipositor with stylomeres 1 and 2 subequal in length, stylomere 1 asetose; stylomere

2 without baso-dorsal projection, with one or two ensiform setae dorsally; preapical sensory

furrow reduced, with one or two nematoid setae, or without these, and without furrow pegs;

Quaest. Ent., 1983, 19 (1,2)

130

Ball and Hilchie

mentum toothed, labial palpomere 2 bi- or plurisetose; apical margin of palpomere 3

subtruncate, or fusiform.

Description. — Standarized Body Length between 4.5 and 7.5 mm. Form slightly varied, from about average for

Carabidae to somewhat flattened and broadened. Color mostly somber: dorsum rufous to black or metallic blue or green,

appendages of most adults same color as that of dorsum, or paler; elytra either concolorous (adults of most species), or

bicolored with various dark markings on paler background.

Microsculpture. Labrum and clypeus: with meshes isodiametric, microlines clearly visible at magnification of 50X.

Frons and vertex with meshes isodiametric or microlines effaced. Pronotum with meshes isodiametric to transverse, or

microlines effaced. Lateral and ventral sclerites of thorax with meshes transverse. Scutellum and elytra with meshes

isodiametric to transverse, or effaced. Abdominal sterna with meshes transverse.

Luster. Dorsum various, dull to shining (most adults), to slightly iridescent.

Standard or fixed setae. Average for lebiines: head with two (or three) pairs of supraorbital setae; submentum and

mentum each with single pair. Pronotum with two to six pairs of lateral setae, posterior pair near posterior angles.

Prosternum with several setae at apex of intercoxal process. Elytra each with two or three discal setae, in interval 3;

umbilical series continuous, 14 to 20 setae included, penultimate setigerous puncture not displaced laterally. Legs with

average setation for carabids: tibia with full complement of spines; tarsomere 5 with row of spines on each ventro-lateral

margin. Abdominal sterna with ambulatory setae, sternum VII with one or two pairs of setae in males, two pairs in

females.

Vestiture and surface. Integument smooth, glabrous, or more or less densely to sparsely punctate, punctures with long

or short slender setae; antennomeres 1-3 generally finely setose, or glabrous with apical ring of setae; tarsomeres dorsally

glabrous or finely setose.

Head. About average in form for carabids. Frontal impressions shallow, broad. Clypeus average, transverse, about

rectangular, anterior margins each slightly concave or truncate. Frons smooth, or rugulose (Fig. 50) laterally. Eyes

average, moderately convex to reduced and flattened. Antennae filiform, antennomere 3 longer than 2 and 4, or subequal

to latter articles; antennomeres each longer than wide, or width and length subequal and antenna shortened.

Mouthparts. Labrum like clypeus, in general form. Mandibles trigonal, average for carabids. Left mandible (Figs.

40. 2A, B, 41A, B, 42A, B, 43A, B, 44A, B) with terebral margin well developed or reduced (most species), cutting edge

retinacular ridge; posterior retihacular tooth small; premolar average, ventral surface with well developed premolar ridge;

molar ridge present or absent; ventral groove average, setose throughout length, or absent. Right mandible (Figs. 40. 2C,

D, 41C, D - 44C, D) similar in overall size and form to left mandible; terebral margin well developed, tooth small or

absent; retinacular ridge prominent or not, anterior retinacular tooth present or absent; premolar tooth present or absent,

premolar ridge well developed; molar ridge present or absent. Maxilla average in form; lacinia (Fig. 45) extensively setose

on ventral surface; galeomere 2 shorter than 1; palpomeres average, 4 fusiform, with apical margin subtruncate. Labium

(Figs. 46-49) average; mentum with well developed tooth, broad or pointed apically, epilobes average; glossal sclerite

broad, truncate and bisetose apically; paraglossae adnate to glossal sclerite, each paraglossa with short setae apically;

palpomere 2 bi-, or plurisetose; palpomere 3 fusiform, with apical margin subtruncate, or in males expanded, securiform.

Thorax. Pronotum transverse, subcordate to subquadrate, surface slightly convex; basal margin beaded, subtruncate to

distinctly lobed medially; anterior margin slightly concave; sides narrow to distinctly explanate; anterior angles broadly

rounded; posterior angles acute to broadly rounded; median longitudinal impression distinct; posterior-lateral impressions

shallow, indistinct. Prosternum with apex of intercoxal process immarginate. Pterothorax average, metepisternum

elongate, with lateral margin greater in length than anterior margin; or subquadrate, with lateral and anterior margins

subequal.

Legs. Average for Carabidae. Tarsomere 4 with apical margin subtruncate, not projected laterally as paired lobes;

tarsal claws smooth or pectinate (Figs. 51-54), three to seven denticles per claw, denticles either sharp (adults of most

species) or apices blunt. Male with front tarsomeres 1-3 (or 4) with adhesive vestiture ventrally; middle tarsomeres 1-4

without or with (adults of Pinacodera) adhesive vestiture.

Elytra. Average for lebiine adults: humeri broadly rounded (or sloped); apical margin subtruncate. Interneurs average,

finely punctate or not; intervals flat to slightly convex.

Wings. Developed normally, or reduced to short stubs; species monomorphic or dimorphic for wing condition. Venation

generally average for carabids: oblongum cell average (Figs. 73A, 74A, 84A, 85A); wedge cell (Figs. 73B, 74B, 84B, 85B)

evident, though more or less reduced.

Abdomen. Abdominal sterna II-VII average for carabids, or sternum VII of males with posterior margin more or less

deeply notched.

Male genitalia. Median lobe (Figs. 70-72; 86-88) cylindrical, anopic (Figs. 70-72) with apical orfice inclined to left

and dorsal surface otherwise sclerotized; or catopic (Figs. 86-88) dorsal surface completely sclerotized; apical portion

various, shorter or longer, narrow to very broad. Internal sac variously armored with vaguely defined fields of microtrichia,

and with or without large, curved apical sclerite. Parameres average for lebiomorphs: left broad, about 0.33 length of

median lobe; right short, but apex free, not fused to median lobe.

Ovipositor and associated sclerites. Tergum and sternum VIII average for lebiomorphs (as in Figs. 76A, B). Tergum X

(Fig. 76C) with sclerotization reduced medially. Valvifer very broad and short. Stylomere 1 broad, slightly wider than

long, asetose. Stylomere 2 (Figs. 55-61) as long or longer than stylomere 1, subcylindrical in form, without baso-dorsal

Cymindine Lebiini of Authors

131

TABLE 2

GEOGRAPHICAL DISTRIBUTION BY REGION, OF THE GENERA AND

SUBGENERA OF CYMINDINA

132

Ball and Hilchie

projection; ensiform setae one or two, longer (Fig. 55A) or shorter (Fig. 56A); trichoid setae few, ventral in position, or

absent; preapical sensory furrow narrow, with one or two short nematoid setae or without these, and without furrow-peg

setae; microsculpture (Figs. 62-65) almost isodiametric, more or less extensive; sculpticells with (Fig. 65) or without

microspines.

Classification. — Eight genus-group taxa are arranged in two genera: Cymindis Latreille,

and Hystrichopus Boheman. Taxa recognized by previous authors as subgenera of Cymindis

are thus accorded lesser rank. Jeannel (1942a: 1039) also recognized within his subfamily

Cyminditae two groups that correspond to the genera that we recognize: tribes

Pseudomasoreini and Cymindini. At the time, however, he did not realize the close affinity

between Psuedomasoreus, Hystrichopus, and Plagiopyga, and thus did not include the latter

two groups in the Pseudomasoreini. A third genus is Ceylonitarus.

Geographical distribution.- This subtribe is basically Megagean in distribution, with one

subgenus extending into the northern part of the Neotropical Region {Pinacodera; to

Honduras, in Central America). Table 2 provides a summary. Details are presented below.

Key to Genera and Subgenera of Cymindina

1 (0 ) Elytron with lateral umbilicate punctures not distinguishable from other serial

setigerous punctures of intervals. Dorsum generally punctate and setose, serial

setae of elytral intervals each more than half length of antennal scape. Head

with three pairs of supraorbital setigerous punctures. Tarsal claws smooth, not

pectinate. Elytra bicolored: flavous with black basal, medial, and apical fasciae

(Fig. 40.1). Specimen from Oriental Region . . . Ceylonitarus, new genus, p. 135

T Elytron with lateral umbilicate punctures distinctly larger than serial punctures

of discal intervals. Dorsum various, glabrous or more or less setose. Head with

two pairs of supraorbital setigerous punctures. Tarsal claws smooth or pectinate.

Elytral color various. Specimen from Megagaea, or from the northern

Neotropical Region 2.

2 (T) Specimen from locality in Nearctic or Neotropical Region 3.

2' Specimen from locality in Palaearctic, Oriental, or Afrotropical Region 4.

3 (2 ) Dorsum glabrous or more or less densely setose. Male with tarsomeres 1-4 of

both front and middle legs slightly widened, ventrally with adhesive vestiture C.

(Pinacodera) Schaum, p. 149

3' Dorsum densely setose. Male with tarsomeres 1-3 of front legs only slightly

widened, ventrally with adhesive vestiture

C. (Cymindis) Latreille(part), p. 156

4 (30 Dorsum with vestiture of short setae, more or less densely punctate, or at least

intervals 4-7 each with several irregular rows of punctures; dorsal surfaces of

tarsomeres sparsely to densely setose 5.

4' Dorsal surface glabrous, elytral intervals impunctate, dorsal surfaces of

tarsomeres sparsely setose or glabrous 6.

5 (4 ) Frons with two sharply defined longitudinal ridges each side. Integument

piceous; lateral margins of pronotum and eltyra rufo-flavous. Pronotum with

broad lateral margins. Range- Indian sub-continent, south of the Himalaya . . .

C. (Afrotarus) Jeannel (part), p. 154

5' Frons laterally smooth, or with indistinct ridges. Integument various, of most

specimens rufous or rufo-piceous, and elytra with or without pale marks.

Cymindine Lebiini of Authors

133

Pronotum with lateral margins various. Range- Atlantic Islands, Africa north

of Atlas Mountains, Palaearctic Region, including upper slopes of the

Himalayan system

C (Cymindis) Latreille"* (part) p. 156

6 (4') Frons laterally smooth or irregularly, sparsely punctate; without two or more

regular ridges. Median lobe of male ca topic (Fig. 86 A). Stylomere 2 with single

ensiform seta (Figs. 55A-57), in basal half 7.

6' Frons each side with two or more regular ridges (Fig. 50). Median lobe of male

anopic (Fig. 69 A). Stylomere 2 of ovipositor with two ensiform setae, located in

posterior half 10.

7 (6 ) Mentum with pair of setae on tooth. Paraglossae glabrous. Antennomere 3

pubescent toward apex; internal sac of male genitalia with rows of small spines

H. (Assadecma) Basilewsky, p. 170

1' Mentum with setae on lateral lobes, only; paraglossae setose. Antennomere 3

pubescent toward apex, or nearly glabrous. Internal sac with or without spines 8.

8 (7') Antennomere 3 sparsely pubescent toward apex; denticles of tarsal claws sharp

(Fig. 52); interval 3 of elytron with two or three setigerous punctures; stylomere

2 of ovipositor with moderate to long ensiform seta (Fig. 5 5 A)

H. (Pseudomasoreus) Desbrochers des Loges, p. 158

8' Antennomere 3 not pubescent, with few long setae apically and preapically;

tarsal claws smooth, or with sharp or blunt denticles; interval 3 of elytron with

three or more setigerous punctures; stylomere 2 of ovipositor with ensiform seta

very short (Figs. 56A and 57) 9.

9 (8') Interval 3 of elytron with four or more setigerous punctures; tarsal claws with

denticles sharp (as in Fig. 52)

H. (Hystrichopus) Boheman, p. 171

9' Interval 3 of elytron with two or three setigerous punctures; tarsal claws smooth,

or with denticles blunt (Fig. 54)

H. (Plagiopyga) Boheman^ p. 172

10 (6') Vertex and frons with isodiametric meshes; metepisternum with lateral margin

distinctly longer than basal width, macropterous; pronotum with sides

explanate; antennomeres 4-10 each distinctly longer than wide; internal sac of

male genitalia (Fig. 69A) without large sclerite

C. (Taridius) Chaudoir, p. 145

10' Vertex and frons smooth medially, microlines effaced; metepisternum with

lateral and basal margins subequal, brachypterous; sides of pronotum narrow;

antennomeres 4-10 each 2.0 longer than wide, or shorter, not more than 1.5

longer than wide; internal sac of male genitala (Figs. 70A, 71 A) with large

'‘According to Antoine 1962: 567), adults of all species of subgenus Cymindis (as delimited

here) have setose elytra, although in some species the setae are very short and sparse. In any

event, glabrous- appearing specimens of subtribe Cymindina occurring to the north of the

Pyrenees Mountains in Western Europe are members of subgenus Cymindis

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Ball and Hilchie

Figs. 40.1-40.2. Photographs of Cymindina, Ceylonitarus ceylonicus, new species. Fig. 40.1: habitus, dorsal aspect (SBL =

5.92 mm.). ; Figs. 40.2A-E: SEM photographs of mandibles, A and D, left, dorsal and ventral aspects, respectively, B and

E, right, ditto; C, basal brush of left mandible, dorsal aspect. Legend, mandibles: art, anterior retinacular tooth; bb, basal

brush; m, molar; part, posterior retinacular tooth; rr, retinacular ridge; tm, terebraf margin. Scale bars= Figs. 40.1 = 1.0

mm.; Figs. 40. 2A, C, D, and E = 200 nm; Fig. 40. 2B = 20

Cymindine Lebiini of Authors

135

sclerite

C. (Afrotarus) Jeannel (part), p. 154

Ceylonitarus new genus

Figs. 40.1-40.6

GENERITYPE: Ceylonitarus ceylonicus, new species (here designated).

Derivation of name. — From the former name of the type area (“Ceylon”), and ''tarus'\ a

name used in various combinations for Cymindis- like forms.

Recognition. — Diagnostic features of this taxon are: habitus Cymindis- like: color of elytra

flavous, with three black fasciae (Fig. 40.1); body generally setose, setae long (Figs. 40. 5A and

B), head with three pairs of supraorbital setae; serial setigerous punctures of discal intervals of

elytron as large as lateral umbilicate punctures, latter not readily distinguishable by eye; frons

laterally shallowly and irregularly grooved; mandibles without ventral grooves (Figs. 40.2E and

D), with large basal brushes (Fig. 40. 2B); left mandible with well developed terebral margin

(Fig. 40. 2A); right mandible without premolar tooth (Figs. 40. 2C and E); tarsal claws smooth;

stylomere 2 of ovipositor narrow at base, falcate, with single long ensiform seta, dorso-lateral in

position (Fig. 40. 6A), ventral sensory furrow well removed from apex (Fig. 40.6B), without

nematoid or furrow-peg setae, and microsculpture meshes of blade transverse, most sculpticells

terminated apically with a microspine (Fig. 40. 6C).

Classification. — Consideration in evolutionary terms of character states of Ceylonitarus

compel us to rank this taxon as a genus, though we are reluctant to recognize monobasic

genera. The following character states suggest that this group is more primitive that either

Cymindis or Hystrichopus: left mandible with well developed terebral margin (Fig. 40. 2A);

maxilla with lacinia sparsely setose ventro-apically (Fig. 40.3; cf. Fig. 45); mental tooth not

ridged (Fig. 40.4; cf. Figs. 46-49); terminal palpomeres narrow apically (Fig. 40.3; cf. Fig. 46);

and stylomere 2 narrow and falcate. The extensive wedge cell of the wing is symplesiotypic for

Ceylonitarus and Hystrichopus.

The following autapotypic features testify to the distinctness of the group: gains- an extra

pair of supraorbital setae, additional antero-lateral marginal setae of pronotum, and the

generally setose body; large basal brushes of the mandibles (Fig. 40. 2B), and acute apices of

lateral lobes of the mentum (Fig. 40.4); losses- ventral grooves, from mandibles; medial

ensiform seta, nematoid setae, and furrow-peg setae, from stylomere 2.

The following apotypic features are shared with other cymindine taxa, but we believe they

were acquired independently in each lineage: color pattern of elytra (shared with some

members of subgenus Taridius); absence of microsculpture from most of dorsal surface, setose

condition of body, and reduced basal ridge of elytron (shared with some members of Cymindis

sensu stricto); stylomere 2 with single ensiform seta, dorso-lateral in position (shared with

females of Hystrichopus sensu lato), and ensiform seta longer than usual (shared with females

of subgenus Pseudomasoreus).

The smooth tarsal claws are difficult to interpret, for they may be primitively smooth, and

thus plesiotypic, or secondarily smooth, and thus apotyic. In any event, this character state is

shared with some members of the subgenus Plagiopyga.

The pattern of shared features suggests that they are examples of convergence, rather than

of close relationship. Further, the plesiotypic features plus probably restricted geographical

range that is peripheral to that of other cymindine groups, suggest to us that Ceylonitarus is a

Quaest. Ent., 1983, 19 (1,2)

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Ball and Hilchie

Cymindine Lebiini of Authors

137

Figs. 40.3-40.6. SEM photographs of structures of Cymindina, Ceylonitarus ceylonicus, new species.— Fig. 40.3: Right

Maxilla, lacinia, palpifer, and palpus, ventral aspect. Fig. 40.4: Labium, ventral aspect, mentum and parts of prementum.

Fig. 40.5: Left elytron— A, basal portion, dorsal aspect, B, basal portion dorso-medial aspect, c, two interneural punctures,

dorso-medial aspect. Fig. 40.6: ovipositor, left stylomere 2— A, lateral aspect; B, ventral aspect; C, apical portion, ventral

aspect. Legend, maxilla: la, lacinia. Legend, labium: el, epilobe; 1, glo sal sclerite; m, mental tooth; pg, paraglossa. Legend,

elytra: br, basal ridge. Legend, ovipositor, stylomere 2: a. lateral ensiform seta; c, sensory furrow; e, trichoid setae; m,

microspine of sculpticell. Scale bars= Figs. 40. 3-40. 5B = 200 /um; Figs. 40.5C-40.6C = 20 ;xm.

Quaest. Ent., 1983, 19 (1,2)

138

Ball and Hilchie

phylogenetic relic, closer to the ancestral stock of the Cymindina than are the other two genera

of this subtribe. The several and striking autapotypic features suggest the possibility of an

extended period of isolation and probably of ethological (as well as structural) divergence from

the other groups of cymindines.

Males are unknown. Features of the male genitalia might shed light on relationships of

Ceylonitarus: if catopic, this would suggest close relationship with Hystichopus\ if anopic, and

with an elaborate apical sclerite on the internal sac, this would suggest close relationship with

Cymindis. We believe that the median lobe will prove to be anopic, and the internal sac either

devoid of armature, or with armature that is strikingly different from that of the taxa of

Cymindis.

We think it possible that Ceylonitarus may include additional species. If so, the character

states of such taxa might provide clues that will make possible a better assessment of the

phylogenetic relationships of this group, and thus provide a better basis for its classification.

Description. — Size small (SBL less that 7.0 mm.), body slender, habitus as in Fig. 40.1. Color fuscous to flavous,

with elytra markedly bicolored.

Microsculpture. Labrum with meshes isodiametric, sculpticells slightly convex. Rest of body surface without

microlines (at least not visible at SOX; very faint vestiges on dorsum seen at lOOOX), surface essentially smooth.

Fixed setae. Standard, except: head with three pairs of supraorbital setae; pronotum with several pairs of marginal

setae in anterior 0.5. Abdominal sternum VII with four setae apically.

Punctation and vestiture. Scape and antennomeres 2-4 generally setose, antennomeres 5-11 more densely so. Eyes

glabrous. Head, prothorax, and lateral and ventral sclerites of pterothorax densely and moderately coarsely punctate,

abdominal sterna more sparsely and finely so. Elytral interneurs more finely punctate than intervals; latter uniseriately

punctate, setae flavous, many more than 0.5 length of antennal scape; serial punctures of discal intervals as large as

umbilicate punctures, latter not readily distinguished by eye.

Head. Frons each side with two or three rather indistinct and irregular ridges. Frontal impressions shallow, indistinct.

Eyes moderately prominent (Fig. 40.1). Antennae with antennomere 3 slightly longer than 4, but shorter than scape.

Mouthparts. Labrum larger than average (1/w 0.59-0.68, x>=0.62). Mandibles as in Figs. 40.2A-E, both without

premolar and ventral groove (Figs. 40.2D-E), and with large basal brush (Fig. 40. 2B); left mandible with well developed

terebral margin (Fig. 40. 2A); right mandible with prominent retinacular ridge and small retinacular teeth (Figs. 40. 2C

and E). Maxillae average, lacinia ventrally with setae near medial margin, not extended to lateral margin pre-apically

(Fig. 40.3), terminal palpomere fusiform, apex narrow. Labium (Fig. 40.4): mentum with lateral lobes acute apically;

epilobes slender throughout, not toothed medially, and terminated at base of mental tooth; latter prominent, broadly

rounded and immarginate apically; glossal sclerite broad, rounded apically; paraglossae membranous apically, setose, with

apices extended slightly beyond plane of apex of glossal sclerite.

Thorax. Pronotum as in Fig. 40.1 (for details, see description of generitype). Ventral and lateral thoracic sclerites

without notable features. Metepisternum with lateral margin clearly longer than basal margin.

Elytra. Dorsal surface deplanate. Base with humerus broadly rounded (Fig. 40. 5A). Apical margin truncate and

markedly sinuate. Basal ridge close to anterior margin, terminated near base of interval 5. Interneurs shallow, punctate.

Epipleuron average.

Wings. Fully developed, veins rather pale and probably slightly sclerotized. Oblongum with short stalk (cf. Figs. 73 A,

74A, and 84A), wedge cell more extensive than in Cymindis (cf. Figs. 73B and 74B).

Legs. Average for Cymindina, except tarsal claws smooth.

Ovipositor. Stylomere 1 glabrous, stylomeres 1 and 2 subequal in length. Stylomere 2 falcate in lateral aspect (Fig.

