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QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department

of Entomology, University of Alberta, Edmonton, Alberta.

Volume 24 1988

CONTENTS

Liebherr- Redefinition of the Supertribe Odacanthitae, and Revision of

the West Indian Lachnophorini (Coleoptera: Carabidae) 1

Book Notice- Lindroth, Carl H. 1985-1986. The Carabidae of

Fennoscandia and Denmark 43

Commentary- Saether, O.A. On the Limitations of Parsimony in

Phylogenetic Analysis 45

Spanton- The Cicindela sylvatica Group: Geographical Variation and

Classification of the Nearctic Taxa, and Reconstructed Phylogeny and

Geographical History of the Species (Coleoptera: Cicindelidae) 5 1

Smetana- Revision of the Tribes Quediini and Atanygnathini. Part II. The

Himalayan Region (Coleoptera: Staphylinidae) 163

Book Review- Jamieson, B.G.M. 1987. The Ultrastructure and Phylogeny

of Insect Spermatozoa 465

Clark- Revision of the Weevil Genus Loncophorus Chevrolat

(Coleoptera: Curculionidae, Anthonominae) 465

Commentary- Ball, G.E. University Training of Systematic Entomologists 519

Williams- Classification, Phylogeny and Zoogeographic Studies of

Species of Sathon Mason (Hymenoptera: Braconidae) 529

Book Review- Fleissner, G. and Gerta Fleissner. 1988. Efferent Control

of Visual Sensitivity in Arthropod Eyes: with Emphasis on Circadian Rhythms ... 639

Index 641

ioiuaest

lones

Entomolosicae

JUN 1? 1988

i-ieRARltS

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

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NUMBER 1

WINTER 1988

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Issued May 1988

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department

of Entomology, University of Alberta, Edmonton, Alberta.

Volume 24 Number 1 1988

CONTENTS

Liebherr-Redefinition of the Supertribe Odacanthitae, and Revision of the

West Indian Lachnophorini (Coleoptera: Carabidae) 1

Book Notice-Lindroth, Carl 1985-1986. The Carabidae of

Eennoscandia and Denmark 43

Commentary: Saether-On the Limitations of Parsimony in Phylogenetic

Analysis

45

Spanton-The Cicindela sylvatica Group: Geographical Variation and

Classification of the Nearctic Taxa, and Reconstructed Phylogeny and

Geographical History of the Species (Coleoptera: Cicindelidae) 51

Figure 1. Male paratype of Anchonoderus darlingtoni, new species, type locality.

REDEFINITION OF THE SUPERTRIBE ODACANTHITAE, AND

REVISION OF THE WEST INDIAN LACHNOPHORINI

(COLEOPTERAiCARABIDAE)

James K. Liebherr

Department of Entomology

Comstock Hall

Cornell University

Ithaca, NY 14853-0999; USA.

Quaestiones Entomologicae

24:1-42 1988

ABSTRACT

Based on cladistic analysis of exemplar generic taxa using 57 morphological

characters of the adult stage, the carabid supertribe Odacanthitae is redefined to

include the tribes Odacanthini and Lachnophorini. Based on shared-derived states

of the female reproductive tract, these sister taxa are hypothesized to be derivatives

of platynine stock. Genera previously considered to comprise the tribe

Pentagonicini possess synapomorphies of Odacanthini, and the more restricted

synapomorphies previously used to distinguish Pentagonicini are shown to be

insufficient to justify recognition of the tribe in a cladistically based classification.

Pentagonicini is thus combined with Odacanthini.

The genera of Lachnophorini are defined using characters of the cladistic

analysis. The tribe comprises the genera Calophaena Klug, Asklepia Liebke,

Phaedrusium Liebke, Eucaerus LeConte, Anchonoderus Reiche, Euphorticus Horn,

Calybe Laporte, and Lachnophorus Dejean, and is restricted to the warmer areas of

the Neotropical and Nearctic regions. A key to genera is provided.

The West Indian lachnophorine fauna is revised, with keys to species, diagnoses,

illustrations, and distributional data provided. Anchonoderus darlingtoni, n.sp., is

described from Hispaniola and Jamaica, and A. jamaicensis, n.sp., is described

from Jamaica. Lectotypes are designated for Anchonoderus humeralis (Bates), n.

comb., A. subtilis Bates, and Lachnophorus leucopterus Chevrolat.