40. 6A), slender, parallel-sided in ventral aspect, apex narrowly pointed. Lateral ensiform seta (Fig. 40. 6A) longer than

average, about half length of stylomere; several trichoid setae medio-ventrally. Following setae lacking: medial ensiform,

nematoid, and furrow-peg. Sensory furrow (Fig. 40. 6B) very narrow, about half way between apex and plane of insertion

point of ensiform seta. Microsculpture: meshes generally transverse, broadly so basally (Fig. 40. 6A), more narrowly so

preapically (Figs. 40. 6B and C); sculpticells flat basally, convex preapically, though not keeled, most terminated with

single microspine (Fig. 40. 6C).

Relationships of genus. — We believe that Ceylonitarus is more primitive than Cymindis or

Hystrichopus, and is the sister group of the ancestral stock of these two genera. See

“Classification”, above, for a discussion of the basis for this hypothesis.

Included species. — Only one, C. ceylonicus, new species, described below.

Cymindine Lebiini of Authors

139

Ceylonitarus ceylonicus, new species

Type material. — HOLOTYPE female, labelled: SRI LANKA Man Dist. 8 mi. SE Mannar black light 15 feet, 6

Nov. 1976; Collected by G. F. Hevel, R. E. Dietz IV, S. Karunaratne, D. W. Balasooriya (USNM). Seven paratypes,

females, labelled: SRI LANKA Man. Dist. 4 mi. NW Mannar black light, 100 ft. 3 November 1976; collector label same

as for holotype (USNM). TYPE LOCALITY: vicinity of Mannar, Sri Lanka.

Derivation of specific epithet. — From the former name of the type area, Ceylon.

Recognition. — Color pattern (Fig. 40.1) of adults of this species is like that of the mainland

species Cymindis (Taridius) stevensi, known from the Nilgiri Hills of India. The two are easily

distinguished, however, by differences in: setation (adults of C. stevensi with dorsal integument

glabrous except for normal fixed setae; adults of C. ceylonicus with dorsal integument generally

setose); microsculpture (pronotum and elytra of C. stevensi with meshes distinct; these surfaces

smooth in C. ceylonicus)-, details of color pattern of elytra (cf. Figs. 40.1 and 75B); pronotum

(sides explanate in C. stevensi adults, not so in members of C. ceylonicus)-, tarsal claws

(pectinate in C. stevensi-, smooth in C. ceylonicus)-, and in setae of stylomere 2 (C. stevensi

females with median and lateral ensiforms, nematoids, and furrow-pegs; C. ceylonicus females

with only lateral ensiform).

Description. — Habitus as in Fig. 40.1, Cymindis- like. Standardized Body Length 5.36-6.64 mm. (x = 6.02 mm).

Color. Head and pronotom dorsally rufo-piceous to piceous, rufous ventrally. Elytra with disc predominantly flavous,

with suture dark, and three black fasciae (Fig. 40.1); epipleura flavous. Metepisterna and abdominal sterna rufo-piceous,

other sclerites of pterothorax rufous; antennae, palpi, and legs flavous.

Microsculpture, setation, form of head, details of mouthparts, thorax (except pronotum), elytra, legs, abdominal sterna

and ovipositor sclerites as described for genus, above.

Pronotum. As wide as or slightly wider than head (Hw/Pwm 0.93-1.00, x = 0.96), slightly wider than long (Pl/Pwm

0.83-0.91, x = 0.87), width near mid-line greater than width at base (PwB/Pwm 0.67-0.73, x = 0.71). Sides narrrow, not

explanate, sharply beaded, markedly sinuate. Anterior lateral angles broadly rounded; posterior-lateral angles rectangular

or acute, distinctly anterior to medial part of basal margin Basal margin not beaded, laterally with short, marked

sinuation. Disc markedly convex medially, sloped downward laterally. Marginal grooves narrow, continuous with narrow

posterior-lateral impressions. Median longitudinal impression shallow; anterior transverse impression indistinct; posterior

transverse impression broad, shallow, continuous with posterior-lateral impressions.

Geographical distribution. — Known only from low altitude localities, in the vicinity of

Mannar, Sri Lanka, where the specimens were taken at light.

Material examined. — Type series, only. We owe a special note of thanks to Terry Erwin,

who drew these specimens to our attention, and made them available for our study.

Cymindis Latreille

Figs. 41-43,45-51, 53, 59-63, 65, and 67-76

Cymindis Latreille, 1806: 190: GENERITYPE: Buprestis humeralis Fourcroy, 1785:57 (monotypy).- Chaudoir,

1873: 53-120.- Csiki, 1932: .- Jeannel, 1942a: 1041-1056.- Antoine, 1962: 564-587.- Jedlicka, 1963: 452-461.- Habu,

1967: 57-74.- Lindroth, 1969: 1070-1086.- Ball, 1982: .

Pinacodera Schaum, 1857: 294. GENERITYPE: Cymindis limbata Dejean, 1831:320 (designated by Lindroth, 1969:

1067).- LeConte, 1861: 24.- Chaudoir, 1875: 2.- Horn, 1881: 156.- 1882: 146.- LeConte and Horn 1883: 45.- Bates,

1883: 187-188.- 1884: 296.- Blatchley, 1910: 142, 152.- Leng, 1920: 67.- Casey, 1920: 279.- Csiki, 1932: 1487.-

Blackwelder, 1944: 62.- Jeannel, 1949: 878.- Ball, 1960: 161.- Lindroth, 1969: 1067-1070. Erwin et ai, 1977: 4.58.- Ball,

1982: . NEW SYNONYMY.

Planesus Motschulsky, 1864: 240(table). GENERITYPE: Cymindis fuscata Dejean, 1831:321 { = Cymindis

platicollis Say, 1823) (original designation by Motschulsky, 1864: 240 (table)).

Taridius Chaudoir, 1875: 71. GENERITYPE: Taridius opaculus Chaudoir, 1875: 7 (monotypy).- Bates, 1892: 416).-

Andrewes, 1930: 342-343.- Csiki, 1932: 1489.- Andrewes, 1935: 204-205.- van Emden, 1937: 123-125.- Jedlicka, 1963:

461.- Ball, 1982:—. NEW SYNONYMY.

Afrotarus Jeannel, 1949: 878. GENERITYPE: Cymindis kilimana Kolbe, 1898: 51 (original designation).-

Basilewsky, 1962: 252.- 1968a: 360.- Ball, 1982:-—. NEW SYNONYMY

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140

Ball and Hilchie

Figs. 41-44. SEM photographs of structures of Cymindina. — Mandibles -A and C, left, dorsal and ventral aspects,

respectively, B and D, right, dorsal and ventral aspects, respectively.— 41, Cymindis (sensu stricto) suturalis Dejean; 42,

C. (Taridius) opacula (Chaudoir); 43, C. (Pinacodera) new species no. 1; 44, Hystrichopus (sensu stricto) near dorsalis

Thunberg. Scale bars = 100 ^m. Legend: art, anterior retinacular tooth; m, molar; pm, premolar; prt, posterior retinacular

tooth; rr, retinacular ridge; tm, terebral margin; tt, terebral tooth; vg, ventral groove.

Cymindine Lebiini of Authors

141

Figs. 45-54. SEM photographs of structures of Cymindina.-Mouthparts and tarsal claws. Fig. 45: C. (Pinacodera) limbata

Dejean, right maxilla, ventral aspect. Figs. 46-49, labium, ventral aspect: 46, C. (P.) limbata-, 47, C. (P.) new species no 2;

48, C. (P.) new species no. 1: 49, C. (Taridius) opacula. Fig. 50: C. (Taridius) opacula, head, dorsal aspect. Figs. 51-54,

tarsal claws, terminal aspect: 51, C. {T.} opacula-, 52, Hystrichopus (sensu stricto) nr. dorsalis-, 53, C, (Afrotarus)

kilimana Kolbe; 54, H. (Plagiopyga) cymindoides (Peringuey). Scale bars = 200 )um. Legend, labium: el, epilobe of

mentum; 1, glossal (or ligular) sclerite; m, tooth of mentum; pg, paraglossa. Legend, tarsus: p, denticle of tarsal claw.

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Cymindine Lebiini of Authors

143

Figs. 55-61. SEM photographs of structures of Cymindina. — Left stylomeres of ovipositor. Fig. 55; Hystrichopus

(Pseudomasoreus) capicola Basilewsky, A and B, medial and ventral aspects, respectively. Fig. 56: H. (sensu stricto) nr.

dorsalis Thunberg, A and B, lateral and ventral aspects, respectively. Figs. 57, 58: H. (Plagiopyga) cymindoides

(Peringuey), lateral and ventral aspects, respectively. Figs. 59-61, A and B, medial and ventral aspects, respectively: 59,

Cymindis (Afrotarus) kilimana Kolbe; 60, C. (Taridius) opacula (Chaudoir); 61, C. (sensu stricto) hieronticus (Reiche).

Scale bars = 20 y.m. Legend: a, lateral ensiform seta; b, medial ensiform seta; c, sensory furrow; d, nematoid seta; S2,

stylomere 2.

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Ball and Hilchie

Figs. 62-65. SEM photographs of structures of Cymindina. — Ovipositor, left stylomere 2. Figs. 62-63, A and B, medial

and ventral aspects, respectively: 62 Cymindis (sensu stricto) suturalis Dejean; 63, C. (Pinacodera) new species no. 1. Figs.

64 and 65, microsculpture: 64, Hystrichopus (sensu stricto) nr. dorsalis Thunberg; 65, C. (P.) new species no. 1 . Scale bars

= 20 um.

Cymindine Lebiini of Authors

145

Recognition. — Diagnostic features of this genus are the following: head with two pairs of

supraorbital punctures; frons with (most members) or without longitudinal ridges and grooves

laterally; elytron with lateral umbilicate punctures larger than, and thus clearly distinguishable

from, setigerous punctures of discal intervals; right mandible without premolar tooth; median

lobe of male genitalia anopic, internal sac with or without sclerites; stylomere 2 of ovipositor

broad at base, with two ensiform setae on dorsal margins.

Classification. — Although the four groups ranked as subgenera are not easy to diagnose for

recognition of individual specimens, we are satisfied that each is monophyletic, and is

reasonably distinctive in combinations of structural features, and in patterns of geographical

distribution. The sequence of subgenera in the text is based on our concept of sister-group

relations, as discussed in more detail, below.

The characteristic form of stylomere 2 of the ovipositor (relatively straight, broad at least

preapically in ventral aspect), with ensiform setae relatively close to the apex (Figs. 59-62)

seems to be apotypic, and is the only such feature for delimiting Cymindis (sensu lato) in

relation to Hystrichopus {sensu lato). It is sufficient, however, to suggest that Cymindis is

monophyletic.

Features interpreted as synapotypic for Cymindis (sensu lato) and Hystrichopus (sensu

lato) and that thus support inference of a sister group relationship for these two taxa are the

ridged mental tooth (Figs. 46-49), and pectinate tarsal claws (Figs. 51-54; denticles reduced in

some members of subgenus Plagiopyga, and interpreted as lost from other members).

Subgenus Taridius Chaudoir, new status

Figs. 42, 49-51, 60, 69, 73, and 75

Diagnostic description. — Size small (SBL ca. 5. 5-7. 4 mm.) to average (ca. 10 mm.). Color: uniformly

rufo-piceous to piceous, or elytra bicolored piceous and flavous (Figs. 75 A, B) appendages paler (rufous or flavous) than

dorsum. Microsculpture: head- vertex and frons with meshes isodiametric, lines clearly developed; pronotum with meshes

transverse; elytra- dorsal surface with meshes isodiametric. Vestiture: dorsum glabrous, except for standard setae, dorsal

surfaces of tarsomeres each with single pair of setae near apex, or very sparsely setose (i.e., one or two additional short

setae). Head: frons each side with from two to seven (Fig 50) sharply defined ridges. Antennae: scape and antennomeres 2

and 3 with ring of setae near apex, otherwise glabrous; antennomeres 4-11 generally setose; antennomeres 3 and 4

subequal in length. Mandibles as in Figs. 42A - D. Labium as in Fig. 49. Metepisternum with lateral margin clearly longer

than anterior margin. Wings fully developed, oblongum and wedge cells as in Figs. 73A and B. Tarsal claws as in Fig. 51.

Median lobe of male genitalia (Figs. 69A, B) with apex simple; internal sac without sclerotized plate. Stylomere 2 of

ovipositor average for cymindoids, though more slender apically than in Pinacodera (Fig. 60A; cf Fig. 63A); microspines

on ventral surface.

Relationships of subgenus. — Compared to other subgenera of Cymindis, Taridius seems

more primitive, and thus likely to be the sister group of Pinacodera - Afrotarus - Cymindis.

Adults share with those of Pinacodera and some adults of Cymindis similar body form and

fully developed wings-but these features are correlated functionally and symplesiotypic.

Females of Taridius and Pinacodera share microspines on stylomere 2 (Figs. 60B and 65).

However, females of Ceylonitarus and Hystrichopus (sensu stricto) also exhibit this feature

(Fig. 64), and so we are disinclined to weight it heavily, for the similarity might be

symplesiotypic or homoplasious. In fact, Taridius seems to be without autapotypic character

states, and thus may be paraphyletic.

The distribution pattern is consistent with a relict status for Taridius: populations of this

structurally plesiotypic group occupy montane areas which are marginal relative to lowland

tropical forests. This suggests displacement from the surrounding lowland tropics. However, it

is also possible that the species are persisting in those areas where forest is still able to persist;

Quaest. Ent., 1983, 19 (1,2)

146

Ball and Hilchie

thus, the distribution pattern is the result of recent ecological circumstances, rather than

temporally remote events.

Included species. — Three species are recognized: C. opacula (Chaudoir, 1875); C.

birmanica (Bates, 1892); and C. stevensi (Andrewes, 1923). The first two species are each

monobasic. C. stevensi includes three subspecies: C. 5. stevensi; C. s. nilgirica (Andrewes,

1935); and C. 5. andrewesi (van Emden, 1937). Adults of these species and subspecies are

distinguished from one another in the following key. Andrewes (1935: 204-205) also included in

Taridius, C. nigra, but adults of this taxon have features of Afrotarus Jeannel, a group with

which Andrewes was unfamiliar when he published his treatment. At that time, Afrotarus had

not been erected, only two of its species had been described (in Cymindis), and they were

known only from the high mountains of East Africa and Abyssinia. Thus, Andrewes could not

be expected to be aware of the true affinities of his new species from the highlands of South

India.

Notes about habitat. — Andrewes (1935: 205) recorded that type material of C. s. nilgirica

was collected among dead leaves at 6000 feet above sea level, in the Nilgiri Hills.

Geographical distribution. — The species of this subgenus are known from the Palni and

Nilgiri Hills of South India, and from the hills of northeastern India, western Burma, and

eastern Java. Andrewes (1930: 343) lists localities.

Relationships of the species.- These are not clear. C. opacula and C. birmanica are

synapotypic in sculpture of the dorsum of the head, whereas C. birmanica and C. stevensi are

synapotypic in color pattern. One of these discordant pairs of synapotypies is homoplasious, but

we are not in position to infer which.

Key to Species of Subgenus Taridius Chaudoir

1 (0 ) Pronotum with more than two pairs of lateral setae; dorsum concolorous,

rufo-piceous C. (Taridius) opacula (Chaudoir), p. \47

r Pronotum with two pairs of lateral setae; dorsum bicolored: each elytron

predominantly flavous, with piceous or black longitudinal marks suturally.

marginally, and transversely in posterior 0.33 (Figs. 75 A, B) 2.

2 (F) Frons each side with at least five longitudinal carinae (cf. Fig. 50)

C. (Taridius) birmanica (Bates), p. 147

2' Frons with not more than three longitudinal carinae each side 3.

3 (2") Pronotum narrow (Hw/Pmw 0.82-0.86), flavous color extended from margins to

larger areas of disc, or confined to reflexed margins; elytron with transverse

dark mark interrupted or very narrow (Fig. 75 A)

C. (Taridius) s. andrewesi (van Emden).

y Pronotum broad (Hw/Pmw 0.73-0.80), with only reflexed margins flavous,

otherwise piceous; elytron with transverse dark mark various 4.

4 (3') Elytron with transverse dark mark continuous, broad medially (Fig. 75B)

C. (Taridius) s. nilgirica Andrewes).

4' Elytron with transverse dark mark interrupted or very narrow medially (Fig.

75A) C. (Taridius) s. stevensi (sensu stricto), p. 147

Cymindine Lebiini of Authors

147

Cymindis (Taridius) opacula (Chaudoir) NEW COMBINATION

Figs. 42A-D, 50-51, and 60A, B

Taridius opaculus Chaudoir, 1875: 8. TYPE AREA.- “le nord de I’Hindostan”.- Bates, 1892: 152.- lakobson, 1907:

396.-Csiki, 1932: 1489.- Andrewes, 1935: 204.- Jedlicka, 1963: 462. Not seen.

Geographical distribution. — This species is known from northern India and northern

Burma, within the range of, but generally at lower elevations than, C. stevensi.

Material examined. — Two males, four females, Assam Lohara Kaziranga 110 m. X.7- 16.61 E. S. Ross, D. Q.

Cavagnaro (CAS). Female, Calcutta (IRSB).

Cymindis (Taridius) birmanica (Bates) NEW COMBINATION

Taridius birmanicus Bates, 1892: 152. TYPE LOCALITY: Teinzo, Karin Cheba, 1300-1400 m.- Andrewes, 1930:

343.-Csiki, 1932: 1489.- Andrewes, 1935: 204.- Jedlicka, 1963: 462.

Geographical distribution.— This species is known only from the Karen Hills of western

Burma. We have not seen any specimens.

Notes. — To judge from the original description, the type specimens of this species have the

flavous marks of the elytra smaller than is characteristic of specimens of C. stevensi.

Cymindis (Taridius) stevensi (Andrewes), NEW COMBINATION

Figs. 75A, B

Taridius stevensi Andrewes, 1923: 689. CO-TYPE labelled: Cotype [circular label, ringed with green]; Gopaldhara,

Darjeeling H. Stevens 1919; H. E. Andrewes coll. BM 1945-97; Taridius Stevensi Andr. cotype H. E. Andrewes det.

[handwritten]. (BMNH). TYPE LOCALITY: Sikkim, Golpadhara (near Darjeeling).- Andrewes, 1930: 343.- Csiki,

1932: 1489.- Andrewes, 1935: 204.

Taridius nilgiricus Andrewes, 1935: 204. LECTOTYPE (here selected) female, labelled: Cotype [circular label,

ringed with green]; Nilgiri Hills H. E. Andrewes; Taridius nilgiricus co-type Andr. H. E. Andrewes det. [handwritten]

(BMNH). TYPE LOCALITY: Nilgiri Hills, India. NEW SYNONYMY.

Taridius andrewesi van Emden, 1937: 123. TYPE LOCALITY: Java, Tengger, Nonkodjadjan. NEW SYNONYMY.

Notes about synonymy. — The specimens included by us in C. stevensi (sensu lato) represent

three nominal species. They are combined on the basis that differences are slight and diagnostic

characters sufficiently variable to suggest at most a pattern of step-clines. One of the principal

diagnostic features claimed by Andrewes for C. nilgirica is extra setae on elytral intervals 3 and

5. However, the elytra of the lectotype are asymmetric in number of setae, and the other

speeimens from the Nilgiri Hills have the normal number of two setigerous punctures in

interval 3. Thus, the lectotype is interpreted as simply an abnormal specimen.

Van Emden (1937: 123) provided a detailed description of C. andrewesi, but the only

differences that seem of diagnostic value are given above, in the key.

Pattern of variation. — Two discordant dines are suggested: decrease in amount of black

pigment of the elytra from southern to northern India; and decreased width of the pronotum

from- northeastern India to western Burma (expressed as increase in value for Hw/Pmw- Table

3). In color pattern, adults of C. s. stevensi are more like those of C. 5. andrewesi, whereas in

proportions of the pronotum, C. 5. stevensi is more like C. 5. nilgirica.

Geographical distribution. — Cymindis s. nilgirica is known only from the Western Ghats

of southern India (Nilgiri and Palni Hills). C. 5. stevensi ranges along the lower southern slopes

of the Himalaya from Sikkim to Haldwani in the United Provinces, and in the Kondmal Hills

of the Eastern Ghats (Andrewes, 1923: 690). Before central India was cleared in historic times

of its forests (Dilger, 1952: 125-127), it seems possible that the ranges of C. 5. stevensi and C. 5.

nilgirica were in contact.

Quaest. Ent., 1983, 19 (1,2)

148

Ball and Hilchie

TABLE 3

DATA ABOUT VARIATION IN STANDARDIZED BODY LENGTH (MM) AND IN

VALUES FOR THE RATIO Hw/Pmw AMONG SAMPLES OF CYMINDIS STEVENSI

(ANDREWES)

SUPSPECIES AND N SBL MEAN Hw/Pmw MEAN

LOCALITY RANGE RANGE

C s. nilgirica

Cymindis s. andrewesi occupies areas to the east of the Irawaddy River, with samples known

from as far north and west as the Karen Hills of Burma and as far south and east as the

Indo-Australian island of Java.

Chorological affinities. — The range of C s. stevensi is overlapped by that of C. opacula,

though the two species have not been recorded from the same locality. The range of C. s.

andrewesi overlaps that of C. birmanica in the Karen Hills. Locality data are not sufficiently

precise to indicate if the two taxa are microsympatric, or if their life cycles are synchronic.

Nonetheless, these geographical contacts are reasonable evidence for reproductive isolation

between C. stevensi and the other two species. Specific distinctness of C. opacula and C.

birmanica is not tested by chorological data.

Material examined. — In addition to type material recorded above, we have seen nine

specimens, as follows.

C. s. nilgirica. Three females: Nilgiri Hills, H. E. Andrewes (BMNH); same locality, collected by G. F. Hampson

(BMNH); and Palni Hills, Kodaikanal, 6900-7200 ft., IZ. 22, S. Kemp (ZSIC).

C. s. stevensi. Male and three females (including one paratype) Gopaldhara, Darjeeling, Sikkim, H. Stevens

(BMNH).

C. s. andrewesi. Male, female, paratypes, O. Java Tengger Nonkodjadjan 1300 m. Wegner (BMNH). Female, Burma

Karen Hills; Taridius sp. H. E. Andrewes det. (BMNH).

Cymindine Lebiini of Authors

149

Subgenus Pinacodera Schaum, new status

Figs. 43, 46-48, 63, 65, and 76

Diagnostic description. — Body moderately flattened, generally elongate. Size about average for carabids, SBL

ca. 5.50-10.50 mm. Color: somber, with dorsum of most specimens darker than venter, and appendages paler than body

integument; head rufous, piceous or black; pronotum rufous, piceous, or black, lateral areas paler or not, than disc; elytra

with dorsal surface rufo-flavous, rufous, piceous, or black, or various combinations of these, epipleura of most specimens

paler than dorsal surface; venter rufous, piceous, or black; antennae rufo-flavous, or rufo-piceous, with scape of most

specimens paler than remaining antennomeres; legs rufo-flavous to black, with femora of most specimens paler than other

articles. Microsculpture: meshes in general, isodiametric, or transverse, but comparatively wide; head- frons and clypeus

with meshes isodiametric, microlines distinct or indistinct, or meshes partially or wholly effaced, ventral surface with

meshes transverse, microlines clear, or partially or wholly effaced; pronotum with meshes uniformly transverse, or

isodiametric postero-laterally, lines clear or partially or wholly effaced; scutellum and elytra with meshes uniformly

isodiametric; prosternum and pterothorax ventrally with meshes transverse, microlines clear, or partially or wholly effaced,

proepisternum with meshes oblique, microlines partially or wholly effaced; abdominal sterna with meshes transverse,

microlines clear or partially or wholly effaced. Vestiture: glabrous (except fixed setae), or sparsely to densely setose and

punctate; elytral intervals impunctate, or with one or more irregular rows of setigerous punctures. Head: frons each side

with two to five irregular longitudinal ridges and grooves more or less distinctly developed. Antennomeres 1-3 either

glabrous (except normal preapical setae) or sparsely setose, setae short; antennomeres 3 and 4 subequal in length.

Mandibles as in Figs. 43A-D. Lacinia as in Fig. 45. Labium as in Figs. 46-48. Metepisternum either distinctly longer than

wide, or width at base and length of lateral margin subequal. Wings fully developed, or reduced to short stubs. Median

lobe of male genitalia with apical portion in lateral aspect straight and narrow, or expanded slightly into knob, internal sac

with sclerotized plate, as in subgenus Cymindis (cf. Fig. 72C), or without armature. Stylomere 2 of ovipositor average for

Cymindis (Figs. 63A, B; cf. Figs. 62A, B); microspines on ventral surface (Fig. 65).

Included species. — Twenty seven species are known of which nine are described. The group

is presently under study by us.

Way of life. — Data about life histories are available for two species, C. platicollis Say, and

C. limbata Dejean (Mahar, 1978). Adults of both species are crepuscular or nocturnal, living

on ground in leaf litter, and in trees. Larvae are terrestrial. C. platicollis is a spring breeder,

while C. limbata breeds during summer. Site of oviposition has not been determined.

Notes about habitat. — Taxa are associated with arboresecent vegetation, from desert,

tropical thorn scrub and thorn forest, to dry oak forests, wet conifer forests and cloud forests in

the mountains of northern Middle America. Altitudinal range of the subgenus extends from sea

level to 2900 meters above sea level.

Geographical distribution. — The range of this subgenus extends from Honduras in Middle

America, to southern Nova Scotia and Ontario, in eastern North America.

Classification. — The species are arrayed in two groups: a more northern group, males of

which have armature in the internal sac, and a more southern group whose males have

unarmored internal sacs.

Phylogenetic relationships. — We believe that Pinacodera is the sister group of the ancestor

of Afrotarus + Cymindis (sensu stricto), based on the inference that males of the common

ancestor of subgenera Pinacodera + Afrotarus + Cymindis exhibited the autapotypic feature

of the complex apical sclerite in the internal sac, as preserved in the descendant stocks. This

requires loss of this sclerite from at least one descendant lineage of Pinacodera

A second hypothesis, based on the pattern of geographical distribution of Cymindis sensu

lato {Pinacodera confined to the New World, two of the other three subgenera confined to the

Old World, and one with most of its species there), is that Pinacodera is the sister group of the

ancestral stoek of the other three subgenera. Such an hypothesis requires either independent

gains of the genital sclerite (once in Pinacodera and once in the Afrotarus + Cymindis stock),

or at least two losses of this sclerite (once from Taridius, and once from Pinacodera). Neither

hypothesis is very well supported, but we think the first is rather stronger, requiring one less

Quaest. Ent., 1983, 19 (1,2)

150

Ball and Hilchie

Cymindine Lebiini of Authors

151

Figs. 66-72. Line drawings of structure of Cymindina. — Pronota and genitalia. Figs. 66-68. Pronota, setae omitted, of

Cymindis (Afrotarus) nigra (Andrewes). Figs. 69-72. Male genitalia of Cymindis. Fig. 69: C. (Taridius) opacula

(Chaudoir), A and B, median lobe, left lateral and ventral aspects (various portions), respectively. Fig. 70: C. {Afrotarus}

kilimana Kolbe, A and B, median lobe, left lateral and ventral aspects, respectively; C and D, left and right parameres,

ventral aspect, respectively. Fig. 1\\C. (A), nigra (Andrewes), A and B, median lobe, left lateral and ventral aspects,

respectively. Figs. 72A-C: Cymindis (sensu stricto) suturalis Dejean, A and B, median lobe, left lateral and ventral

aspects, respectively; C. internal sac, inverted position. Figs. 72D-E: C. (sensu stricto) hieronticus (Reiche), median lobe,

left lateral and ventral aspects, respectively.

Quaest. Ent., 1983, 19 (1,2)

Cymindine Lebiini of Authors

153

Figs. 73-76. Line drawings of structures of Cymindina. — Figs. 73-74. Wing veins and cells of Cymindis species, A and B,

oblongum and wedge cells, respectively: 73, C (Taridius) opacula (Chaudoir); 74, C. {sensu stricto) suturalis Dejean. Fig.

75. Color pattern of left elytron of Cymindis (Taridius) stevensi (Andrewes); A, C. 5. andrewesi (van Emden); B, C. s.

nilgirica (Andrewes). Fig. 76. Terminal abdominal sclerites of C. (Pinacodera) new species no. 1: A, Tergum VIII, dorsal

aspect; B, Sternum VIII, ventral aspect; Tergum X, dorsal aspect. Legend. Wing cells: O, oblongum; W, wedge. Veins: A,

anals; Cu, cubitus; M. media; R, radius.

Quaest. Ent., 1983, 19 (1,2)

154

Ball and Hilchie

loss of a complex structure, and not requiring its independent development in two lineages.

Subgenus Afrotarus Jeannel, new status

Figs. 53, 59, 66-68, and 70-71

Diagnostic description. — Size small SBL ca. 5.0-7. 0 mm. Color: body rufo-piceous to almost black,

appendages somewhat paler. Microsculpture: head- vertex with meshes isodiametric, frons with meshes transverse or

absent; pronotum with meshes transverse; elytra with meshes transverse or isodiametric; ventral surface with meshes

generally transverse. Luster: head shining; pronotum and elytra shining to slightly iridescent; ventral surface generally

slightly iridescent. Vestiture: dorsal surface of body glabrous (adults of most species) or sparsely setose (adults of one

species), and dorsal surfaces of tarsomeres glabrous or setose. Head: frons each side with two or three sharply defined

ridges extended longitudinally. Antennae various: antennomeres 4-10 each either average in proportions (1/w ca. 2.00) or

shortened (1/w ca. 1.50). Pronotum (Figs. 66-68) transverse or only slightly wider than long, sides sinuate, lateral margins

slightly elevated posteriorly. Metathorax reduced, metepisternum with lateral and basal margins subequal in length, or

lateral margin distinctly longer than basal margin. Wings reduced to short stubs. Median lobe of male genitalia with apex

hooked or straight; internal sac with sclerotized plate, as in subgenus Cymindis (Figs. 70A, 71 A). Stylomere 2 of ovipositor

with dorsal ensiform setae longer (Figs. 59A, B) or average.

Included species. — This subgenus includes five species: four African; one Arabian; and one

Indian. The African species are: C. kilimana Kolbe; C. leleupi (Basilewsky); C. meruana

(Basilewsky); and C. raff ray i Fairmaire. The Arabian species is C. scotti Basilewsky, and the

Indian species is C. nigra (Andrewes).

Notes about habitat. — The species are known from mountain forests and their environs.

Geographical distribution. — The range of this subgenus includes the high mountains of

east Africa (Kilimanjaro and Meru, in Tanzania), Ethiopia, the southern mountains in the

Arabian Peninsula, and the hills in southern India. Two species (C. kilimana and C. leleupi)

are known from a single massif (Kilimanjaro). The remaining species are allopatric relative to

one another. However, all of these taxa are known from very few specimens.

Phylogenetic considerations. — Basilewsky (1962: 205-207) suggested that the extant

species of Afrotarus (known to him only from Africa and Arabia) represented a Palaearctic

lineage derived from Cymindis stock. Presence of the group in southern India suggests that it

probably occupied lands farther north, an inference that could also be derived from

Basilewsky’s hypothesis. However, the evidence at hand does not require that Afrotarus be a

southern derivative of a northern group. We think it likely that Afrotarus is relict, and is the

sister group of subgenus Cymindis.

Key to Species of Subgenus Afrotarus Jeannel

1 (0 ) Dorsal surface and eyes sparsely setose; median lobe of male genitalia with

apical hook (Fig. 71)

C. (Afrotarus) nigra (Andrewes), p. 155

V Dorsal surface and eyes glabrous; apex of median lobe hooked or not 2.

2 (!') Pronotum about as long as wide, impunctate, marginal grooves each broad

throughout length, lateral margins elevated. Elytra subovoid, disc markedly

flattened, marginal grooves deep, lateral margins elevated, microlines of

intervals distinct. Head elongate, smooth, eyes small, hardly prominent, temples

long. Antenna long, slender, three articles extended past base of pronotum.

Labial tooth broad, rounded apically. Head and pronotum rufo-testaceous,

elytra castaneous medially, testaceous marginally, and with large humeral mark

testaceous. Length 8.00 mm. or more

Cymindine Lebiini of Authors

155

TABLE 4

DATA ABOUT VARIATION IN STANDARDIZED BODY LENGTH (SBL) AND IN

VALUES FOR THE RATIO Pl/PwB AMONG SPECIMENS OF CYMINDIS NIGRA

(ANDREWES)

LOCALITY AND SEX SBL (mm.) Pl/PwB

Palni Hills 9 5.46 0.94

Cardomon Hills (5 5.02 1.03

Maharashtra 6 6.54 1.02

C. (Afrotarus) scotti Basilewsky.

2' Pronotum transverse, wider than long, disc punctate or not, lateral margins not

elevated. Head broad, eyes large and prominent, temples shorter. Antennomeres

4-10 short and wide, extended past base of pronotum only slightly more than

two articles. Mentum with apex of tooth acute. Elytron with lateral margin

plane, not elevated. Length 7.00 mm. or less 3.

3 (2') Elytral intervals with microlines indistinct, meshes transverse, surface slightly

iridescent. Pronotum 1.40 times wider than long, sides markedly constricted

posteriorly, but not sinuate; posterior angles slightly projected. Elytra

castaneous, with vague testaceous humeral markings in basal 0.25 of intervals 5

and 6 C. (Afrotarus) raffrayi Fairmaire.

3' Elytral intervals with microlines less well developed, meshes isodiametric, dull 4.

4 (3') Elytra generally castaneous, each with testaceous mark on interval 3 anterior to

middle, and another short one preapically . C. (Afrotarus) leleupi (Basilewsky).

A' Elytron with interneurs shallow, impunctate, uniformly piceous or bicolored

with well developed humeral marks. Median lobe without apical hook

C. (Afrotarus) kilimana Kolbe.

Cymindis (Afrotarus) nigra (Andrewes), NEW COMBINATION

Figs. 66-68, and 71

We record a few observations about the limited material that we have seen of this species:

three specimens from the proximally located Palni (type locality) and Cardomon Hills, and one

specimen from Maharashtra, some 500 km. to the north. The male from Maharashtra differs

rather strikingly from the more southern specimens: dorsal integument and eyes of the former

are more evidently setose, lateral margins of the pronotum (Fig. 68 cf. Figs. 66 and 67) are

more reflexed, elytral humeri are less constricted, the metepisternum is distinctly longer than

wide, and body size is larger (Table 4). These differences suggest specific distinctness. On the

other hand, the adult of C. nigra from the Cardomon Hills differs from specimens in the Palni

Hills in form and proportions of pronotum (pronotum longer than wide. Table 4), and in color,

though the localities are close together. This suggests that C. nigra is inherently variable, so

Quaest. Ent., 1983, 19 (1,2)

156

Ball and Hilchie

that one could imagine that populations that are far apart geographically could differ

strikingly. Further, the apex of the median lobe of the northern male is identical in form to that

of a male paratype. For now, we prefer to include all of these specimens in a single species, with

the expectation that additional material will eventually be accumulated and will serve as the

basis for a revision of this interesting complex.

One might expect that the Indian species of Afrotarus would exhibit a combination of

features that would set it apart from the more western species which are geographically close to

one another. In fact, this expectation is not realized, and the Indian species seems to differ no

more from the African species than the latter differ from one another.

Material examined. — We have studied specimens of subgenus Afrotarus, labelled as

follows.

C. kilimana Kolbe. Male and female, TANZANIA, Kilimanj. Sjostedt; Kiboscho 3000 m Mus Paris coll. Ch. Alluaud

(MNHP).

C. nigra (Andrewes). Male, INDIA Madero Ind. Or.; Staudinger and Bang-Haas, 1933; H. E. Andrewes Coll. BM

1945-97 (BMNH). Female, Shores of Kodaikanal Lake, 6550 ft Palni Hills S. India VII. 22 (under stones), S. Kemp

(ZSIC). Female, S. INDIA 8 mi. NE Munnar, 6200' III. 20. 62; E. S. Ross, D. Q. Cavagnaro (CAS). Note: this locality is

in the Cardomon Hills of the Western Ghats, and is also known as Pallivasal. Male, INDIA, Maharashtra Mahableshwar

1250 m. 11.13.62; E. S. Ross, D. Q. Cavagnaro (CAS).

C. raffrayi Fairmaire. Male, ETHIOPIA Simien Derasghie c. 9800 ft. 22.XII.1952; from grove of tall juniper trees

north of town; N. Ethiopia 1952-1953, Hugh Scott, 3 m, 1953-335; Afrotarus raffrayi Fairmaire, det. Basilewsky 1953

(BMNH).

Subgenus Cymindis Latreille, NEW STATUS

Figs. 41A-D, 61A-B, 62A-B, 72A-B, 74A-B

Major recent faunal treatments of Cymindis are for: France, by Jeannel (1942a); Morocco,

by Antoine (1962); eastern Asia, by Jedlicka (1963); Japan, by Habu (1967); and the Nearctic

Region, by Lindroth (1969a). V. M. Emetz is actively studying the eastern Palaearctic species.

Perhaps in the near future, he will be able to write a general revision of the subgenus on a

world-wide basis, and thus establish a general system.

Diagnostic description. — The faunal treatments listed above offer detailed

characterizations of adults. Especially useful are the descriptions by Antoine and Lindroth.

Here, we record only those features that serve to differentiate Cymindis adults from those of

other subgenera, and that offer promise for establishing phylogenetic relationships.

Size average for carabids (length 7.0 - 12.0 mm.). Color various, dorsum varied from nearly black to rufous, or

bicolored, with head and prothorax rufous, elytra darker (black or metallic blue); venter generally paler than dorsum; legs

and mouthparts piceous to testaceous, generally paler than dorsum; elytra concolorous, or dark with pale humeral marks.

Microsculpture developed or not, but if developed: meshes isodiametric on head and elytra, slightly transverse on

pronotum. Luster: dorsal surface generally dull, pronotum more shiny than head or elytra, metallic in members of few

species (group designated Menas Motschulsky). Vestiture: surface generally setose, though markedly reduced in members

of some species (for example, C. suturalis Dejean, and C. hieronticus Reiche) to very sparse and very short setae, visible

only in lateral aspect at high magnification; tarsomeres dorsally and antennomeres 1-3 setose. Surface generally punctate,

though punctures reduced in specimens with reduced setation. Fixed setae average, though members of some species with

additional lateral setae on pronotum; elytron with three or four setigerous punctures in interval 3. Head: frons each side

with or without longitudinally extended ridges and grooves, frontal impressions indistinct. Antennae average for carabids,

antennomere 3 distinctly longer than 4; antennomeres 4-10 each longer than wide. Mouthparts: mandibles as in Figs.

41A-D, right mandible with well developed anterior terebral tooth (Fig. 41 A); terminal palpomeres similar in form, or

labial of males of some species broader than maxillary; labial palpomeres of females unmodified. Pronotum slightly or

markedly transverse, sides slightly sinuate or incurved evenly posteriorly; disc slightly convex; posterior angles rectangular

or more or less rounded; base arcuate; metepisternum with lateral margin longer than anterior margin {i.e., longer than

wide). Elytra average for lebiines. Macropterous or brachypterous. Tarsal claws with denticles well developed, three or

more (adults of most species), or very small, one or two. Median lobe of male genitalia with apical portion short, either

straight or curved, apical orifice on left side (Figs. 72A-E); internal sac with fields of microtrichia, with apical sclerite

(Fig. 72C). Stylomere 2 of ovipositor average for Cymindis (Figs. 62A-B).

Cymindine Lebiini of Authors

157

Included species. — This subgenus is moderately diverse, including more than 150 species,

but it is not very divergent (Antoine, 1962: 566). Thus it seems to be, in terms of its state of

evolution, in full flower.

As Chaudoir (1873: 53) noted, Cymindis became a dumping ground (“une espece de

magazin”), including a collection of unrelated lebiines that shared in common terminal

securiform palpomeres. He traced development of the concept of the genus that he had, which

is essentially the one we use for the nominotypical subgenus. Csiki (1932: 1464) listed

Iscariotes Reiche and Psammoxenus Chaudoir as subgenera of Cymindis (Chaudoir treated

these groups as genera close to Cymindis). The subgenus Tarulus Bedel, 1906 was accepted as

such by Csiki (1932: 1464). Jeannel (1942a) excluded C. canigoulensis (placed in the subgenus

Pseudomasoreus by Desbrochers des Loges (1904), and similarly ranked by Csiki (1932:

1464)). Additionally, subgenera have been added to those listed by Csiki (1932: 1464-1465):

Pseudocymindis Habu, 1967; Paracymindis Jedlidka, 1968; Assadera Mandl, 1973;

Pteroritziella Mandl, 1973; Neopsammoxenus Emetz, 1973; and Pseudomastes Emetz, 1972.

We believe that all of these groups should be subordinate to the rank of subgenus, but that

remains to be determined.

Notes about habitat. — According to Lindroth (1969a: 1071), and from our experience with

the Nearctic fauna, the species are xerophilous, individuals living in open country with sparse

vegetation. In forested regions, adults are found in open meadows, or along edges of forests.

Populations of some species (for example, C. arizonensis Horn) inhabit deserts, while those of

other species (for example, C. borealis LeConte) occur on dry arctic tundra.

Geographical distribution. — This subgenus is widespread in the Holarctic Region. In the

Old World, it ranges from the Atlas Mountains in the west and Himalaya in the east northward

to the Arctic. The range in the New World is similar: from northern Chihuahua, Mexico, to the

Arctic Islands of Canada. Most species, however, are concentrated in the middle latitudes of

the Northern Hemisphere.

Phylogeny and zoogeography. — We have three points to make. Judging from the numerous

species identified by means of few characters, and wide continuous range of the subgenus, we

believe its differentiation to be relatively recent. Because the group is much more diverse and

divergent in the Old World than the New (all Nearctic species probably belong to a single

Holarctic species group), and because its seemingly closest relatives are also in the Old World,

we believe that Cymindis {sensu stricto) is of Old World origin. We also believe that the

xerophily of the species is a derived feature that enabled the group to spread and diversify with

the later Tertiary development of dry temperate habitats.

Material examined. — We have seen specimens of most Nearctic species, but we have

studied and dissected only the following Old World forms.

Cymindis suturalis Dejean. Male and female, labelled: ALGERIA Biskra, Van Dyke Coll. (CAS).

Cymindis hieronticus Reiche. Four males labelled: W. PAKISTAN 10 mi. SW Kohat 650 m. XII. 19. 1961; E. S. Ross

and D. Q. Cavagnaro. Four females, labelled: W. PAKISTAN, 2 mi. w. Cherat, 1200 m., XII.20.1961; E. S. Ross and D.

Q. Cavagnaro.

Hystrichopus Boheman

Figs. 44A-D, 52, 54, 58, 64, 77-88C

Hystrichopus Boheman, 1848: 42. GENERITYPE: Hystrichopus angusticollis Boheman, 1848 (subsequent

designation by Basilewsky, 1954b: 15).- Peringuey, 1896: 212-218.- Basilewsky, 1954b: 7-80.- 1958: 297-302.- 1960:

85-86.- 1961b: 122-126.- 1962: 207-212.- 1976: 717.

Plagiopyga Boheman, 1848: 75. GENERITYPE: Plagiopyga ferruginea Boheman, 1848: 76 (monotypy). NEW

SYNONYMY.- Peringuey, 1896: 212-221.- Basilewsky, 1945b: 80-94.- 1958: 302.

Quaest. Ent., 1983, 19 (1,2)

158

Ball and Hilchie

Pseudomasoreus Desbrochers des Loges, 1904: 140, 143, 163. GENERITYPE: Cymindis canigoulensis Fairmaire

and Laboulbene, 1854: 32 (monotypy).- Bedel, 1906: 241.- Porta, 1923: 227.- Antoine, 1938: 171.- Jeannel, 1941: 62.-

1942a: 1039-1041.- 1949: 878-881.- Basilewsky, 1953b: 57-58.- 1954c: 89-96.- 1958a: 296-297.- 1962: 212-216.-

Antoine, 1962: 562.- Mateu, 1970a: 173-175.- Basilewsky, 1976: 717.- Mateu, 1980: 16-23.

Assadecma Basilewsky, 1982: 20. GENERITYPE: Assadecma madagascariensis Basilewsky, 1982: 22 (original

designation and monotypy). NEW SYNONYMY.

Data about synonymy are provided in conjunction with treatments of subgenera.

The following features of adults characterize this genus: head with two pairs of supraorbital

setigerous punctures; frons without ridges and grooves laterally; elytron with lateral umbilicate

punctures larger and thus clearly distinguishable from other setigerous punctures; right

mandible with well developed premolar (Figs. 44B and D); median lobe of male catopic,

internal sac without sclerites (Figs. 86C-88C); valvifer of ovipositor with extra lobe, stylomere

2 elongate, slender, with single ensiform seta (Figs. 55-57).