La supertribue des carabides Odacanthitae est redefinie, bas^e sur une analyse cladistique de genres

selectionnes, utilisant 57 characteres morphologiques du stade adulte afin d’y inclure les tribues

Odacanthini et Lachnophorini. Basee sur des charactires derives-communs du systime reproductif femelle,

nous emettons I’hypothise que ces taxons-soeurs sont derives d’un groupe platynine souche. Des genres

precedamment considerees comme faisant partie de la tribue Pentagonicini possedent des synapomorphies

de la tribue Odacanthini, et les synapomorphies les plus restrictives precedamment utilisees pour distinguer

Pentagonicini sont demontrees insuffisantes pour justifier la reconnaissance de cette tribue dans une

classification basees sur la cladistique. Pentagonicini est done combinee avec Odacanthini.

RESUME

2

Liebherr

Les generes Lachnophorini sont definis utilisant les characteres de I’ analyse cladistiqiie. La tribue

comprend les genres Calophaena King, Asklepia Liehke, Phaedrusium Liehke, Eucaerus LeConte,

Anchonoderus Reiche, Euphorticus Horn, Calybe Laporte, et Lachnophorus Dejean, et est restreinte aux

zones plus chaiides des regions neotropicales et nearctiques. Une clef d’ identification aux genres est

presentee.

La faune lachnophorine de I'lnde de I' Quest est revisee, et des clefs d’ identification d I'especes, de

meme que des descriptions diagnostiques, des illustrations et des informations sur la distributions sont

presentees. Anchonoderus darlingtoni, n.esp., est decrite pour L Hispaniola et la Jamaique, et A.

jamaicensis, n.esp., est decrite pour la Jamaique. Des lectotypes sont designes pour Anchonoderus

humeralis (Bates), n. comb., A. subtilis Bates, et Lachnophorus leucopterus Chevrolat.

INTRODUCTION

Both the composition and placement of the tribe Lachnophorini have proved

enigmatic to carabid systematists. I believe there are two major reasons for the

uncertainty regarding this group. First, the Lachnophorini are principally distributed

in the New World tropical and subtropical regions. The carabid fauna of these areas

has been extensively described {e.g., Bates, 1881-1884), but only recently has

attention turned to detailed investigation of the natural affinities of taxa within this

fauna. Secondly, some Lachnophorini exhibit a derived ant-like habitus. Past

classificatory attempts have focused on this suite of unique derivations, at the

expense of searching for shared-derived characters held with other taxa.

The initial purpose for this study was to revise the West Indian species of

Lachnophorini as part of a group effort to monograph the West Indian carabid fauna

(Erwin and Sims, 1984). So little was known of the affinities of Lachnophorini,

however, that I determined to provide a better definition of the tribe, and elucidate

its phylogenetic affinities. To this end, a cladistic analysis was performed, which

was based on characters of the adult stage of Lachnophorini. Representatives of 14

other carabid genera previously implicated as relatives to lachnophorines were also

studied. Based on this investigation, the Odacanthini and Lachnophorini proved to

possess synapomorphies in the female reproductive tract, leading me to propose the

Odacanthini and Lachnophorini as sister groups. Analysis of taxa within

Lachnophorini provided characters useful for diagnosing the genera of the tribe. The

resulting generic diagnoses are provisional, and it will require an extensive revision

of mainland species to confirm what is presented here. Nonetheless, based on the

characters examined, and their analysis using cladistic principles (i.e., grouping of

taxa by shared-derived similarity), the Lachnophorini is judged to be a natural

assemblage.

Taxonomic History

LeConte (1853) proposed Lachnophori to comprise the genera Lachnophorus

Dejean and Ega Laporte {=Calybe Laporte). The following year Lacordaire (1854)

proposed Anchonoderides for Anchonoderus Reiche, Lachnophorus, Calybe, Ega,

and three other genera currently placed elsewhere: Callistus Bonelli, Lasiocera

Odacanthitae and West Indian Lachnophorini

3

Dejean, and Camptotoma Reiche. Horn (1881), impressed by the many peculiar

characters of the genus Ega, proposed the monogeneric tribe Egini. The first concept

of Lachnophorini largely reflecting current ideas was that of Bates (1883), who

proposed Lachnophorinae for Anchonoderus, Lachnophorus, Euphorticus Horn,

Calyhe, and Ega.