Jeannel (1942a) established the tribe Pseudomasoreini for Pseudomasoreus Desbrochers

des Loges, adding to it (1949) Hystrichopus Boheman. Subsequently, Basilewsky (1954b), who

treated this group as a tribe of Lebiinae equivalent to the Cymindini and other lebiine tribes,

added Plagiopyga Boheman. Three years later, Basilewsky (1957: 240) noted that male

dromiines of Klepturus and Klepteromimus also had catopic median lobes, and concluded that

it would be necessary to re-evaluate this character state as a basis for ranking the

pseudomasoreine group as a tribe of Lebiinae. With Mateu (1980: 17) we believe that this

feature does not constitute a sufficient basis for giving pseudomasoreines a high rank, though in

contrast to him, we believe that catopy was evolved only once in the Cymindina, and thus

delimits a monophyletic group.

Subgenus Pseudomasoreus Desbrochers des Loges, 1904, NEW STATUS

Characteristics. — This subgenus is adequately characterized by Jeannel (1942a and 1949),

Basilewsky (1954a), and Antoine (1962). To the features noted by these authors, we add:

stylomere 2 of ovipositor slender throughout length, more or less tubular, with dorso-lateral

ensiform setae moderate to long (Fig. 55A).

Included species. — This subgenus includes 13 previously described species, and four more

from localities in the Union of South Africa, are described below.

Geographical distribution. — The range of Pseudomasoreus extends from the Cape Region

of the Union of South Africa to the Pyrenees Mountains of Spain and France, and includes

Madagascar. However, the range is discontinuous: five species are known only from South

Africa (see below); nine species are known only from Madagascar (Mateu, 1980); two species

are known only from the high mountains of East Africa (Basilewsky, 1962); and P.

canigoulensis is known only from localities in and to the north of the Atlas Mountains.

Notes about phylogeny and zoogeography. — Mateu (1980: 15), in conjunction with his

useful revision of the Madagascan species, notes that they seem to be montane- adapted and

possibly xerophilous, but that little is known about their way of life. Most species (including

those in Africa) are known from single localities, only. Mateu also noted the relative abundance

of species of Madagascar compared with those on the African continent, and suggested that

additional species may be discovered. He stated that the subgenus, though truly old, seems to

have been revitalized on Madagascar, for its species there are very similar to one another, and

thus seem to have evolved recently.

Jeannel (1942b: 316-317), when species were known only from Madagascar and north of the

Atlas Moutains, suggested that Pseudomasoreus had arisen in eastern Africa during the

Cymindine Lebiini of Authors

159

Cretaceous, had reached the two areas listed above, and survived there, but had become extinct

on mainland Africa, to the south of the Sahara. Subsequent discovery of H. capicola, H.

kivuanus, and H. uluguruanus confirmed Jeannel’s prediction that the group had been in

Afrotropical mainland Africa. Basilewsky (1954c: 91), when he described H. capicola,

predicted that the group would also be represented on the mountains of East Africa. Thus, his

more detailed prediction was confirmed with discovery of H. kilimanus hkb,f>and H.

uluguruanus.

Both Jeannel and Basilewsky stated that Pseudomasoreus had arisen in Cretaceous time,

and had once been continuously distributed. Thus, the present discontinuous range is

interpreted as relict, and these authors suggested that Pseudomasoreus is on the way to

extinction. This may be true, but it is interesting to note that the northern periphery of the

range of the group is occupied by a species whose adults are macropterous. It thus seems

possible that this lineage may be a recent arrival in the north. However, until phylogeny can be

reconstructed and used as a basis for interpreting the distribution pattern, it seems best to avoid

making additional inferences.

The Species of Pseudomasoreus of East and South Africa

We have had the opportunity to study 16 specimens of Pseudomasoreus from Afrotropical

localities. These represent material of three described species and four undescribed species. We

describe the latter and provide a key to these and to the previously described mainland

Afrotropical species.

Description. — We list here features shared by adults of all of the species.

Color. Generally somber, dorsal surface various, lateral margins of pronotum and elytra generally a bit paler than

medial areas; ventral surface constantly dark (rufo-piceous to piceous); appendages flavous.

Microsculpture. Dorsum of head (including clypeus) and labrum with meshes isodiametric; pronotum with meshes

various; meshes of pro- and pterothoracic sterna and metepisterna transverse; meshes of pro- and mesepisterna elongate;

meshes of abdominal sterna transverse. Luster of dorsal surface various; of ventral surface, iridescent.

Eyes. Moderate in size, flattened, not prominent.

Pronotum (Figs. 77-82). Subcordate to cordate, sides incurved posteriorly; posterior angles obtuse; disc with median

longitudinal impression and laterally with irregular shallow impressions, without transverse impressions; surface slightly

convex, laterally slightly sloped; lateral grooves well developed, posterior lateral impressions irregular shallow basins, more

or less continuous with lateral grooves. Two pairs of lateral setae.

Elytra flat; posterior margins subtruncate. Interneurs average, impunctate; scutellar interneur developed. Intervals

slightly convex, sparsely punctate. Parascutellar setigerous puncture developed. Disc with two setigerous punctures in

interval 3; umbilical series including about 14 setigerous punctures.

Relationships. — The data are not available to do more than arrange the species in order of

inereasing apotypy, based on inferred morphoelines of ehanges in mieroseulpture (from

isodiametric to transverse meshes) and structural changes associated with loss of wings

(reduction of metathorax, and change in form of elytral humeri). We also assume that these

speeies are more elosely related to one another than they are to the Madagasean speeies of

Pseudomasoreus or to the Mediterranean P. canigoulensis.

Key to Eastern and Southern African Species of the Subgenus Pseudomasoreus Desbrochers des

Loges

1 (0 ) Elytron with mieroseulpture meshes isodiametrie 2

V Elytron with mieroseulpture meshes transverse, surfaee irideseent or not 3.

2(1) Metepisternum elongate (w/l = ca. 1.50); size smaller (SBL ea. 5.00 mm) ....

H. {Pseudomasoreus} reticulatus new species, p. 164

1' Metepisternum short (w/1 ea. 1.00); size larger (SBL more than 6.00 mm) ....

Quaest. Ent., 1983, 19 (1,2)

160

Ball and Hilchie

Cymindine Lebiini of Authors

161

A

Figs. 77-84. Line drawings of structures of Cymindina. — Figs. 77-82. Pronota, dorsal aspect, of Hystrichopus

(Pseudomasoreus) species; 77, H. reticulatus, new species; 78, H. capicola (Basilewsky); 79, H. uluguruanus

(Basilewsky); 80, H. basilewskyi, new species; 81, //. thoracicus, new species; 82, H. mateui, new species. 83A, H.

kivuanus (Basilewsky); 83B, H. mateui. new species. Fig. 84. Wing cells and associated veins of Hystrichopus (sensu

stricto) massaicus Basilewsky: A. oblongum cell; B, wedge cell. Legend. Wing cells: O, oblongum; W, wedge. Veins: A,

anals; Cu, cubitus; M, median; R, radius.

Quaest. Ent., 1983, 19 (1,2)

162

Ball and Hilchie

88A

Cymindine Lebiini of Authors

163

Figs 85-88. Line drawings of structures of Cymindina. — Fig. 85. Wing cells and associated veins of Hystrichopus

(Plagiopygaj chaudoiri Peringuey: A, oblongum cell; B, wedge cell. Figs. 86-88. Male genitalia, median lobe of

Hystrichopus (Pseudomasoreus) species. A, B, C, left lateral, dorsal, and ventral aspects (basal lobe excluded),

respectively, of: 86, H. uluguruanus (Basilewsky); 87, H. basilewskyi, new species and 88, H. mateui, new species.

Legend. Wing cells: O, oblongum; W, wedge. Veins: A, anals; Cu, cubitus; M, media; R, radius.

Quaest. Ent., 1983, 19 (1,2)

164

Ball and Hilchie

H. (Pseudomasoreus) capicola Basilewsky, p. 167

3 (T) Metepisternum long and slender (w/1 more than 1.50); microsculpture meshes

of elytron wide, surface not iridescent; elytron with dark mark behind middle

(Fig. 83 A) H. (Pseudomasoreus) kivuanus Basilewsky, p. 167

3' Metepisternum short (w/1 less than 1.50); elytra iridescent, bicolored or not . . 4.

4 (3') Elytron with distinct dark mark near suture, rest of surface rufo-flavous 5.

4' Elytron concolorous, rufo-piceous to piceous 6.

5 (4 ) Specimen from locality in South Africa; pronotum less markedly constricted

posteriorly; dark mark of elytron extended more anterad (Fig. 83B); male with

apical portion of median lobe long, inclined to left (Figs. 88B, C)

H. (Pseudomasoreus) mateui, new species, p. 170

5' Specimen from Tanzania; pronotum markedly narrowed posteriorly; dark mark

of elytron not extended anterad; male with apical portion of median lobe

straight H. (Pseudomasoreus) uluguruanus Basilewsky, p. 168

6 (4') Metepisternum slightly longer than wide (w/1 1.25- 1.38); smaller (SBL ca.

5.00 mm.) H. (Pseudomasoreus) basilewskyi new species, p. 168

6' Metepisternum about quadrate, length less than width; larger (SBL ca. 7.00

mm) H. (Pseudomasoreus) thoracicus, new species, p. 169

Hystrichopus (Pseudomasoreus) reticulatus, new species

Fig. 77

HOLOTYPE female, labelled: COLL MUS TERVUREN Cape Prn: Clanwilliam distr.,

Sederberg VII.1958 1500m J. Smith (MACT).

ion of specific epithet. — A Latin adjective, meaning net-like, in allusion to appearance of

the isodiametric microsculpture of the elytra.

Recognition. — In addition to features cited in the key, the single female of this species

differs from females of H. capicola having a wider pronotum in relation to length (see Table 7).

From females of other species, this one differs in having the pronotum narrow in relation to

head width (see Table 6).

Description. — Values for SBL and for ratios Hw/Pmw, Pl/Pmw and MES; 1/w are presented in Tables 5-8.

Color. Dorsum piceous; epipleura of elytra rufous.

Microsculpture. Dorsum of head and elytra with isodiametric meshes, pronotum with wide, transverse meshes.

Luster. Surface generally dull.

Pronotum. As in Fig. 77. Sides not markedly constricted posteriorly.

Elytra. Humerus rounded, not projected anteriorly. Basal ridge not markedly sinuate.

Ovipositor. Stylomere 2 longer and straighter, and with ensiform setae shorter than in H. mateui.

Geographical distribution. — This species is known only from the type locality in the Union

of South Africa.

Relationships. — The long metepisternum, rounded elytral humeri, and isodiametric

microsculpture of the elytra indicate that this is the most primitive species of Pseudomasoreus

on mainland Africa.

Material examined. — Known only from the holotype.

Cymindine Lebiini of Authors

165

TABLE 5

DATA ABOUT VARIATION IN STANDARDIZED BODY LENGTH (MM) AMONG

EAST AND SOUTH AFRICAN SPECIES OF PSEUDOMASOREUS

MALES FEMALES

TABLE 6

DATA ABOUT VARIATION IN VALUES FOR THE RATIO Hw/Pmw

AMONG EAST AND SOUTH AFRICAN SPECIES OF PSEUDOMASOREUS

MALES FEMALES

Quaest. Ent., 1983, 19 (1,2)

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Ball and Hilchie

TABLE 7

DATA ABOUT VARIATION IN VALUES FOR THE RATIO Pl/Pmw AMONG EAST

AND SOUTH AFRICAN SPECIES OF PSEUDOMASOREUS

MALES FEMALES

TABLE 8

DATA ABOUT VARIATION IN VALUES FOR THE RATIO MES: 1 /w

AMONG EAST AND SOUTH AFRICAN SPECIES OF

PSEUDOMASOREUS

MALES FEMALES

Cymindine Lebiini of Authors

167

Hystrichopus (Pseudomasoreus) capicola (Basilewsky), NEW COMBINATION

Figs. 55A-B, and 78.

Pseudomasoreus capicola Basilewsky 1954c: 93. HOLOTYPE female, labelled: HOLOTYPUS [orange paper];

COL.MUS. CONGO. Cape Colony Dunbrody Co. P. Basilewsky; Pseudomasoreus capicola n. sp. P. Basilewsky det. 1954.

(MACT). TYPE EOCAEITY.- Dunbrody Cape Colony, South Africa.- Basilewsky, 1958a: 296-297, Fig. 40.

Recognition. — Large size, slender pronotum in relation to length and width of head, short

metepisterna, isodiametric microsculpture of elytra, and slightly transverse microsculpture

meshes of pronotum distinguish this species from others of Pseudomasoreus.

Description. — Tables 5-8 provide data about variation in SBL, and in values for ratios Hw/Pmw; Pl/Pmw and

MES: 1/w.

Color. Dorsum piceous, epipleura of elytra rufous.

Microsculpture. Head and elytra with meshes isodiametric, pronotum with meshes slightly transverse.

Luster. Dull.

Pronotum (Fig. 78) Narrow, sides slightly constricted posteriorly.

Elytra. Humeri projected forward, basal ridge markedly sinuate.

Geographical distribution. — This species is known only from the Union of South Africa, in

Cape Province and Basutoland.

Relationships. — These have not been determined. The predominantly isodiametric

microsculpture of the dorsum suggests that this species is primitive. However, the metathorax is

appreciably shortened, suggesting some derivativeness.

Material examined. — In addition to the holotype, we have seen two females labelled: s. Afr.

Basutoland Makhere Mts. 15 miles ENE Mokhotlong 8. IV. 51 No. 268 Swedish South Africa expedition 1950-51 Brinck

Rudebeck 9500 ft; COLL MUS CONGO (ex Lund Mus) Coll. P. Basilewsky (MACT). Six more specimens are known

from this locality (Basilewsky, 1958a: 297), but we have not seen them.

Hystrichopus (Pseudomasoreus) kivuanus (Basilewsky), NEW COMBINATION

Fig. 83A

Pseudomasoreus kivuanus Basilewsky, 1962: 215. HOLOTYPE, female, labelled: HOLOTYPUS [orange paper];

COLL MUS CONGO Tanganyika Terr: Kilimanjaro Marangu Versant S.E. 1800-2200 m 20/27. VII. 57; Resideu de

foret transition [blue paper]; Mission Zoologiq. IRSAC en Afrique orientale P. Basilewsky et N. Leleup; Pseudomasoreus

kivuanus n.sp. P. Basilewsky det. 1960 (MACT).

Recognition. — In addition to character states listed in the key, the female of H. kivuanus is

distinguished by moderate size (SBL- 6.28 mm.) (smaller than the type of H. thoracicus, about

same as females of H. capicola,), and the pronotum very broad in relation to both head width

(Table 6) and pronotum length (Table 7).

Description. — Data about Standardized Body Length, and ratios Hw/Pmw, Pl/Pmw, and MES: w/1 are

presented in Tables 5 to 8.

Color. Head and pronotum piceous, elytra generally rufous with broad, irregular, transverse dark mark in posterior

half (Fig. 83A); elytral epipleura rufous.

Microsculpture. Meshes of head and pronotum isodiametric; meshes of elytra transverse.

Luster. Head and pronotum dull. Elytra shining, but not iridescent.

Pronotum. Very broad, sides rounded, moderately constricted posteriorly.

Elytra. Humeri average, not extended anteriorly as prominent lobes; basal ridge not markedly sinuate. Hind wing with

cells as in Figs. 84A, B.

Basilewsky (1962: 215) stated that adults of this species were apterous. However, wing

rudiments are about half the length of the elytra.

Geographical distribution and habitat. — The single known female was collected in

Tanzania, on Mount Kilimanjaro, in forest, between 1800 and 2200 meters above sea level.

Relationships. — Color pattern and geographical proximity suggest that H. kivuanus and

H. uluguruanus are more closely related to one another than to other species of

Quaest. Ent., 1983, 19 (1,2)

168

Ball and Hilchie

Pseudomasoreus. Their immediate common ancestor was probably from a primitive stock, for

H. kivuanus has very slightly derived microsculpture of the dorsal surface, has retained long

wing rudiments and features associated with wing reduction are not well developed; that is, the

metathorax is large (as shown by long metepisterna), and the elytral humeri are not projected

forward.

Hystrichopus (Pseudomasoreus) uluguruanus Basilewsky, NEW COMBINATION

Figs. 79 and 86

Pseudomasoreus uluguruanus Basilewsky, 1962: 213. HOLOTYPE male, labelled: HOLOTYPUS [orange paper] ;

vert foret/ombrophile dans I’humus [blue paper] ; COLL MUS CONGO Tanganyika Terr: Bunduki, Uluguru Mts., 1300

m. 2.5.1957; Mission Zoologique IRSAC en Afrique Orientale (P. Basilewsky and N. Leleup); Pseudomasoreus

uluguruanus n. sp. P. Basilewsky det. 1960. (MACT). PARATYPE male, similarly labelled to holotype, but collected on

summit of Mt. Kidunda, 1800-1950 m., 3. V. 1957. (MACT). PARATYPE female, similarly labelled to holotype, but

collected on Mgeta, 1300 m., 30.IV- 2.V. 1957. (MACT).

Recognition. — In addition to character states listed in the key, members of this species are

recognized by a combination of small size (SBL less than 6.00 mm.), and cordate pronotum

(Fig. 79).

Description. — Data about variation in Standardized Body Length, and in ratios Hw/Pmw, Pl/Pmw, and MES:

1/w are presented in Tables 5 to 8.

Color. Head piceous; pronotum with disc piceous, broad lateral area rufous; elytra generally rufo-flavous, with broad

irregular transverse dark mark in posterior half, less distinct in similarly marked female of H. kivuanus. Elytral epipleura

flavous.

Microsculpture. Head with meshes isodiametric; pronotum with meshes isodiametric to transverse; elytra, with meshes

transverse, narrow.

Pronotum. Form as in Fig. 79, cordate, sides constricted posteriorly.

Elytra. Humeri produced anteriorly as lobes, basal ridge of elytron markedly sinuate.

Male genitalia.- Median lobe as in Figs. 86A-C: apical portion in dorsal aspect short and broad; internal sac with

basal microtrichial fields short, concentrated near apical orifice of median lobe (two males dissected).

Basilewsky (1962: 213) stated that specimens of this species are winged. However, each

wing comprises a small stub only, no longer than the combined lengths of two abdominal terga.

Geographical distribution and habitat. — This species is known only from the type locality-

The Uluguru Mountains in Tanzania, at elevation of 1800-1900 m. Adults were collected in

mountain forest, in leaf litter.

Relationships. — Color pattern and geographical distribution indicate that H. uluguruanus

and H. kivuanus are sister species. However, H. uluguruanus shares with the more derived

species of Pseudomasoreus transverse microsculpture and iridescent luster of the elytra,

reduction of wings, marked reduction of the metathorax (indicated by short metepisterna), and

produced elytral humeri. Evidently, these states were developed convergently with the same

states in other members of the subgenus.

Hystrichopus (Pseudomasoreus) basilewskyi, new species

Figs. 80 and 87

HOLOTYPE male, labelled: COLL MUS TERVUREN Cape prov. Swellendam distr., Grootvadersbos J. Smith

VII. 1958. (MACT). ALLOTYPE female, same label as holotype. Holotype and female paratypes returned to MACT;

male paratype deposited in CAS.

Derivation of specific epithet. — We take pleasure in naming this species for Pierre

Basilewsky, distinguished specialist of the African carabid fauna and of African biogeography.

Recognition. — In addition to key character states, small size, concolorous elytra, and

metepisternum slightly longer than wide, distinguish members of this species from all others.

Cymindine Lebiini of Authors

169

Males are further distinguished by the very short and broad apical portion of the median lobe

(Fig. 87B, C).

Description. — Data about variation in Standardized Body Length, and in ratios Hw/Pmw, Pl/Pmw, and MES:

1/w are presented in Tables 5 to 8.

Color. Head, rufo-testaceous; pronotum rufous; elytra rufo-piceous to piceous.

Microsculpture. Head with meshes isodiametric. Pronotum with meshes transverse, but not especially narrow; elytra

with meshes transverse, very narrow.

Luster. Dorsum of head dull; pronotum with surface shining, not iridescent. Elytra with surface iridescent.

Pronotum. Form as in Fig. 80, moderately broad in relation to head, sides not markedly constricted posteriorly.

Elytra. Humeri projected anteriorly, basal ridge markedly sinuate.

Male genitalia. Median lobe (Fig. 87A-C) with apical portion very short and broad. Internal sac with basal

microtrichial fields long, extended anteriorly in inverted position.

Geographical distribution. — This species is known only from the type locality, in South

Africa.

Relationships. — This is a markedly derived species, adults having iridescent elytra and

humeri projected. However, the metathorax is only partially reduced, and microsculpture of the

pronotum is not modified enough to provide iridescence. Iridescence of the elytra is an apotypic

character state shared with P. thoracicus and P. mateui, and on this basis we locate P.

basilewskyi in an informal group with these species.

Material examined. — This species is known only from the type series.

Hystrichopus (Pseudomasoreus) thoracicus, new species

Fig. 81

HOLOTYPE female, labelled Grahamstown 14.1.1904 (J. O’N) [handwritten]; Platynus

sp. nov. [handwritten]; Pseudomasoreus sp. ign. [handwritten] South African Museum.

PARATYPES, two females, labelled: [G or A]? T, 15.V,12; S. Africa Cle Deux acc. 67769.

(USNM). And as above, except “Cle Doux” (USNM).

Derivation of specific epithet. — This is an adjectival form of “thorax”, and draws attention

to the large pronotum that is characteristic of specimens included in this species.

Recognition. — Large size (SBL about 7.00 mm.), iridescent pronotum and elytra, and

broad pronotum with wide lateral grooves distinguish adults of this species from other known

Afrotropical species of Pseudomasoreus.

Description. — Data about variation in Standardized Body Length and in values for ratios Hw/Pmw, Pl/Pmw,

and MES: 1/w are presented in Tables 5 to 8.

Color. Dorsum of head, pronotum and elytra piceous, elytral epipleura rufous.

Microsculpture.- Head with meshes isodiametric; pronotum and elytra with meshes transverse, narrow.

Luster. Dorsum of head dull; pronotum and elytra with surfaces iridescent.

Pronotum. As in Fig. 81, cordate, lateral grooves wider than usual.

Elytra. Humeri projected anteriorly, basal ridge markedly sinuate.

Ovipositor. Stylomere 2 average for subgenus Pseudomasoreus.

Geographical distribution. — This species is known from the Union of South Africa, only.

Relationships. — This is a derived species in that its adults are characterized by iridescent

pronotum and elytra, markedly reduced metathorax, and produced elytral humeri. These

character states are shared with adults of H. mateui, new species, which we regard as the sister

species of H. thoracicus.

Quaest. Ent., 1983, 19 (1,2)

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Ball and Hilchie

Hystrichopus (Pseudomasoreus) mateui, new species

Figs. 82, 83B and 88A-C

HOLOTYPE male, labelled: Malvern, Natal; G. A. K. Marshall 1917-55 [blue line in

middle of label]; Cymindide gen et sp nova? Per. [handwritten] (BMNH). ALLOTYPE

female, labelled: NATAL Ekombe For. 39 mi. N. of Kranskop 1520 m. IV. 10.58; E.S. Ross

and R.E. Leech, collectors (CAS). PARATYPE female, labelled: Mbabang Swaziland

[handwritten] ; Pseudomasoreus n. sp. (ap. capicola Basilw) P. Basilewsky det. 1962 other

specimens are necessary [Note: the left mandible missing]. (SAMC).

Derivation of specific epithet. — This is based on the surname of Dr. Joaquin Mateu, who

has published extensively about carabids of the tropics of the world, especially about lebiines.

Recognition. — In addition to the key character states, specimens of this species are

distinguished by small size (SBL less than 6.00 mm.) and cordate pronotum (Fig. 82). Males

have a long apical portion of the median lobe, with apex spatulate (Figs. 88B, C).

Description. — Data about variation in Standardized Body Length, and in ratios Hw/Pmw, Pl/Pmw, and MES:

1/w are presented in Tables 5 to 8.

Color. Head rufo-piceous; pronotum piceous to disc rufo-piceous, with lateral areas rufo-flavous (specimen may be

slightly teneral); elytra with dorsal surface generally rufous to rufo-flavous, medially with dark mark extended along

suture into basal half.

Pronotum. As in Fig. 82, cordate, sides markedly constricted posteriorly.

Elytra. Humeri projected anteriorly, basal ridge markedly sinuate. Hind wing with cells as in Fig. 84B.

Male genitalia. Median lobe (Figs. 88A-C) long, with apical portion long, spatulate. Internal sac with microtrichial

fields long, extended basally.

Ovipositor. Stylomere 2 average for Pseudomasoreus.

Geographical distribution. — This species is known only from two localities in the Union of

South Africa.

Relationships. — This species seems to be the sister species of H. thoracicus, new species.

Subgenus Assadecma Basilewsky, NEW STATUS

Characteristics.- The most striking features of this subgenus are: size of specimens (overall

length 14 to 15 mm., estimated SBL 12 to 13 mm.), relative size of pronotum (almost half the

length of elytra), its form (parallel-sided, base and apex about equal in width), very short and

broad tarsomere 4, armature of the male internal sac (several rows of spines), and long, slender

apical portion of the median lobe. Mandibles are markedly different from those of other

cymindines, but we judge from the illustration (Basilewsky, 1982: Fig. 2c) of the ventral

surface of the right mandible that it is worn. If so, the features exhibited are not of taxonomic

value.

Other character states are shared with members of other subgenera of Hystrichopus. Even

though females of Assadecma are not available, we believe that they will be found to have

stylomeres characteristic of Hystrichopus (sensu latoj, and probably characteristic of subgenus

Pseudomasoreus.

Included species. — The single known species of this subgenus, H. madagascariensis

(Basilewsky), is based on two males, collect at different localities in eastern Madagascar. The

type locality is Hiaraka (1000 meters), on the Masoala Peninsula. The holotype is in MNHP,

the paratype in MACT.

Notes about phylogeny. — Relatively large size of its members, a markedly distinctive

combination of other character states, and seeming isolation of the single known species on

Madagascar suggest that Assadecma is a phylogenetic relic, rather than representing a recently

Cymindine Lebiini of Authors

171

evolved descendant of one of the other extant subgenera. Thus, this taxon is likely to be of

substantial importance in reconstructing the evolutionary history of Hystrichopus (sensu latoj.

Subgenus Hystrichopus {sensu stricto) Boheman, new status

Figs. 44A-D, 52, 56A-B, 64, and 84A-B

Notes about synonymy. — Basilewsky (1954b: 13) listed the following genus-group names as

junior synonyms: Ctenoncus Chaudoir, 1850; and Assotatus, Assoterus, Astus, and Aspastus

Peringuey, 1896. He discussed the nomenclatural history of Hystrichopus and its junior

synonyms (1954a: 15-16). Details are not reviewed here.

Characteristics. — This subgenus was adequately characterized by Basilewsky (1954b:

13-14), for purposes of identification. He did not, however, examine the mandibles (Figs.

44A-D), hind wings (oblongum cell large. Fig. 84A, wedge cell small. Fig. 84B) or stylomere 2

of the ovipositor (Fig. 56A): note the very short ensiform seta).

Notes about classification. — This subgenus includes 58 described, and two undescribed

species. In spite of this diversity, Hystrichopus seems quite homogeneous, so .uch so that

Basilewsky (1954b: 16) chose not to recognize formal subgenera, but instead arranged the

species in two sections and 1 1 groups, to which he also assigned those species that he described

subsequently.

Although many of the most closely related taxa are allopatric, Basilewsky elected not to use

the subspecies category. He argued that more information was required to establish subspecies

than species, and that he did not have the requisite information because of a shortage of

specimens. He also argued that the brachypterous montane vicariads had been isolated long

enough to have differentiated to the species level.

The two sections of Hystrichopus are distinguished by differences in development of the

metathorax, which are in turn associated with wing development: adults of Section I have long

metepisterna, dehiscent elytra, and are either macropterous or brachypterous; those of Section

II have reduced metepisterna, elytra more or less fused together along the suture, and wings

absent.

Although wing loss is characteristic of both groups, Basilewsky stated that processes of

change were probably different: reduction of the flight function in Section II he recognized as

an orthogenetic process, whereas wing loss by members of Section I was adaptive. The

important point to note here is that he conceived the sections (as well as “Groups“) as

phylogenetically valid assemblages.

Nonetheless, in our view, wing loss in both sections is the result of the same process, that of

adaptation. Furthermore, in the absence of additional evidence that Section II is monophyletic,

this taxon must be suspect in a phylogenetic system: reduction of the metathorax could have

taken place in a number of lineages independently. Section I is based on a symplesiotypy, and

may not be monophyletic, either.

Habitat. — Basilewsky (1954b: 18) classified the species as “lapidicoles” or “humicoles”.

Lapidicoles are found under stones at lower elevations in savanna land, mainly in South Africa,

but also at high altitudes in East Africa. Humicoles are residents of mountain forests, and are

found in damp humus (leaf litter, et al. Adults do not show a tendency for reduction of pigment

or eye size, so it seems certain that they are not associated with deep litter, nor are they

troglophiles.

Quaest. Ent., 1983, 19 (1,2)

172

Ball and Hilchie

Altitudinal range extends from near sea level in the south to between 3000 to 4000 meters

on Mount Kilimanjaro, in Tanzania. Generally, the more northern taxa live at elevations above

1800 meters.

Geographical distribution. — The range of this subgenus extends from southernmost South

Africa northward to the southeastern mountains of the Arabian Peninsula, and with an isolated

species (H. nimbanus Basilewsky) on the massif of Nimba, in French Guinea, West Africa.

The range is discontinuous, because the East African species occur on mountains, at high

elevations.

The species of Section II seem to be restricted to South Africa and Zimbabwe. The range of

the species of Section I is co-extensive with range of the subgenus as a whole.

Notes about phytogeny and zoogeography. — Basilewsky did not attempt to reconstruct the

phytogeny of subgenus Hystrichopus in detail. However, he considered the topic in a general

way (1954b: 21-23; see also 1962: 207-212). The distribution pattern (especially the marked

isolation of H. nimbanus in the mountains of French Guinea), suggested to him that the group

is ancient, at least earlier in origin than development of the Red Sea. The group was formerly

widespread in Africa, and had wider ecological tolerances than have the extant stocks. Thus,

the latter are relics- that is, they do not represent a temperate-adapted stock that recently

spread north from South Africa.

On the contrary, it seems to us that the distribution pattern could be subject to a very

different interpretation. However, what must come first is a phylogenetic analysis of the

species, so that relationships can be hypothesized, and thus sense can be made of the different

distributions of macropterous and brachypterous taxa, especially those of Section I. Basilewsky

(1954a: 22) suggested that occurrence of brachypterous forms of Section I at high altitude is a

function of “stenohygrothermy”, for adults of such species live in damp humus in mountains

forests. This may be so, but it is no help in understanding phylogenetic relationships of taxa

with macropterous and brachypterous adults.

Material examined. — Our observations are based on the following material, from

collections of the California Academy of Sciences.

Hystrichopus dorsalis Thunberg. Three males, three females- South Africa Cape Province George X.28.49 B.

Malkin.

Hystrichopus massaicus Basilewsky. Two females, from Kenya. Nairobi XII. 3 1.1 959, E. S. Ross. 17 mi. SE Nakuru

1900 m; E. S. Ross, R. E. Leech.

Hystrichopus rufipes Dejean. Male- South Africa 6mi. NW Port Beaufort 70 m 14.1.1967; E. S. Ross, K. Lorenza.

Subgenus Plagiopyga Boheman, NEW STATUS

Figs. 50, 57, 58, and 85A-B

Notes about synonymy. — Basilewsky (1954b: 80) listed Diaphoroncus Chaudoir, 1850 as a

junior synonym of Plagiopyga, and cites as well earlier references to this subgenus.

Characteristics. — Basilewsky (1954b: 80-81) provided a useful characterization of adults of

this subgenus, contrasting their character states with those of Hystrichopus (sensu stricto). To

these we add: stylomore 2 of ovipositor with very short ensiform seta (Fig. 57), and hind wing

with oblongum cell larger (Fig. 85 A), wedge cell absent (Fig. 85B).

Notes about classification. — Ten species are included in this subgenus. Basilewsky (1954a)

characterized them, but did not provide an infrageneric arrangement: the species are treated in

the sequence in which their names appear in the key (pp. 83-85).

Notes about habitat. — Little information is available about this topic. As Basilewsky

(1954a: 82) noted, testaceous body color and tendency for reduction in eye size exhibited by

Cymindine Lebiini of Authors

173

adults indicates that they avoid light. In fact, adults of some species have been collected from

caves, rodents’ nests, and from under larger rocks. However, the winged condition of adults

indicates that dispersal by flight is normal, so that the species are not confined to subterranean

situations.

Geographical distribution. — This subgenus is predominantly South African, with one

species {H. cyclogonus Chaudoir) ranging as far north as Tanzania, and four species being

known from Zaire, only.

Notes about phytogeny and zoogeography. — These topics have not been addressed

previously, with reference to this subgenus. Without representative material of all species, we

can only make suggestions about a line of investigation to pursue. Because of many shared

similarities with members of the surface-dwelling Pseudomasoreus and Hystrichopus, one can

assume that the ancestor of Plagiopyga must have been a surface-dweller, also, with average

eyes, pectinate tarsal claws (Figs. 52 and 54), and darker color. Extant species whose adults are

thus characterized are near the ancestral stock. Reduction of these features probably occurred

in more derived stocks that had developed a more apotypic mode of existence. These

considerations plus vicariant distribution patterns of species thought to be closely related should

provide the clues necessary to reconstruct the phylogeny of the extant species of Plagiopyga.

Material examined. — Our observations are based on the following material, collected in

South Africa.

H. (Plagiopyga) cymindoides Peringuey. Three females, - E. Cape Province, Congo Caves X.30.49 B. Malkin (CAS).

Three males, female; Stormsriver, W. Humansdorp, 2403415, 4-10. XII. 1981, 525 Peck (from a cave).

H. (Plagiopyga) chaudoiri Basilewsky. Female, Queenstown E. T. Wells 1902-19 (BMNH). Female, Natal Estcourt

G. A. K. Marshall 1917-55 (BMNH).

Subtribe CALLEIDINA

We include here two groups of nominal genera that were originally assigned to the

Cymindina: Anomotarus assemblage- Anomotarus Chaudoir, Lithostrotus Blackburn,

Dromiotes Jeannel, and Cephalotarus Mateu; and the Trigonothops assemblage-

Trigonothops Macleay, Phloeocarabus Macleay, and Diabaticus Bates. We add to the latter

assemblage Speotarus Moore.

By including Anomotarus in the Calleidina, we declare the latter name and Anomotarina

synonymous. Calleidina is the older name, and is thus valid for this group.

Habu (1967: 117), who established the subtribe Anomotarina, recognized a close

relationship of the single included genus with the calleidines, citing as evidence similarity in

form of mandibles and female genitalia. Terrestrial modifications of tarsi of adult Anomotarus

satisfied Habu that this genus should not be included in the Calleidina. However, adults of

some calleidine taxa are basically terrestrial, and do not have structurally generalized tarsi that

one might expect. We conclude that either tarsi modified for climbing were part of the ground

plan of the Calleidina, and were relatively recently lost from some (but not all) groups that

became terrestrial secondarily, or that arboreal modifications occurred after calleidines had

evolved, and thus were not part of the ground plan. This argument is basic for combining

anomotarines and calleidines in a single subtribe.

Recognition. — The following features are diagnostic: labrum transverse; without suborbital

setae; right mandible with broad, edentate retinacular ridge; maxilla with lacinia and galea

sparsely setose; mentum toothed; elytron with umbilical setigerous punctures in continuous line,

penultimate puncture not displaced laterally; tarsomeres broad, apical margin of tarsomere 4

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174

Ball and Hilchie

91

Cymindine Lebiini of Authors

175

Figs. 89-92. Photographs of Calleidina.— Habitus, dorsal aspect, of specimens of Trigonothops: 89, T. (Diabaticus)

australis (Erichson) (SBL= 10.29 mm); 90, T. (Diabaticus) pauper (Blackburn) (SBL = 6.89 mm); 91, T. (Abaditicus)

collaris (Blackburn) (SBL = 7.88 mm); 92, T. (Abaditicus) meyeri, new species (SBL = 7.88 mm).

Quaest. Ent., 1983, 19 (1,2)

Cymindine Lebiini of Authors

177

Figs. 93-96. SEM photographs of structures of Calleidina, genus Trigonothops.—Vi%. 93: T. (Diabaticus) pauper

Blackburn, mandibles-A and C, left, dorsal and ventral aspects, respectively; B and D, right, dorsal and ventral apical

aspects respectively. Figs 94 and 95; T. (Phloeocarabus) nigricollis MacLeay, head and pronotum, respectively, dorsal

aspect. Fig. 96: T. (sensu stricto) longiplaga Chaudoir, left stylomere 2-A, B-lateral aspect; C, D-medial aspect;

E-ventral aspect; F-apical branched seta. Scale bars: 93A-96A, and 96C= 100 /lim, 96B, D, E, F=5 ^ni. Legend, features

of mandibles: m, molar; pm, premolar; prt, posterior retinacular tooth; rr, retinacular ridge; tt, terebral tooth; vg, ventral

groove. Legend, stylomere 2: e, branched seta.

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178

Ball and Hilchie

oom

Cymindine Lebiini of Authors

179

Figs. 97-100. SEM photographs of structures of Calleidina, genus Trigonothops.— Ovipositors, left stylomeres. Figs. 97

and 98, T. (Phloeocarabus) nigricollis Blackburn and T. (Abaditicus) meyeri, new species, respectively, stylomere 2; A, B,

C, D, and E, lateral, lateral (apical portion); medial, medial (apical portion), and ventral aspect respectively. Fig. 99, T.

(Diabaticus) australis Erichson: A, valvifer, and stylomeres 1 and 2, lateral aspect; B-E, stylomere 2— B, lateral aspect; C,

lateral aspect, apical portion; D, medial aspect; E, ventral aspect. Fig. 100, T. (Diabaticus) pauper (Blackburn); A, B, C,

D, and E, lateral (apical portion), medial aspect, medial (apical portion), and apico-ventral aspect, respectively. Scale bars:

97A-C, 98A-C, 99B, D, and 100 A-E = 50 /um; 97B, D, E, 98B, D, E, and 99C and E = 5 ^m. Legend: a, lateral ensiform

seta; b, medial ensiform seta; d, ventral setae; e, branched apical seta.

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Ball and Hilchie

Cymindine Lebiini of Authors

181

Figs. 101-102. Photographs of Calleidina, Trigonothops (Speotarus) lucifuga (Moore).— Fig. 101: habitus, dorsal aspect

(SBL=6.98 mm.). Fig. 102: ovipositor, left stylomere 2— A, lateral aspect; B, lateral aspect, apical portion; C,

apico-ventral aspect. Scale bars=50 um.

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182

Ball and Hilchie

Cymindine Lebiini of Authors

183

Figs. 103-106. Line drawings of Calleidina, genus Trigonothops.—y^^.\Q genitalia— A and B, median lobe, left lateral and

ventral (106B, dorsal) aspects, respectively; C and D, left and right parameres, respectively, ventral aspect, of; 103, T.

(sensu stricto) longiplaga Chaudoir; 104, T. (Phloeocarabus) nigricollis Blackburn; 105, T. (Abaditicus) meyeri, new

species; 106, T. (Speotarus) lucifuga (Moore).

Quaest. Ent., 1983, 19 (1,2)

184

Ball and Hilchie

Cymindine Lebiini of Authors

185

Figs. 1 07-1 1 2. Line drawings of Calleidina.— Wing cells and associated veins— A, oblongum cell, B, wedge cell,

respectively, of: 107, Trigonothops (sensu stricto) longiplaga Chaudoir; 108, Anomotarus {sensu stricto) crudelis

Newman; 109, T. (Phloeocarabus) nigricollis Blackburn; 1 10, T. (Speotarus) lucifuga (Moore); 111, Anomotarus (sensu

stricto) stigmula Chaudoir; \ \2, A. (Dromiotes) maculipennis Mateu. Legend: cells-O, oblongum; W, wedge. Veins— A,

anals; Cu, Cubitals; M, Media; R. Radius.

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186

Ball and Hilchie

sub-truncate or moderately to deeply notched; median lobe of male genitalia with apical orifice

hemiopic, on left and ventral side; ovipositor with stylomere 1 glabrous, stylomere 2

approximately cylindrical, tapered, ensiform setae short or absent; apical or preapical portion

more or less setose; nematoid setae present or absent.

The Trigonothops assemblage

Figs. 89-92

It is easy to see why Phloeocarabus± and Diabaticus were included in the cymindines:

tarsi, though moderately broad, are not markedly so, and tarsomere 4 is only notched, without

well developed lobes and pads of specialized setae. The tarsi of Trigonothops adults are clearly

different from those of cymindines, but in other features this group and the former two seem

quite close to one another.

It is also easy to see why Moore (1964; 73) placed Speotarus (Fig. 101) near Anomotarus:

in external features, adults of the two groups look much alike. However, the stout tarsi of

described Speotarus adults suggest a calleidine affinity, and this is borne out by structure of

the male genitalia and stylomeres of the ovipositor.

Adults of the four calleidine groups of the Trigonothops assemblage exhibit so much

similarity to one another that they are here included in a single genus. Furthermore, these

groups are confined to the same zoogeographic area, the Australian Region, and this provides

additional evidence for inferring close relationship. Additional details are provided below.

Trigonothops MacLeay

Trigonothops MacLeay, 1864: 110. GENERITYPE: Calleida pacifica Erichson: 1842: 124 (original designation). -

Chaudoir, 1877: 221.- Sloane, 1898:493. 1920: 170.-Csiki, 1932: 1488.- Darlington, 1968: 184.

Phloeocarabus MacLeay, 1871: 85. GENERITYPE: Phloeocarabus mastersi MacLeay, 1871: 85 (monotypy).-

Sloane, 1898:494- 499.- Blackburn, 1901: I12.-Csiki, 1932: 1488.- Darlington, 1968: 183- 184 NEW SYNONYMY.

Notoxena Chaudoir, 1877: 226. GENERITYPE: Trigonothops nigricollis M?ichQ2Ly, 1864: 111 (monotypy).- Sloane,

1898: 499.- Darlington, 1968: 183- 184.

Diabaticus Bates, 1878: 324. GENERITYPE: Plochionus australis Erichson, 1842: 124 (monotypy)- Csiki, 1932:

1489. NEW SYNONYMY.

Speotarus Moore, 1964: 71. GENERITYPE: Speotarus lucifugus Moore, 1964: 73 (original designation).- Matthews,

1980: 10. NEW SYNONYMY.

Abaditicus, new subgenus. GENERITYPE: Diabaticus collaris Blackburn, 1901: 111 (here designated).

Notes about classification. — Inclusion of these taxa in a single genus makes the latter

difficult to define in terms of external features. However, we feel confident that this assemblage

is monophyletic, and we feel that it is more important to emphasize relationships inferred from

complex internal structures than to emphasize differences, which, though easily perceived,

seem of less importance. Attention is thus drawn to an underlying unity, and we hope that this

will stimulate future workers on Australian carabids to undertake study of the group as a

whole.

To draw attention to divergence within Trigonothops, we recognize four previously named

assemblages as subgenera: Trigonothops (sensu strictoj, Phloeocarabus, Diabaticus, and

Speotarus. However, the nominal genus Diabaticus seems to be paraphyletic, including two

species that are less closely related to its Xy^Q;D. australis Erichson, than to other group of

Trigonothops. Therefore, we erect a fifth subgenus, Abaditicus, no previously published names

being available.

Cymindine Lebiini of Authors

187

The generitype of Notoxena Chaudoir is included in Phloeocarabus± . Chaudoir (1877:

226), when he described Notoxena, did not cite MacLeay (1871). Hence, he must have been

unaware that a genus had already been proposed that would include T. nigricollis.

Descriptive notes. — To the characterization of Trigonothops by Darlington (1968: 183), we add the following.