The Lachnophorini was later dismembered by Liebke (1936, 1938), who placed

Calyhe in the Colliurini (=Odacanthini), separated from Lachnophorus. He also

included in Colliurini genera previously placed in Ctenodactylini and Platynini.

More recently, Lachnophorini has been recognized as a tribe distinct from

Odacanthini and Platynini (Reichardt, 1977, Liebherr, 1983) and as a component of

an expanded Pterostichini (Lrwin et ai, 1977).

Eucaerus LeConte was considered “an osculant form which is equally out of

place in any tribe here defined” (LeConte, 1861). Ball and Hilchie (1983), through

an investigation of taxa previously considered members of the subtribe Cymindina

of Lebiini, placed Eucaerus in Lachnophorini. They considered it part of a eucaerine

complex also comprising Asklepia Liebke and Phaedrusium Liebke. Reichardt

(1974) had previously noted the similarity in palpal structure between Asklepia and

Calyhe.

Laporte (1835) first proposed the Odacanthidae. LeConte (1861) included the

Casnoniae (genera Casnonia [= CoUiuris Dejean] and Leptotrachelus Latreille),

Lachnophori (genera Ega and Lachnophorus) and Lucaeri (genus Eucaerus) in his

interpretation of Odacanthini. Jeannel (1948) proposed Odacanthomorphi at

superfamilial rank, comprising his families Perigonidae, Peleciidae,

Lachnophoridae, Odacanthidae, and Ctenodactylidae. Kryzhanovskiy (1976) united

the tribes Odacanthini and Ctenodactylini in the supertribe Odacanthitae, whereas

Lachnophorini was joined with Perigonini and Omphreini in the Perigonitae.

MATERIALS AND METHODS

Taxonomic Material

Specimens used in this study were obtained from the following institutions

through the assistance of their respective curators;

AMNH American Museum of Natural History, New York, L.H.

Herman;

BMNH British Museum (Natural History), London, N.E. Stork, M.J.D.

Brendell;

CNC Canadian National Collections, Ottawa, A. Smetana;

CUIC Cornell University Insect Collection;

HDEO Hope Dept, of Entomology, Oxford, G.C. McGavin, D.S.

Smith;

lOJK Institute of Jamaica, Kingston, T.H. Parr;

IREC Institut de Recherches Entomologique de la Carafbe, P.

Quaest. Ent., 1988, 24 (1)

4

Liebherr

Chalumeau;

IZPW Institut Zoologique, Academic Polonaise des Sciences,

Warsaw, S.A. Slipinski;

MCZ Museum of Comparative Zoology, Harvard University,

Cambridge, MA, J.M. Carpenter, A.F. Newton, S.R. Shaw;

MNHP Museum National d’Histoire Naturelle, Paris, H. Perrin;

NMNH National Museum of Natural History, Smithsonian Institution,

Washington, D.C., T.L. Erwin;

TAMU Texas A. & M. University, College Station, H.R. Burke;

UASM University of Alberta Strickland Museum, Edmonton, G.E. Ball

and D. Shpeley;

UMMZ University of Michigan Museum of Zoology, Ann Arbor, B.M.

O’Connor;

RLDC Robert L. Davidson Collection, c/o Carnegie Museum of

Natural History, Pittsburgh, PA.

Taxonomic Methods

Microscopic techniques follow Liebherr (1986b). Where possible, types of West

Indian Lachnophorini have been examined. Lectotype designations are included

where appropriate. Complete listings are included for West Indian localities, with

extra-limital circum-Caribbean records summarized from museum findings and

publications.

Body length was measured as the distance from mandibular apex to elytral apex.

This total was the sum of head length from mandibular apex to cervical collar,

median pronotal length, plus elytral length from base of scutellum to elytral apex.

Scanning electron micrographs are used to illustrate certain aspects of these

beetles’ structure. Specimens were sonicated for 30 seconds in 95% ethanol,

mounted on double-sided scotch tape, and sputter coated for four minutes with

gold/palladium mixture. The micrographs were taken on an AMR lOOOA

miscroscope at 5kV.

Methods of Cladistic Analysis. — An initial set of taxa that included all West

Indian Lachnophorini, plus representatives of 14 other genera, was examined to

determine taxa to be included in the cladistic analysis (Fig. 2). This study set

included representatives of five genera hitherto included in Lachnophorini.