Adults with eyes large, prominently bulged, temples small (Fig. 94), or only slightly convex, with temples well developed

(Fig. 101). Tarsomere 4 notched or bilobed; tarsal claws pectinate or smooth. Male genitalia with median lobe hemiopic,

internal sac with large flagellum- like sclerite (Figs. 103-106). Ovipositor with stylomere 1 asetose, stylomere 2 cylindrical,

ensiform setae two or absent, ventral surface setose; apical portion extended and attenuate (Figs. 97A, C) or not (Figs.

98A, C).

Way of life.— Adults of Trigonothops (sensu stricto), Phloeocarabus, ± and Abaditicus are

arboreal. We do not know where adults of Diabaticus live, but we assume that they spend at

least part of their lives on trees. Adults of Speotarus are known only from caves.

Evolutionary trends. — If, as we believe, calleidines are basically arboreal, then the arboreal

groups of Trigonothops are likely to be closer to the ancestral stock of the genus, with the

cave-inhabiting Speotarus being the more remote. If this is correct, the smooth tarsal claws and

rather flattened eyes of adults of Speotarus are probably apotypic, denticles having been lost

from the ancestral stock of this subgenus, and the compound eyes reduced.

Key to Subgenera of Trigonothops

1 (0) Tarsal claws smooth; eyes slightly convex, temples large (Fig. 101); pronotum

with narrow lateral grooves, only slightly transverse (Fig. 101)

Speotarus Moore, p. 1 9 1

V Tarsal claws pectinate; eyes markedly convex and bulged, temples small (Figs.

89-92, and 94); pronotum with wider lateral grooves, more transverse (Figs.

89-92, and 95) 2

2 (F) Tarsomere 4 cleft apically, with pair of large lobes, ventrally with modified

setae Trigonothops (sensu stricto}, p. 188

T Tarsomere 4 notched apically, lobes short, without modified vestiture ventrally .

3

3 (2') Head with pair of distinct longitudinally directed lateral ridges, especially

prominent between supraorbital setigerous punctures, and extended to posterior

pair (Fig. 94); eyes very large, entire lateral area of head occupied; pronotum

very broad, basal margin distinctly lobed (Fig. 95)

MacLeay, p. 188

3' Head without longitudinally directed ridges, or these shorter, not extended to

posterior pair of supraorbital setigerous punctures; temples short, eyes average

in size (Figs. 89-92), though prominent; pronotum narrower, more elongate,

basal margin convex, but not distinctly lobed 4

4 (30 Head sharply constricted posteriorly, in form of neck (Figs. 91, 92); elytron with

microsculpture meshes isodiametric, not transverse

Abaditicus, new subgenus, p. 189

4" Head not sharply constricted in form of neck (Figs. 89, 90); elytron with

microsculpture meshes transverse Diabaticus 188

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188

Ball and Hilchie

Subgenus Trigonothops (sensu stricto), NEW STATUS

Figs. 96A-F, 103A-B and 107 A-B

Descriptive notes. — To Darlington’s (1968: 184-185) characterization of this taxon, we add the following, based

on study of 12 specimens (CAS) of two species, from various localities in Queensland and New South Wales. Hind wing

with average oblongum cell, and wedge cell long, narrow (Figs. 107A, B) or both cells reduced (Figs. 108A, B). The basal

portion of the flagellar sclerite of the internal sac is almost as long as the main part (Figs. 103 A, B), that is, this structure

is relatively short. Stylomere 2 of the ovipositor is as follows: form as in Figs. 96A, C, base broad, tapered markedly about

half length, then paralled-sided; apex blunt; microsculpture with meshes distinct at base, isodiametric, each scale with

acuminate tip; in apical 0.50, meshes elongate microlines shallow; apical 0.33 with few spines (Figs. 96D, E), one large

ensiform seta dorso-laterally (Fig. 96D), and apical 0.20 with branched (Figs. 96B and F) and unbranehed long setae

extended from microscales.

Subgenus Phloeocarabus MacLeay NEW STATUS

Figs. 94, 95, 97A-E, 104A-B, and 109A-B

Descriptive notes. — To Darlington’s (1968: 183) characterization of this taxon, we add the following, based on

examination of his series of T. nigricollis, from various localities in New Guinea and Australia. Wing with cells large

(Figs. 109A, B). The internal sac of the male genitalia contains a large reverse “J” shaped sclerite (Figs. 104A, B).

Stylomere 2 of the ovipositor as in Figs. 97A-E, elongate, tapered gradually to narrow apex; microsculpture rather

irregular, microlines fine, meshes isodiametric basally, elongate apically, (Figs. 97B, C); few setae mainly on lateral and

dorsal surfaces, about half way between base and apex, without ensiform setae; short seta-like projections extended from

microscales in apical 0.20 (Figs. 97B, D, and E).

Subgenus Diabaticus Bates, NEW STATUS

Figs. 89, 90, 93A-D, 99A-E and lOOD-E

Having had the opportunity to see specimens of the three described species that were

previously included, and of a fourth related but undescribed species, and having reached the

conclusion that two subgenera are represented rather than one, it seems appropriate to offer a

more extended analysis of this complex.

Structures that seem best to show relationships in Trigonothops are male genitalia and

ovipositor. Unfortunately, we have both males and females of only one species of the Diabaticus

complex, T. meyeri, new species. T. australis (Erichson) and T. pauper (Blackburn) are

represented by females, only; and T. collaris, by a single male. However, because of the general

pattern that we perceive, we feel certain that the missing pieces of evidence will fit in, when

they are eventually found.

Stylomere 2 of ovipositors of T. australis (Fig. 99) and T. pauper (Fig. 100) is markedly

different in form and sculpture from stylomere 2 of a T. meyeri female (Fig. 98). The latter

stylomere is more like that of Trigonothops (sensu stricto) and Phloeocarabus females. We

think it likely that T. australis and T. pauper females exhibit the plesiotypic form, and that

the other forms are apotypic.

With T. meyeri, we group T. collaris Blackburn because of striking similarities in the

genitalia and in form of head.

Descriptive notes. — Form as in Figs 89 and 90.

Color: body piceous, appendages rufous, elytra concolorous. Microsculpture and luster: dorsum of head (including

labrum and clypeus) with meshes isodiametric, surface dull; pronotum with meshes transverse, surface shining, but not

iridescent; lateral and ventral thoracic sclerites (including mesepisterna) with meshes transverse; abdominal sterna with

meshes transverse, surface iridescent; scutellum with meshes isodiametric. Dorsal surface glabrous (except standard fixed

setae), or sparsely setose. Pronotum subcordate, sides sinuate posteriorly, margins broadly curved upward; posterior angles

Cymindine Lebiini of Authors

189

approximately right. Ovipositor: stylomere 2 as in Figs. 99 and 100, blade-like, apical 0.33 straight (Fig. 99C), or

slightly twisted (Fig. lOOE); microsculpture predominantly of isodiametric meshes (Figs. 99D-E), elongate on

apico-medial surface of D. pauper (Fig. lOOD), lines deep, each scale with acuminate tip; apico-dorsal 0.33 with 15-20

trichoid setae, two ensiform setae pre-apically, one lateral, one medial, short (Fig. 99E), or longer (Fig. lOOE); apex

with (Figs. lOOD, E) or without (Figs. 99C, E) fine setae extended from microscales.

Geographical distribution. — This subgenus is known from Tasmania and southeastern

Australia, only.

Key to Species of Subgenus Diabaticus

1 (0 ) Dorsum of body and dorsal surfaces of tarsomeres generally punctate and

setose; metepisternum short, with anterior and lateral margins subequal

T. (Diabaticus) pauper, Blackburn, p. 189

r Dorsum of body and dorsal surfaces of tarsomeres glabrous, generally

impunctate; metepisternum long, lateral margin longer than width at anterior

margin T. (Diabaticus) australis (Erickson), p. 189

Trigonothops (Diabaticus) australis (Erichson), NEW COMBINATION

Figs. 89 and 99A-E

Plochionus australis Erichson, 1842; 124.

Diabaticus australis-. Bates, 1878: 324.- Blackburn, 1901: 17.-Csiki, 1932: 1489.

Descriptive notes. — Standardized Body Length of two females: 8.60 and 8.96 mm. Values for Vwm/Hw 0.59

and 0.62. Lateral margins of pronotum only slightly elevated; broad lateral grooves markedly narrowed near anterior

setigerous punctures, these in bottom of lateral grooves, clearly removed from margin. Elytron with basal ridge complete,

extended from humerus to suture, near scutellum.

Bates (1878: 325) noted the superficial similarity in body form between adults of this

species and those of Cymindis (Pinacodera) punctigera (LeConte).

Specimens examined. — Two females (BMNH), both determined by T.G. Sloane: one labelled Hobart, 91-88

[ovipositor dissected]; the other, V D Ld 77- 19; 146 [abdomen lacking].

Trigonothops (Diabaticus) pauper (Blackburn), NEW COMBINATION

Figs. 90 and lOOA-E

Diabaticus pauper Blackburn, 1901: 111. HOLOTYPE female, labelled: Tazm [red print] T; Type [circular label,

ringed with red]; Blackburn Coll 1910- 236; Diabaticus pauper, Blackb. [handwritten] (BMNH).- Csiki; 1932: 1489.

Descriptive notes. — Form as in Fig. 90. Standardized Body Length (five females): 5.60-(6.17)- 6.76 mm. Range

of values for ratio width of neck to width of head: 0.53-0.65. Dorsal surface of frons and pronotum laterally rugulose, and

elytral striae deeper than in adults of T. australis. Pronotum more narrowed posteriorly, and lateral margins more

elevated; lateral margins of elytra crenulate; humeri narrowed (associated with wing loss and reduction of metathorax),

and marginal ridge terminated near base of interneur 4. Ovipositor with stylomere 2 as in Figs. lOOA-E.

Geographical distribution. — This species is known from Tasmania, only.

Material examined. — in addition to the holotype, we have seen four females (BMNH): two labelled Franklin,

Tasmania, 91-88; and two labelled Hobart, 91-88.

Abaditicus, new subgenus

Figs. 91, 92, 98A-E, and 105A-D

This taxon is established to include Diabaticus collaris (Blackburn) and Trigonothops

meyeri, new species.

Quaest. Ent., 1983, 19 (1,2)

190

Ball and Hilchie

Derivation of subgeneric name. — This is an anagram of Diabaticus, the name of the group

to which T. collaris was originally assigned.

Recognition. — The markedly constricted posterior part of the head (Figs. 91 and 92) is

sufficient to distinguish adults from those of other subgenera of Trigonothops. Additionally,

the basal ridge of the elytron is extended only to the base of interneur 3; females have stylomere

2 of the ovipositor short and stout and without ensiform setae (Fig. 98); and males have a

moderately long sclerite (Fig. 105B) in the internal sac.

Descriptive notes. — Form as in Figs. 91 and 92.

Color: body and appendages rufous; elytra concolorous (rufous) or bicolored (Fig. 92). Microsculpture and luster:

dorsum of head (including labrum and clypeus) with meshes isodiametric, surface dull; pronotum with meshes transverse,

surface shining but not iridescent; most lateral and ventral thoracic sclerites with meshes transverse, mesepisternum with

meshes isodiametric; abdominal sterna with meshes transverse, surface iridescent. Dorsal surface glabrous (except

standard fixed setae). Pronotum subcordate, sides sinuate posteriorly, lateral margins broadly curves upward; posterior

angles approximately right. Elytron with basal ridge terminated near base of interneur 3, not extended to sutural margin.

Internal sac of male genitalia with large, reverse “J” -shaped sclerite (Fig. 105B).

Stylomere 2 of ovipositor as in Figs. 98A-E, short, broad, constricted slightly medially, broadened apically, apical

margin very broad (Eig. 98 B). Microsculpture meshes isodiametric or slightly elongate, microlines generally distinct,

scales without acuminate tips. Apical 0.25 with setae on lateral and dorsal surface, but not on medial surface, without

ensiform setae; apex with long setae extended from microscales.

Geographical distribution. — This subgenus is known from southeastern Australia

(Victoria), only.

Relationships. — We believe Abaditicus is the primitive sister group of the subgenus

Phloeocarabus, ± based on inferred transformation series in armature of the internal sac, and

details of stylomere 2.

Key to Species of SwhgtnvLS Abaditicus

Elytra concolorous, rufo-piceous

T. (Abaditicus) collaris (Blackburn), p. 190

Elytron sharply bicolored, most of surface piceous, with apex and extensive area

of disc rufous (Fig. 92)

T. (Abaditicus) meyeri, new species, p. 190

Trigonothops (Abaditicus) collaris (Blackburn), NEW COMBINATION

Fig. 91

Diabaticus collaris Blackburn, 1901: 111. HOLOTYPE male labelled: 6954 H. Wick [red print] T; Type [circular,

ringed with red]; Blackburn coll 1910- 236; Diabaticus collaris Blackb. [handwritten] (BMNH).- Csiki, 1932: 1489.

Descriptive notes. — Eorm as in Eig. 91. Standardized Body Length 6.88 mm. Value for ratio width of neck to

maximum width of head 0.49. Pronotum with lateral grooves broader than in adults of T. australis, and hardly narrowed

anteriorly; anterior pair of setigerous punctures nearly marginal. Median lobe as in Pig. 105, internal sac with reverse

“J”-shaped sclerite.

Material examined. — Holotype, only.

Trigonothops (Abaditicus) meyeri, new species

Figs. 92, 98A-E, and 105A-E

1 (0)

r

Type material.— ElOLOTYPE male, labelled: Woodhouse Ck., Nunniong Pit. Vic. 16.5.66.

P. Meyer; under bark of E. delegatensis (CSIRO). Three paratypes, from the same locality:

Cymindine Lebiini of Authors

191

collected on May 16- female (CAS); collected on May 26- male (BMNH); female (MCZ).

Derivation of specific epithet. — From the surname of the collector, Peter A. Meyer,

Heidelberg, Victoria, Australia, to whom the senior author is grateful for the gift of these and

other specimens.

Recognition. — This is the only known species of Abaditicus whose adults have spotted

elytra.

Description. — Character states of subgenus, and the following. Form as in Fig. 92. Standardized Body Length,

males 6.9- 7.12 mm., females 6.88- 7.08 mm. Body form Calleida- like. Neck evident (W vertex min./Hw males 0.52-

0.53, female 0.52. Hw/Pl- males, 0.87- 0.89, females, 0.86- 0.92; Pl/El- males, 0.300- 0.32, females 0.29- 0.30.

Color. Appendages and body except elytra rufous; elytron with following rufous- epipleura, lateral groove, apical 0.16,

and irregular discal area from interval 2 to 6, extended to humerus on interval 5; following black- interval 1, in basal 0.84,

triangular area near scutellum, transverse band in apical 0.33, and intervals 7- 9 throughout most of length.

Microsculpture. As described for subgenus. Surface slightly shining, pronotum more so than head or elytra.

Fixed setae. Average for Calleidina: both pairs of pronotal setae on lateral margins.

Head. Frons and anterior part of vertex depressed. Frontal impressions extended diagonally to anterior pair of

supraorbital setigerous punctures. Eyes average for subgenus; occipital area markedly constricted. Mouthparts average,

including mental tooth, axiniform ultimate labial palpomeres, and bisetose penultimate palpomeres.

Pronotum. Moderately transverse, anterior margin shallowly concave, posterior margin convex, but not clearly lobed;

lateral margins distinctly to slightly sinuate; anterior angles broadly rounded, posterior angles about right; lateral margins

elevated, more broadly so posteriorly; disc broad, only slightly convex medially; median longitudinal impression sharply

delimited, extended from near anterior to near posterior margin; posterior-lateral impressions indistinct, shallow, broadly

continuous with broad lateral grooves.

Elytra. Humeri broadly rounded, apical margin subtruncate; basal ridge terminated near base of interneur 3, not

extended to parascutellar setigerous puncture; interneurs shallow, intervals hardly convex.

Male genitalia. As in Figs. 105A-D, average for Calleidina.

Ovipositor. Stylomere 2 as in Figs. 98A-E.

Notes about habitat. — According to the labels, specimens in the type series were collected

under bark of a eucalyptus tree. Probably, then, this species is arboreal. Interestingly the color

pattern of these specimens is like that of many arboreal Australian lebiines (for example,

Trigonothops longiplaga Chaudoir). Darlington (1971: 250-251) suggested that mimicry

might be involved as an explanation for similarity in color pattern exhibited by some tree

trunk-inhabiting lebiines, though he did not refer specifically to the pattern characteristic of T.

meyeri. This suggestion seems reasonable to us, and we extend it in terms of Mullerian

mimicry, to the many groups of Australian lebiines that are colored like adults of T. meyeri.

Erwin (1978 and 1979) discussed tests of defense mechanisms that showed them to be powerful

for adults of Agra and other lebiines. This is supporting evidence that this group of insects has

the necessary equipment to form the basis for development of complexes of protected mimics.

Geographical distribution. — This species is known only from the type locality, which is in

the general range of the previously described species of Abaditicus.

Phylogenetic relationships.-— Adults of this species share with those of T. collaris a head

with constricted occipital area, and elytra with basal ridges incomplete. These synapotypic

features satisfy us that these two species are more closely related to one another than to the

other known species of Trigonothops.

Subgenus Speotarus Moore, new status

Figs. 101, 102A-C, 106A-D, and llOA-B

Descriptive notes. — The following details are added to the original description of Speotarus (Moore, 1964: 91).

These notes are based on two specimens of T. lucifugus Moore, 1964; male, Cocklebiddy Cave, Eucla Basin, S. Australia,

bat piles 12.1.66, J. Lowrey; female, bat cave, Naracoorte, 9 Mar, 1963, E. Hamilton-Smith.

Habitus as in Fig. 101. Standardized Body Length, male 6.86 mm., female, 7.06 mm.

Quaest. Ent., 1983, 19 (1,2)

192

Ball and Hilchie

Microsculpture. Dorsum of clypeus and anterior part of frons smooth, microlines not evident, vertex with meshes

isodiametric, microlines shallow; thoracic and abdominal sclerites, and elytra with meshes transverse. Surface generally

shining, especially head.

Head. Eyes though extensive in area, only slightly convex, not protuberant (Fig. 101).

Pronotum. Narrow, slightly transverse, lateral grooves narrower than in Trigonothops (sensu stricto) adults.

Legs. Anterior and middle femora with more setae than usual, posterior face of middle femur with more than 12

setae.

Wings. Completely developed, not reduced. Oblongum cell fusiform (Fig. IlOA), wedge cell very small (Fig. HOB).

Median lobe of male genitalia hemiopic, apical orifice to left and ventrad (Figs. 106A, B). Internal sac with reverse

“J”-shaped sclerite, and small sclerite. Parameres as in Figs. 106C, D.

Ovipositor (Figs. 102A-C). Stylomere 1 asetose. Stylomere 2 elongate, apical portion tapered, preapically with pair

of ensiform setae (one lateral, one medial), and several trichoid setae on ventral surface. Microsculpture with

sculpticells elongate, each with small spine directed apically; microsculpture otherwise simple.

Notes about way of life. — Also included in subgenus Speotarus is a second species, T.

princeps (Moore, 1964). Both species are known only from caves. Although pale color of

integument and rather reduced eyes are cavernicolous adaptations, the normally proportioned

metathorax and rather long wings of Speotarus adults suggest that they are not troglobitic.

Moore {in litt.) advised us: “the species are undoubtedly troglophiles (guanophiles) and are

plentiful in certain caves, notably on the Nullarbor Plain, where there are no trees and no

surface litter.” Further, he stated that the beetles have not been found in the course of extensive

surveys of the litter-fauna, in southern Australia. This counters our first thought that habitus of

the beetles suggests adaptation to life in deep litter.

Moore (personal communication) advises us that additional specimens of Speotarus have

been found in additional caves. These beetles exhibit some differences from the previously

described species, and may represent undescribed taxa.

Evolutionary considerations. — In the letter cited above, Moore advanced an hypothesis to

explain the cave-inhabiting way of life of a stock that might have been arboreal. He suggested

that the extant species of Speotarus were derived from tree-dwelling calleidines that took up

life in tree-roosting bat colonies, and became adapted to living in association with guano. It

would be but a rather short evolutionary step from that stage to life in caves inhabited by bats.

As he noted, support for this hypothesis would come from discovery of Speotarus specimens in

association with arboreal bats. We think that Dr. Moore’s idea has merit, and hope that he

succeeds in his quest for confirmatory evidence.

The Anomotarus assemblage

For reasons stated below, we combine the named genera of this complex in a single genus,

Anomotarus Chaudoir. Further, we have considered seriously the possibility of a close

relationship between Anomotarus {sensu lato) and Trigonothops {sensu lato). However, we

were unable to identify synapotypic features to support this alliance.

Anomotarus Chaudoir

Figs. 108 A, B, and 113-117

Anomotarus Chaudoir, 1875; 48. GENERITYPE; Anomotarus blivaceus Chaudoir, 1875; 48 (monotypy).- Sloane,

1898: 494.- 1917: 435.- 1920: 170.-Csiki, 1932; 1492- 1493.- Jedlicka, 1963: 300, 450.- Moore, 1964: 73.- Habu, 1967:

118- 121.- Darlington, 1968: 186- 191.- Mateu, 1970b: 148.- 1972; 44.- Moore, 1967a: 183- 184.

Uvea Fauvel, 1881: CXVIII. GENERITYPE: Cymindis stigmula Chaudoir, 1852: 57 (monotypy).

Nototarus Chaudoir, 1875: 19. GENERITYPE: Nototarus australis Chaudoir, 1875: 19 (monotypy).- Sloane, 1898:

494.-Csiki, 1932: 1492.- Moore, 1963:442.- 1967b: 442-445.- Darlington, 1968: 185- 186. NEW SYNONYMY.

Lithostrotus Blackburn, 1894; 200. GENERITYPE; L. coerulescens Blackburn, 1894: 200 (monotypy).- Sloane,

1898: 494.- Csiki, 1932; 1492. NEW SYNONYMY.

Cymindine Lebiini of Authors

193

Lestianthus Sloane, 1894: 451. GENERITYPE: Lestianthus sculpturatus Sloane, 1894: 452 (monotypy)

{ = Lithostrotus coerulescens Blackburn).

Dromiotes Jeannel, 1949: 914. GENERITYPE: Lebia stigmula Fairmaire, 1901: 126 {—A. jeanneli Mateu, 1972:

47, not A. stigmula Chaudoir, 1852: 57) (original designation).- Mateu, 1972: 44.

Cephalotarus Mateu, 1970b: 150. GENERITYPE: Cephalotarus maculipennis Mateu, 1970b: 151 (monotypy).-

1972: 46.

Notes about names and classification. — By inclusion in Anomotarus of Dromiotes, the

type species of the latter {Lebia stigmula Fairmaire, 1901) becomes a secondary junior

homonym of A. (sensu stricto) stigmula (Chaudoir, 1852). For the name L. stigmula

Fairmaire, therefore, Mateu (1972: 94) proposed the new name Anomotarus (Dromiotes)

jeanneli. Character states diagnostic for these taxa seem too slight and too few to warrant

ranking as genera. Thus, we think it best to include all of the species in a single genus.

However, we also think it desirable to indicate the pattern of divergence in the genus by

recognition of three subgenera: Dromiotes Jeannel; Anomotarus (sensu stricto); and Nototarus

Chaudoir (including Lithostrotus Blackburn).

Justification of synonymy of Nototarus and Lithostrotus is required. Distinctive features of

adults of Lithostrotus are: dorsal integument metallic blue, surface densely, coarsely punctate

(Fig. 113), and setose, with microsculpture generally effaced; eyes small, temples large;

Pronotum (Fig. 113) with very sharp posterior angles and sharply defined basal lobe. Our

material of Nototarus includes adults of eight species (mostly unnamed). None exhibit metallic

color, but two have a pattern of punctation similar to that of Lithostrotus, with the pronotum

similar in form, and eyes similarly reduced. Adults of two species are less coarsely punctate,

and are glabrous; the others are impunctate, and have rather larger eyes. In brief, the

differences are bridged between the Lithostrotus adults and those of the more typical

Nototarus species. Thus, a transformation series seems to be indicated, with one end

represented by Lithostrotus. It will no doubt be desirable to recognize species groups, in

conjunction with revision of the species of this subgenus.

Key to Subgenera of Anomotarus (sensu lato)

1 (0 ) Mentum toothed. Metepisternum elongate, lateral margin longer than width at

anterior margin. Elytron with microsculpture meshes more or less transverse.

Eyes large. Internal sac with or without sclerite. Stylomere 2 of ovipositor with

sculpticells flat (Figs. 114 and 115) 2

V Mentum edentate. Metepisternum short, lateral and anterior margins subequal.

Wings reduced. Elytron without microlines, or meshes transverse. Eyes reduced,

though head large. Internal sac with large sclerite. Stylomere 2 of ovipositor

with surface rugose, sculpticells raised as keels; pair of long slender, curved

setae near apex (Fig. 1 16D) or not (Fig. 1 17D)

Subgenus Nototarus Chaudoir.

2 (F) Wing with oblongum cell broad (Fig. 112A), wedge cell small (Fig. 112B).

Internal sac of male with well developed sclerites, flagellum-like or not.

Stylomere 2 without long, curved setae (Figs. 1 15A-E). Species Afrotropical . .

Subgenus Dromiotes Jeannel.

1! Wing with oblongum cell narrow, wedge cell absent (Figs. lllA, B); or small

(Fig. 108B). Internal sac without sclerites. Stylomere 2 of ovipositor with long,

slender, curved setae (Figs. 1 14B and D). Species Oriental or Australian

Anomotarus (sensu stricto).

Quaest. Ent., 1983, 19 (1,2)

194

Ball and Hilchie

Fig. 113. Photograph of CdAXcxdmz.— Anomotarus (Nototarus) coerulescens (Blackburn), habitus, dorsal aspect

(SBL = 4.89 mm.).

Cymindine Lebiini of Authors

195

Figs. 114-117. SEM photographs of Calleidina, genus Anomotarus.—O\'\pos\ior%, left stylomeres 1 and 2, or 2, only. A, B,

C, D, and E— lateral, lateral (apical portion), medial, medial (apical portion), and apico-ventral aspects, respectively, of:

1 14, A. (sensu stricto) stigmula Chaudoir; 115 A. (Dromiotes) maculipennis Mateu; 1 16, A. (Nototarus) coerulescens

(Blackburn); \ \1,A. (N.) tumidiceps (Blackburn). Scale bars: 1 14A, C, E, 1 15A, C, E, 1 16A, B, and 1 17A, C = 50 Mm;

1 14B, D, 1 15B, D, 1 16C, D, E, and 1 17B, D, E = 10 Mm. Legend: a, lateral ensiform seta; b, medial ensiform seta; c,

sensory furrow; d, nematoid setae; e, branched apical seta; SI, stylomere 1 ; S2, stylomere 2.

Quaest. Ent., 1983, 19 (1,2)

196

Ball and Hilchie

Darlington (1968: 185-187) provided useful descriptions of Anomotarus and Nototarus, to

which we add that species with brachypterous members and with mental tooth probably belong

to Anomotarus (sensu stricto). Hence, both of these subgenera have brachypterous members.

Moore (1964: 73) suggested that it may be necessary to erect a new genus to include several

species that seem to have adult characteristics similar to those of A. tumidiceps Blackburn.

The seeming scarcity of specimens of A. coerulescens makes it desirable to have a detailed

account available, for the benefit of workers on Australian carabids.

Anomotarus (Nototarus) coerulescens Blackburn, NEW COMBINATION

Figs. 113, and 1 16A-E

Lithostrotus coerulescens Blackburn, 1894: 200. HOLOTYPE female, labelled: 5274 Viet [red print] T; Type

[circular, ringed with red]; Blackburn coll 1910- 236; Lithostrotus coerulescens Blackb [handwritten] (BMNH).

Lestianthus sculpturatus Sloane, 1894: 451 (type not seen).- 1898: 494.

Lithostrotus planior Blackburn, 362. HOLOTYPE female, labelled: B7 MCS 7755 [red print] T; Type HT [circular,

ringed with red]; Australia Blackburn coll BM 1910- 236; Lithostrotus latior Blackb [handwritten]; This must be the type

of planior. The name latior was evidently written in error. No such name as latior has been published. A.M. Lea 6/9/12

[handwritten] (BMNH). TYPE LOCALITY: Australia New South Wales, Blue Mountains, 3000 feet.- - Lea, 1912:

xxviii. NEW SYNONYMY.

Notes about synonymy. — We have seen the above-listed holotypes. They are so similar to

one another that it seems they must be conspecific, and we regard them as such.

Recognition. — The following combination of character states sets adults of this species

apart from others included in Anomotarus: dorsum metallic blue-green; microlines on dorsal

surface not visible at magnification of SOX, except labrum with meshes isodiametric; dorsal

surface punctate, each puncture with long seta; elytral intervals each uniseriately punctate,

each puncture extended about width of interval, except punctures of interval 1 smaller; frontal

impressions of head, median longitudinal impression of pronotum, and scutellar interneur very

deep; eyes small, temples tumid, large; pronotum markedly cordate, base sharply lobed,

posterior angles acute; metasternum short, metepisternum quadrate; hind wings short stubs;

elytra with humeri sharply ridged, projected forward; stylomere 2 of ovipositor with

microsculpture very coarse (Figs. 116A-E), extended apically as ridges and spines.

Standardized Body Length 3.80- 4.04 mm. (three specimens).

Notes about relationships. — Adults of this southeastern Australian species most closely

resemble those of a probably undescribed species, known from a single female collected in

southern West Australia (Margaret River; MCZ). The single male of A. angusticollis (Sloane)

(Wiluna; MCZ) shares with the above species the coarse, generally punctate dorsum. However,

it is much larger, and the basal lobe of the pronotum is less distinctly developed.

Subtribe DROMIINA

The exact composition of this subtribe has not been settled. Jeannel (1949: 990) chose to

include in the subfamily Dromiitae (family Lebiidae) the dromiines (sensu stricto) and the

demetriines, ranking these groups as tribes. He excluded Apristus Chaudoir, placing this genus

in the family Lionychidae. Habu (1967) chose to rank demetriines and dromiines as subtribes

of Lebiini, and to include the lionychid genera in the Dromiina. We elect to follow Habu,

though we exclude Celaenephes Schmidt- Goebel.

Jeannel (1949: 915) also erected the tribe Singilini (subfamily Lebiinae) to include a

number of genera whose adults are characterized by small size, and pale, hairy integument.

Cymindine Lebiini of Authors

197

Mateu (1963) revised this complex, pointing out that three groups were included, which he

ranked as tribes: Lichnasthenini, Singilini (sensu stricto), and Somotrichini. Ball (1975: 152)

transferred the somotrichine to the subtribe Pericalina {sensu lato). It seems to us that

lichnasthenines and singilines, as understood by Mateu, can best be accommodated in the

Dromiina, and we place them here. For the present, the names Singilini and Lichnasthenini are

treated as junior synonyms of Dromiina.

On the basis of shared similarities in details of ovipositor sclerites and form of median lobe,

we add to the singiline assemblage of the Dromiina the following taxa that were included by

Csiki (1932: 1497- 1498) in the subtribe Cymindina: Metaxymorphus Chaudoir, 1850;

Periphobus Peringuey, 1896; and Callidomorphus Peringuey, 1896. Members of these taxa are

so similar to one another that it is inappropriate to rank them as genera. Nonetheless, adults of

each group are distinguished from one another on the basis of body form (see key, below).

Consequently, we rank each as a subgenus of Metaxymorphus, the senior name.

Notes are also included about Coptoptera, for reasons given below.

Metaxymorphus Chaudoir, SENSU NOVO.

Figs. 118A-B and 126A-B

Metaxymorphus (sensu stricto) Chaudoir, 1850: 370. GENERITYPE: Dromius frenatus Dejean, 1831: 351 (original

designation). Peringuey, 1896: 205.- Csiki, 1932: 1497.- Basilewsky, 1958a: 295.- 1961c: 216- 217.

Periphobus Peringuey, 1896: 204, 211. GENERITYPE: P. ferox Peringuey, 1896: 211 (monotypy).- Csiki, 1932:

1498.- Basilewsky, 1956: 236- 242.- 1958a: 296. NEW SYNONYMY.

Callidomorphus Peringuey, 1896: 204, 210. GENERITYPE: Metaxymorphus vittiger Chaudoir, 1877: 234

(monotypy).- Csiki, 1932: 1498.

We are not in position to give diagnostic features of adults of this genus, for we do not know

the other genera of dromiines well enough. We note, however, that the basis for assigning

Metaxymorphus to the Dromiina is: head without suborbital setigerous punctures; elytron with

penultimate umbilical setigerous puncture not laterad of antepenultimate and ultimate

punctures; scutellar interneur separate from interneur 1, base of latter present; tibiae average,

spinose; tarsomeres slender, glabrous dorsally, male front tarsomeres expanded slightly, with

biseriate adhesive vestiture ventrally; tarsal claws pectinate; median lobe of males with basal

bulb very small (Figs. 124A - 226A), right paramere very small (Fig. 125D); ovipositor with

stylomeres 1 and 2 subequal, both glabrous, stylomere 2 with preapical “orifice” (membranous

area. Figs. 121-123), preapical sensory furrow absent.

Description. — Smaller than average. Standardized Body Length ca. 3. 2-4. 5 mm. Form about average for

Carabidae. Color somber: uniformly rufous to testaceous, or elytra striped alternately rufo-testaceous and testaceous;

appendages paler than dorsum.

Microsculpture. Dorsum with meshes generally isodiametric to transverse on pronotum and elytra, microlines clearly

visible at 50X. Venter and lateral sclerites of thorax with meshes transverse.

Luster. Dorsal surface dull; ventral surface faintly iridescent.

Fixed setae. Average for lebiines. Head with two pairs of supraorbital setigerous punctures; submentum and mentum

each with single pair. Pronotum with two pairs of lateral setigerous punctures, posterior pair near posterior angles. Elytron

with two discal setigerous punctures in interval 3; umbilical series continuous, of 13 or 14 setigerous punctures. Legs with

average setation: tibiae with full complement of spines; tarsomere 5 with row of setae on each ventro-lateral margin.

Abdominal sternum VII of both males and females with four setigerous punctures.

Vestiture and surface. Dorsal and ventral surfaces essentially glabrous, impunctate. Antennomere 1 with single seta;

antennomeres 2 and 3 each with ring of setae preapically; remaining antennomeres average for lebiines.

Head. Average in form for lebiines, as broad or broader than average. Frontal impressions indistinct or well developed.

Clypeus transverse, about rectangular, or with anterior margin distinctly incised, concave. Eyes average. Antenna filiform,

antennomere 3 distinctly longer than 4; antennomeres each longer than wide.

Mouthparts. Labrum transverse, about rectangular. Mandibles (Figs. 118A-D) trigonal, but overall asymmetric, left

with anterior portion of terebra much narrower than anterior portion of right terebra (Figs. 1 18A and B). Left mandible

Quaest. Ent., 1983, 19 (1,2)

198

Ball and Hilchie

Figs. 118-123. SEM photographs of structures of Dromiina, genus Metaxymorphus.—¥'\%. 1 18: M. (Periphobus)

confusus (Basilewsky), mandibles— A and C, left, dorsal and ventral aspects, respectively; B and D, right, dorsal and

ventral aspects, respectively. Fig. 1 19: A/. (P.) confusus, right maxilla, ventral aspect. Fig. 120: M. (P.) confusus, tarsal

claw, terminal aspect. Figs. 121-123: Left stylomere 2. Fig. 121: Metaxymorphus (sensu stricto) species-A and B, lateral

and ventral aspect, respectively. Fig. 122: M. (P.) confusus-A and B, medial and apico-ventral aspects, respectively. Fig.

123: M. (Callidomorphus) vittiger (Peringuey)-A and B, lateral and ventral aspects, respectively. Scale bars: 1 18A-D,

and 1 19 = 100 ^ni; 120-123B= 50 itm. Legend, mandibles: art, anterior retinacular tooth; m, molar; pm, premolar; prt,

posterior retinacular tooth; tm, terebral margin; tt, terebral tooth; vg, ventral groove. Legend stylomere 2: m, preapical

membrane.

Cymindine Lebiini of Authors

199

Figs. 124-126. Line drawings of structures of Dromiina, genus Metaxymorphus.— Male genitalia, A and B, median lobe,

left lateral and ventral aspects, respectively; C and D, left and right parameres, respectively, ventral aspect. Fig. 124: M.

(sensu stricto) species. Figs. 125: M. (Periphobus) confusus (Basilewsky). Fig. 126: M. (Callidomorphus) vittiger

(Peringuey).

Quaest. Ent., 1983, 19 (1,2)

200

Ball and Hilchie

(Figs. 118A and C) with blunt, broad terebral tooth, terebral margin distinct for most of length of terebra; cutting edge

retinacular ridge, anterior retinacular tooth small; posterior retinacular tooth prominent, with well developed ridge

internally; molar tooth prominent, clearly isolated from premolar tooth; ventral groove (Fig. 118C) short, asetose. Right

mandible (Figs. 118B and D) with cutting edge terebral margin anteriorly, retinacular ridge posteriorly; terebral tooth

blunt, not as broad as that of left mandible; retinacular ridge prominent, anterior tooth conical, prominent, posterior

tooth well developed, with well developed ventral ridge; ventral groove short, setae few. Maxilla average in form; lacinia

with single row of setae on ventral surface (Fig. 119); galeomere 2 shorter than 1 coarsely sculptured; palpomeres

slender, palpomere 4 appreciably longer than 3, fusiform, narrowed apically. Labium average, mentum with well

developed tooth, and epilobes widened apically; glossal sclerite broad, apically with pair of long setae, and several

shorter setae; paraglossae adnate to glossal sclerite, about as long as latter, each with row of rather large setae apically;

palpomeres average in form, sparsely setose, palpomere 2 longer than 3, with two long setae; palpomere 3 fusiform,

narrowly truncate at apex.

Thorax. Pronotum markedly to slightly transverse, constricted posteriorly; all margins sharply beaded; anterior

margin slightly concave; posterior margin curved, but not lobate; sides rounded, incurved evenly posteriorly, not sinuate;

disc slightly convex, lateral grooves narrow; median longitudinal impression well developed. Prosternum with apex of

intercoxal process immarginate. Metathorax reduced, metepisternum either quadrate or wider than long (i.e., length of

anterior margin greater than that of lateral margin).

Legs. Average for lebiines. Tibiae with well developed spines. Front tarsomeres 1- 3 of males slightly expanded,

each with two rows of adhesive vestiture ventrally. Tarsomere 4 with apical margin truncate. Claws pectinate,

pectinations small (Fig. 120) few (one-three per claw).

Elytra. Average for lebiine adults, though humeri more sloped than average; apical margin subtruncate to truncate.

Interneurs shallow, impunctate; intervals flat to slightly convex; basal ridge sinuate, extended from humerus to edge of

scutellum.

Wings. Short stubs.

Abdominal sterna II- VII average for lebiines.

Male genitalia. Median lobe (Figs. 124A, B - 126A, B) cylindrical, anopic; basal bulb markedly reduced; apical

orifice on left side. Internal sac with various sclerites. Right paramere reduced (Fig. 125D).

Ovipositor and associated sclerites. Stylomeres 1 and 2 without ensiform or nematoid setae (Figs. 121-123),

subequal. Stylomere 2 with part of ventral surface membranous, membrane seemingly exsertile; without preapical

sensory furrow and associated sense organs of ventral surface.

Key to Subgenera of Metaxymorphus (sensu lato)

1 (0 ) Clypeus sloped ventrally rather abruptly, depressed medially, or not. Head

broad, body robust Periphobus (sensu lato) Peringuey.

r Clypeus sloped gradually anteriorly, surface plane, not depressed medially.

Head narrower, body slender, agonoid 2.

2 (T) Elytra bicolored, laterally testaceous, medially with more or less extensive,

irregular, rufo-testaceous to piceous dark mark

Subgenus Metaxymorphus Chaudoir. p. 197

2' Elytra bicolored, pattern regular, margin and intervals 1, 3, 5, and 7 testaceous,

intervals 2, 4, 6, and 8 rufo- piceous to piceous

Subgenus Callidomorphus Peringuey.

Notes about classification. — We explained above our reasons for including the species of

Metaxymorphus, Callidomorpha, and Periphobus in a single genus.

Csiki (1932: 1497- 1498) listed the names of 19 valid species of Metaxymorphus (sensu

strictoj, to which Basilewsky (1961c: 216- 217) added M. flaviceps Motschulsky, and M.

discopennis Motschulsky, having transferred them from Charopterus. Most of the species were

described by Peringuey. According to Basilewsky (1958a: 295), it is impossible to interpret

with certainty most of Peringuey’s descriptions. It will be necessary, therefore, to revise this

group, on the basis of a careful study of type material.

Basilewsky (1956: 236- 242) revised Periphobus Peringuey. Noting that the striking sexual

dimorphism recorded by Peringuey was the result of combining material of two species under a

single name, Basilewsky included the female co-type of P. ferox Peringuey (type locality-

Cymindine Lebiini of Authors

201

Oudtshoorn) in the new species P. confusus Basilewsky. He provided illustrations of habitus

(Fig. 1, P.ferox; Fig. 4, P. confusus) and of the male genitalia (Fig. 2a, P. confusus; Fig. 2b, P.

ferox) for both species. (As noted on reprints, captions for Figs. 3 and 4 were reversed). The

habitus illustration was reproduced as Fig. 39 in “South African Animal Life” (Basilewsky,

1958a: 296).

According to the description and key (Basilewsky, 1956: 238), heads of P.ferox specimens

are more markedly modified than are heads of P. confusus. Furthermore, the heads are

sexually dimorphic, especially those of P. ferox. However, this dimorphism is not as extreme as

Peringuey believed.

Notes about habitat. — We did not locate information for Metaxymorphus. We surmise,

however, on the basis of brachyptery, color, and form of adults, that they inhabit dry, open

area, and live on the ground.

Geographical distribution. — This genus is known only from localities in the Union of South

Africa.

Specimens examined. — We have seen 40 specimens of Metaxymorphus (sensu lato), from the following

localities in South Africa.

M. (Metaxymorphus) atriceps Peringuey. Male, Cape Colony, Uitenhaage, Rv. J. O. Neil 1917- 55 (BMNH). Male,

Cape Colony, Port Elizabeth G. A. K. Marshall 1917- 55 (BMNH).

M. (M.) cursor Peringuey. Male, female, Capetown, G. A. K. Marshall 1917-55 (BMNH).

M. (Metaxymorphus) species?- 12 males, 15 females, all from Cape Province. IV. 1958 E. S. Ross, R. E. Leech

(CAS). Male, two females, 19 mi. SE Garies 220 m V.2. 58; E. S. Ross, R. E. Leech (CAS). Four males, two females, 3

mi. SW Ladysmith 475 m. IV. 24. 58; E. S. Ross, R. E. Leech(CAS). Four males, female, Strandfontein XI. 13.49 B.

Malkin (CAS). Three males, nine females, Urendenburg XI.19.49 B. Malkin (CAS).

M. (Callidomorphus) vittiger Chaudoir. Male, Capland, Algoa Bay Dr. Brauns (BMNH). Female, Cape Colony

Uiteahage Rev. J. O. Neil 1917- 55 (BMNH).

M. (Periphobus) confusus Basilewsky. Four males. Cape Province 5 mi. W. Herold 600 m. IV. 24. 58; E. S. Ross, R. E.

Leech (CAS). Three females. Cape Province 3 mi. SW Ladysmith 475 m. IV. 24. 58; E. S. Ross, R. E. Leech (CAS).

Coptoptera Chaudoir

Coptoptera Chaudoir, 1837: 5. GENERITYPE: Coptoptera brunnea Chaudoir, 1837: 5 (monotypy).— Peringuey,

1896: 230 (in part).- Basilewsky, 1956: 401.

Klepsiphrus Peringuey, 1896: 223, 237. GENERITYPE: Klepsiphrus pugnax Peringuey, 1896: 237 (monotypy).

Syndetus Peringuey, 1896: 204, 222. GENERITYPE: Syndetus simplex Peringuey, 1896: 222

(monotypy).-Basilewsky, 1958b: 340-341.

Notes. — The genus Syndetus was included in the Cymindina by Peringuey (1896: 223),

though he pointed out that specimens of S. simplex had dromiine featues, as well. Basilewsky,

who examined the type of S. simplex, subsequent to his revision of Coptoptera (1956),

concluded that this species was not only a dromiine, but also that it was a species of

Coptoptera. We have not seen specimens of this species, but we accept Basilewsky’s judgement.

Tribe ZUPHIINI

The genus Agastus Schmidt-Goebel was included by Jedlicka (1963: 451) in the

Cymindina, but this genus clearly belongs in the Zuphiini- where Csiki (1932: 1567) placed it.

The senior author saw in the British Museum (Natural History) a specimen of A. ustulatus

Gestro from Java, and another with an indecipherable locality label, that was compared with

the type.

Quaest. Ent., 1983, 19 (1,2)

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Ball and Hilchie

CONCLUDING REMARKS

This paper began with the seemingly limited objective of seeking for the sister group of a

New World taxon of lebiines. It developed into a taxonomic treatment, based on barely

adequate material of groups ranging in rank from intra-specific to subtribal. Because so much

of the work centered around dismembering of a taxon treated previously as if it were a

taxonomically valid entity, and because of a shortage of time as well as of material for study,

the paper was frustrating to write. Taxonomists, like most other scientists, prefer to build,

rather than to take apart. Building for taxonomists consists primarily of description of new

taxa, and locating such in the system of previously described taxa. Nonetheless, re-organization

of groups like the cymindines of authors is required if future workers are to have a more secure

basis for proceeding with classification of the Lebiini.

We reiterate our belief that future progress will be along lines that Habu pioneered. We

wish to present briefly our views about how research on lebiines should proceed to produce

maximally useful results in minimum time. It seems to us that development of a general system

of classification for the Lebiini could be obtained in two stages. The first is undertaking of

regional studies, zoogeographical region by region. Publications could consist of

broad-spectrum reviews, based on dissections of representative members of each of the

described genera, in order to test further those characters that seem to be important, and to

assign these taxa to proper subtribes. At the same time, keys to genera ought to be written, and

species names catalogued.

Stage 2 would have a taxonomic focus, with all of the genera of the world of each subtribe

being assembled on the basis of inferred phylogenetic relationships. Persons doing this work

would have the data base assembled by regional studies to guide them. Additionally,

inter-regional comparisons would likely unearth additional character systems for use in

classification. At this stage, the search for sister groups both within and between tribes would

be of substantial importance and might lead to re-defining the limits of the Lebiini, either by

exclusion of some subtribes, or by inclusion of other lebiomorph tribes.

Because much taxonomic research is on a regional basis, we believe that the initial regional

approach advocated here to re-classification of lebiine genera will lead quickly to publications

that are of immediate interest and use. Such publications are likely to provide the impetus for

accumulation of additional data that will be of use in the world-wide treatment of genera of

individual subtribes.

A second general issue about which comments seem appropriate is ranking of taxa. So long

as one works within a geographically limited fauna, one can adopt the generic concepts that

have been applied by previous workers in that area. However, a study of a group on a

world-wide basis requires adoption of a uniform treatment. In this study, we were required to

deal with the discrepancy between a broad concept of genera advocated by Lindroth (1969b:

XVII) as applied to Holarctic carabids, and the more restricted one advocated explicitly by

Basilewsky (1968b: 185) in his studies of African taxa, and applied by Mateu in his studies of

tropical lebiines, generally. We believe that more broadly defined genera are more useful to

biologists other than taxonomists, and that units more difficult to recognize and more restricted

geographically can be named, but ranked at a lower level. Thus, we have defined genera

broadly, in spite of the discomfort that will be caused to some of our colleagues.

Procedure in ranking is not a matter of right and wrong, but one of taste and preference-

unless one adheres strictly to the tenets of cladistics. We hope that our re- ranking of

Cymindine Lebiini of Authors

203

well-known taxa will be judged on the merits advocated, and will be found satisfactory for

general use. We hope that our judgements will not be rejected out of hand.

Reference above to biologists other than taxonomists recalls the interrelationships between

these two groups, specifically with reference to the Lebiini. So far, study of lebiines has been

principally the playground of taxonomists. In the course of their studies, such workers have

discovered clues suggestive of modes of life and behavior that ought to excite interest of

ecologists and ethologists, as well as of economic entomologists. When such workers take up the

challenges inherent in determining life histories, host-parasite relationships, other ecological

relationships, and behavior patterns, the data produced will be of great value to taxonomists,

and will no doubt help in resolving vexing taxonomic problems.

Finally, we return to the initial purpose of this paper: a search for a sister group, specifically

that of Pinacodera Schaum. We think that we have found it, though we are not sure. At least

we have shed some light on the problem, and will develop hypotheses on the basis of our work.

Hennig (1966: 139) noted that an important task of phylogenetic systematics is search for sister

groups of monophyletic taxa. By accepting his formulation of tasks of systematics, we have

been able to examine a range of interesting problems. As others have stated, Hennig’s methods

seem fruitful. They should be used widely to seek understanding of important practical

taxonomic problems, rather than to serve as the basis for the futile and arid debate that rages in

current issues of “Systematic Zoology” and elsewhere, the tone of which is reminiscent of the

writings of Medieval scholastics addressing theological problems that seem now of little

consequence.

ACKNOWLEDGEMENTS

For the loan of material on which this study is based, we thank the following curators: M. E.

Bacchus and P. M. Hammond (BMNH); D. H. Kavanaugh (CAS); B. P. Moore (CSIRO); G.

Demoulin (IRSB); P. Basilewsky (MACT); R. J. McGinley and A. E. Newton (MCZ); J.-J.

Menier and H. Perrin (MNHP); V. Whitehead (SAMC); T. L. Erwin (USNM); and T. Sen

Gupta (ZSIC). Peter A. Meyer, Heidelberg, Victoria, Australia, presented to the Strickland

Museum various Australian lebiines that were very important to this study, and also provided

some of the inspiration for its undertaking.

For courtesies extended during visits to their respective institutions, the senior author is

grateful to: M. E. Bacchus and P. M. Hammond; P. Basilewsky; G. Demoulin; T. L. Erwin; D.

H. Kavanaugh; and R. J. McGinley and A. E. Newton.

James Liebherr (Department of Entomology, University of California, Berkeley, California)

made available to us a manuscript about affinities of lachnophorines that caused us to alter our

original view about this topic.

A preliminary draft of the typescript was reviewed by Terry L. Erwin, Ronald B. Madge

(CIE- BMNH), and Barry P. Moore, who made valuable suggestions for improving the

substance of the text, and who located numerous minor errors of omission and commission.

We received substantial assistance with preparation of illustrations from members of the

technical staff of our Department. We note especially contributions of D. Shpeley and G. B.

Braybrook, in preparing specimens and taking photographs with the Scanning Electron

Microscope. We also thank J. S. Scott for his meticulous attention to detail in preparing the

plates for publication. Various versions of the manuscript were typed by I. E. Bergum, and C.

Shirt, and we appreciate very much the care taken and the interest shown in producing the final

Quaest. Ent., 1983, 19 (1,2)

204

Ball and Hilchie

manuscript.

We also acknowledge assistance of J. E. O’Hara, who undertook a search of the literature

for current publications about cymindines.

NOTE ADDED IN PROOF

After this paper was nearly ready for publication, R. B. Madge {in litt.) advised us that

Habu (1982:113) had erected the Celaenephina as a new subtribe for Celaenephes

Schmidt-Goebel, though he expressed doubt {loc. cit.: 110) that this genus belonged in the

Lebiini. He also diagrammed {loc. cit.: 1 14, Fig. 29) his views about evolution of the stylomeres

of truncatipennian carabids, with those of Celaenephes either in an ancestral position, or

outside this taxonomic complex. Thus, our views, expressed above, are basically in agreement

with those of Habu.

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1871. Notes on a collection of insects from Gayndah. Ibid. 2: 79-158.

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212

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1920. The Carabidae of Tasmania. 45: 113-180.

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Cymindine Lebinii of Authors

213

INDEX TO NAMES OF TAXA

(Synonyms in italics)

FAMILY GROUP TAXA

Agrina, 112

Anomotarina, 110, 173

Apenina, 99, 112, 120, 124

Calleidina, 99, 110, 111, 112, 173

Calleiditae, 109, 116

Callidi, 110

Carabidae, 103

Catascopi, 109, 110

Catascopina, 110

Colliurini, 101

Coptoderitae, 109

Cymindidae, 101

Cymindina, 96, 98, 99, 108, 109, 1 10, 112,

129, 132, 133, 138, 158, 173, 197

Cymindini, 132, 158

Cyminditae, 109, 132

Demetriina, 110, 112

Dromii, 109, 110

Dromiina, 99, 110, 112, 196, 197

Dromiitae, 109, 196

Gallerucidiini, 1 1 1

Lachnophorini, 98, 101, 102, 103

Lebidii, 109, 111

Lebidiina, 110

Lebii, 109

Lebiidae, 109, 196

Lebiides, 101, 109, 110

Lebiina, 110, 116

Lebiinae, 158, 196

Lebiini, 96, 98, 101, 102, 108, 109, 110,

111, 116, 120, 196

Lebiitae, 109

Lichnasthenini, 99, 197

Lionychidae, 109, 196

Nemotarsina, 112

Pericalides, 109, 110

Pericalina, 98, 1 10, 1 1 1, 1 16, 197

Pericalitae, 109

Physoderi, 109

Physoderina, 110

Platynina, 98, 99

Pseudomasoreini, 132, 158

Pterostichini, 98, 99, 102

Singilini, 99, 196, 197

Somotrichini, 197

Thyreopteridae, 109, 116

Thyreopteritae, 109

Thysanotini, 98, 116

Zuphiini, 98, 99, 201

GENERA AND SUBGENERA

Abaditicus, new subgenus, 99, 186, 187,

189, 190, 191

Afrotarus Jeannel, 99, 139, 145, 146, 149,

154, 156

Agastus Schmidt-Goebel, 99, 201

Agra Fabricius, 108, 109, 111, 129, 191

Anarmosta Peringuey, 98, 99, 101

Anchonoderus Reiche, 102

Anomotarus (sensu stricto), 193, 196

Anomotarus Chaudoir, 173, 186, 192, 193,

196

Antimerina Alluaud, 98, 116

Apenes {sensu stricto), 125, 126

Apenes LeConte, 98, 99, 103, 124, 125,

126, 129, 132, 135

Apristus Chaudoir, 196

Asklepia Liebke, 101, 102, 103

Aspastus Peringuey, 171

Assadecma Basilewsky, 99, 129, 133, 158,

170

Assadera Mandl, 157

Assotatus Peringuey, 171

Assoterus Peringuey, 1 7 1

Astus Peringuey, 171

Callidomorphus Peringuey, 99, 197, 200

Calybe Castelnau, 101

Celaenephes Schmidt-Goebel, 112, 196

Cephalotarus Mateu, 173, 193

Ceylonitarus, new genus, 99, 129, 132,

135, 138, 145

Charopterus Motschulsky, 200

Coptoptera Chaudoir, 99, 197, 201

Cymindis {sensu stricto), 132, 133, 135,

149, 156, 157

214

Ball and Hilchie

Cymindis Latreille, 99, 109, 120, 127, 129,

132, 135, 138, 139, 145, 146, 149, 154,

156, 157

Cymindoidea (sensu stricto), 125, 126,

127, 128, 129, 133

Cymindoidea Castelnau, 99, 124, 126,

127, 128, 129

Diabaticus Bates, 99, 173, 186, 187, 188,

189, 190

Diaphoroncus Chaudoir, 172

Didymochaeta Chaudoir, 99, 125, 126

Dromiotes Jeannel, 173, 193

Dromius Bonelli, 109

Eucaerus (sensu stricto), 102, 107

Eucaerus LeConte, 98, 101, 102, 103, 107,

108

Eucheila Dejean, 109

Euplenes Darlington, 99

Euplynes Schmidt-Goebel, 98, 99, 101

Euproctinus Leng and Mutchler, 1 1 1

Eurycoleus Chaudoir, 1 1 1

Gallerucidia Chaudoir, 1 1 1

Habutarus, new subgenus, 99, 124, 125,

126, 127, 128, 129

Haplopeza Boheman, 101

Hystrichopus (sensu stricto), 120, 145,

171, 172

Hystrichopus Boheman, 99, 129, 132, 133,

135, 138, 145, 157, 158, 170, 171, 173

Inna Putzeys, 109

Iscariotes Reiche, 157

Klepsiphrus Peringuey, 201

Klepteromimus Peringuey, 158

Klepturus Peringuey, 158

Lachnaces Bates, 98, 101, 102, 103, 107,

108

Lachnophorus Dejean, 101

Lebia Latreille, 109

Lebidia Morawitz, 1 1 1

Leptosarcus Peringuey, 99, 116, 120

Leptotrachelus Latreille, 101

Lestianthus Sloane, 193

Lithostrotus Blackburn, 173, 192, 193

Madecassina Jeannel, 98, 116

Malisus Motschulsky, 99, 125, 126

Menas Motschulsky, 156

Metaxymorphus (sensu stricto), 197, 200

Metaxymorphus Chaudoir, 99, 197, 200,

201

Mormolyce Hagenbach, 109

Nemotarsus LeConte, 108

Neopsammoxenus Emetz, 157

Nominus Motschulsky, 125

Nototarus Chaudoir, 192, 193, 196

Notoxena Chaudoir, 186, 187

Paracymindis Jedlicka, 157

Periphobus Peringuey, 99, 197, 200

Phaedrusium Liebke, 98, 101, 102, 103

Philophuga Motschulsky, 110

Philotecnus Mannerheim, 126

Phloeocarabus Macleay, 99, 186, 187, 188

Phloeoxena Chaudoir, 1 20

Pinacodera Schaum, 96, 99, 129, 132, 135,

139, 145, 149, 203

Plagiopyga Boheman, 99, 129, 132, 133,

135, 145, 157, 158, 172, 173

Planesus Motschulsky, 139

Platytarus Fairmaire, 99, 124, 125, 126,

127, 129

Psammoxenus Chaudoir, 157

Pseudocymindis Habu, 157

Pseudomasoreus Desbrochers des Loges,

99, 129, 132, 133, 157, 158, 159, 164,

167, 168, 169, 170, 173

Pseudomastes Emetz, 157

Pteroritziella Mandl, 157

Selenorites Jeannel, 116

Selenoritus Alluaud, 98, 116, 117

Speotarus Moore, 173, 186, 187, 191, 192

Sphalera Chaudoir, 99, 124, 125, 126, 129

Sphenopalpus Blanchard, 125

Sphenopselaphus Gemminger and Harold,

125

Syndetus Peringuey, 99, 201

Taridius Chaudoir, 99, 129, 133, 135, 139,

145, 146, 149

Tarulus Bedel, 157

Tecnophilus Chaudoir, 1 10

Thyreopterinus Alluaud, 98, 116, 117, 118

Thyreopterus (sensu stricto), 118

Thyreopterus Dejean, 98, 116, 117

Thysanotus Chaudoir, 98

Cymindine Lebinii of Authors

215

Trigonothops (sensu strictoj, 186, 187,

188, 192

Trigonothops Macleay, 99, 147, 173, 186,

187, 188, 190, 191, 192

Trymosternus Chaudoir, 99, 121, 124,

126, 128, 129

Uvea Fauvel, 192

Xatis Fairmaire, 99

SPECIES AND SUBSPECIES

aenea Dejean, Cymindis, 125

alluaudi (Jeannel), Thysanotus, 116

andrewesi (van Emden), Cymindis, 146,

147, 148

andrewesi van Emden, Taridius, 147

angusticollis (Sloane), Anomotarus, 196

angusticollis Boheman, Hystrichopus, 157

arizonensis Horn, Cymindis, 157

atriceps Peringuey, Metaxymorphus, 201

australis (Erichson), Trigonothops, 188,

189, 190

australis Chaudoir, Nototarus, 192

australis Erichson, Diabaticus, 186, 189

australis Erichson, Plochionus, 186, 189

badestrinus Bates, Eucaerus, 108

basilewskyi, new species, Hystrichopus,

164, 168

birmanica (Bates), Cymindis, 146, 147,

148

birmanicus Bates, Taridius, 147

bisignata Dejean, Cymindoidea, 126

borealis LeConte, Cymindis, 1 57

brunnea Chaudoir, Coptoptera, 201

callidoides Chaudoir, Euplynes, 99, 101

canigoulensis Fairmaire and Laboulbene,

Cymindis, 157, 158

canigoulensis Fairmaire and Laboulbene,

Pseudomasoreus, 158, 159

capicola (Basilewsky), Hystrichopus, 159,

164, 167, 173

capicola Basilewsky, Pseudomasoreus, 167

ceylonicus, new species, Ceylonitarus, 135,

138, 139

coerulescens Blackburn, Anomotarus, 196

coerulescens Blackburn, Lithostrotus,

192, 193

collaris (Blackburn), Trigonothops, 188,

190, 191

collaris Blackburn, Diabaticus, 186, 189,

190

confusus Basilewsky, Metaxymorphus,

201

confusus Basilewsky, Periphobus, 201

cordatus Rambur, Trymosternus, 128

cursor Peringuey, Metaxymorphus, 201

cyanipennis Schmidt-Goebel, Euplynes, 99

cyclogonus Chaudoir, Hystrichopus, 173

cymindoides Peringuey, Hystrichopus, 173

discopennis Motschulsky,

Metaxymorphus, 200

dispar Peringuey, Anarmosta, 99, 101

dispar Peringuey, Euplynes, 101

dorsalis Thunberg, Hystrichopus, 172

elegans Alluad, Antimerina, 116

famini Dejean, Cymindis, 126

ferox Peringuey, Periphobus, 197, 200,

201

ferruginea Boheman, Plagiopyga, 157,

201

flaviceps Motschulsky, Metaxymorphus,

200

frenatus Dejean, Dromius, 197

fuscata Dejean, Cymindis, 139

geminatus Bates, Eucaerus, 108

haitianus Darlington, Eucaerus, 108

hamigera Chaudoir, Didymochaeta, 125

hessei Basilewsky, Leptosarcus, 1 20

hieronticus Reiche, Cymindis, 156, 157

hilaris Bates, Eucaerus, 108

humeralis Fourcroy, Buprestis, 139

insularis Darlington, Eucaerus, 108

jeanneli Mateu, Anomotarus, 193

kilimana Kolbe, Cymindis, 139, 154, 155,

156

kivuanus Basilewsky, Hystrichopus, 159,

164, 167, 168

kivuanus Basilewsky, Pseudomasoreus,

167

kivuanus Basilewsky, Thyreopterus, 118

lebioides Bates, Eucaerus, 108

216

Ball and Hilchie

leleupi (Basilewsky), Cymindis, 154, 155

limbata Dejean, Cymindis, 139, 149

longiplaga Chaudoir, Trigonothops, 191

lucidula Dejean, Cymindis, 125

lucifugus Moore, Speotarus, 186

lucifugus Moore, Trigonothops, 191

maculipennis Mateu, Cephalotarus, 193

madagascariensis (Basilewsky),

Hystrichopus, 170, 172

madagascariensis Basilewsky, Assadecma,

158

mastersi MacLeay, Phloeocarabus, 186

mateui, new species, Hystrichopus, 164,

169, 170

meruana (Basilewsky), Cymindis, 154

meyeri, new species, Trigonothops, 188,

189, 190, 191

nigra (Andrewes), Cymindis, 154, 155,

156

nigricollis MacLeay, Trigonothops, 186,

187, 188

nigripes Fairmaire, Xatis, 99

nilgirica (Andrewes), Cymindis, 146, 147,

148

nilgiricus Andrewes, Taridius, 147

nimbanus Basilewsky, Hystrichopus, 172

olisthopoides Bates, Eucaerus, 108

onychina Dejean, Cymindis, 128

onychinus (Dejean), Trymosternus, 128

opacicollis Bates, Eucaerus, 108

opacula (Chaudoir), Cymindis, 146, 147,

148

opaculus Chaudoir, Taridius, 139, 147

pacifica Erichson, Calleida, 186

papua (Darlington), Cymindoidea, 128

papua Darlington, Nototarus, 127

parallelus Blanchard, Sphenopalpus, 125

pauper (Blackburn), Trigonothops, 188,

189

pauper Blackburn, Diabaticus, 189

planior Blackburn, Lithostrotus, 196

platicollis Say, Cymindis, 139, 149

porrectus Peringuey, Leptosarcus, 1 20

postica Dejean, Cymindis, 125

princeps (Moore), Trigonothops, 192

ptolemaei Alluaud, Selenoritus, 1 16, 1 17

ptolemaei Alluaud, Thyreopterus, 1 18

pugnax Peringuey, Klepsiphrus, 201

pulchripennis Bates, Eucaerus, 108

punctigera (LeConte), Cymindis, 189

punctulata Dejean, Cymindis, 125

raffrayi Fairmaire, Cymindis, 154, 155,

156

reticulatus, new species, Hystrichopus,

159, 164

rufipes Dejean, Hystrichopus, 172

scotti Basilewsky, Cymindis, 154, 155

sculpturatus Sloane, Lestianthus, 196

sericatus, new species, Eucaerus, 107, 108

sericeus Bates, Eucaerus, 107, 108

sericeus Bates, Lachnaces, 107

simplex Peringuey, Syndetus, 201

sinuata (Say), Cymindis, 125

stevensi (Andrewes), Cymindis, 139, 146,

147, 148

stevensi Andrewes, Taridius, 147

stigma Mannerheim, Philotecnus, 126

stigmula Chaudoir, Anomotarus, 193

stigmula Chaudoir, Cymindis, 192

stigmula Fairmaire, Lebia, 193

strandi Liebke, Asklepia, 102, 103

striatus Bates, Eucaerus, 108

sulcatus Bates, Eucaerus, 108

suturalis Dejean, Cymindis, 156, 157

suturalis Liebke, Phaedrusium, 103

thoracicus, new species, Hystrichopus,

164, 167, 169, 170

titschacki Liebke, Phaedrusium, 103

truncatus Rambur, Trymosternus, 128

tumidiceps Blackburn, Anomotarus, 196

uluguruanus Basilewsky, Hystrichopus,

159, 164, 167, 168

uluguruanus Basilewsky, Pseudomasoreus,

168

ustulatus Gestro, Agastus, 201

varicornis LeConte, Eucaerus, 107

variegata Dejean, Cymindis, 125

vittiger Chaudoir, Metaxymorphus, 197,

201

Book Review

Malicky, H. 1983. Atlas of European Trichoptera/ Atlas der europaischen Kocherfliegen/

Atlas des Trichopteres d’Europe. Series Entomologica 24, x + 298 pp. Dr. W. Junk Publishers,

The Hague, Boston, London. ISBN 90-6193-134-7. Dfl. 175.00 ($US 76.00 approx.).