Fifty-seven unit characters of the adult stage were examined (Appendix 1).

Inspection of these data and published information on larvae determined the taxa to

be included in the cladistic analysis.

A number of the unit characters represent further derivation of a more

generalized derived state. Such pairs of more general and more derived

synapomorphies are characters 1 and 2, 3 and 4, 5 and 6, 11 and 12, 13 and 14, 19

and 20, 24 and 25, 26 and 27, 31 and 32, 34 and 35, 36 and 37, 45 and 46. The final

cladograms were scanned to determine that the changes to the most derived states

Odacanthitae and West Indian Lachnophorini

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provisional

in-group

unresolved

provisional out-groups

Figure 2. Five lachnophorine genera included in cladistic analysis, and 14 other genera provisionally allied

with Lachnophorini in earlier classifications.

occurred above changes to the intermediate states.

Character state polarity was determined using out-group comparison (Watrous

and Wheeler, 1981). The ancestral character state for the study set was determined

based on three preliminary taxonomic groupings (Fig. 3), and the polarity algorithm

derived from Farris (1970) by Maddison et al. (1984). Using this method, the

character states at the secondary out-group node (Fig. 3) were determined, these

states serving as the basal states for cladogram construction. For those instances in

which this method could not unambiguously assign ancestral character states,

additional species of Glyptolenus Bates and Platynus Bonelli were used as

unresolved secondary out-groups.

Quaest. Ent., 1988, 24 (1)

6

Liebherr

tf) <0

3 3

k. (Ok.

0> 3 0

Figure 3. Provisional lachnophorine in-group, genera considered close neighbors based on preliminary

investigations, and platynine genus Glyptolenus used as the out-group for the cladistic analysis. All taxa

possess synapomorphy of bipartite spermatheca.

Cladistic network building was accomplished using the Phylogeny Inference

Package (PHYLIP) Mix algorithm, version 2.0 of Felsenstein,' and the Phylogenetic

Analysis Using Parsimony program (PAUP) of Swofford (1984). Based on character

state polarities determined as above, the Mix Algorithm was run using: 1) Wagner

Parsimony (Farris, 1970) for all characters, 2) Camin-Sokal (1965) coding for all

characters, and 3) a variety of Wagner and Camin-Sokal coding for various

characters. PAUP was run with Glyptolenus chalybaeus Dejean specified as the

outgroup, hold trees = 5, swap = local, and the mulpars option employed. The PAUP

cladograms did not depend on prior character polarity assessment, and are simply

*J. Felsenstein, Dept, of Genetics, Univ. of Washington, Seattle, WA 98195

Odacanthitae and West Indian Lachnophorini

7

the shortest network rooted at the outgroup.

The assumption of irreversibility of character evolution implicit in Camin-Sokal

coding is not considered to represent the general case for character evolution. The

probabilities of gains or losses for the characters in this study are largely unknown,

and use of Wagner parsimony is preferred in most cases. Three characters, however,

were judged to be largely irreversible, leading to use of mixed coding in the

PHYLIP Mix algorithm. Characters 37-39, respectively, involve complete loss of

the oblongum cell, loss of the wedge cell, and loss of the 1st anal vein on the flight

wings. Simplification of wing venation is a trend generally observed during the

evolution of brachyptery (Darlington, 1936). In the taxa observed herein, vein and

cell loss appears correlated with smaller body size. It is assumed for this study that

loss of wing veins and cells is irreversible. This assumption is based on the many

examples of vein and wing loss observed throughout the Carabidae. Rederivation of

lost veins and cells is deemed possible, but because of the prevalence of brachyptery

and venational simplification, it is considered much less likely than repeated losses.

The shortest cladogram may be preferred based on adherence to the principle of

parsimony but other factors must also be considered. First, it is recognized that

characters of greater complexity most likely have more complex genetic and

developmental bases. While parsimony is an objective criterion useful for

comparing cladograms (Wheeler, 1986), a less parsimonious cladogram may

express a more likely sum of character transformations. This might occur if

reduction in changes of a conservative character is offset by additional changes in

several labile characters. I judge the utility of a character in indicating phylogenetic

affinity by the number of times the character changes state on the cladogram: i.e.,

the fewer state changes the better the character (Sober, 1986). In this analysis, I

present several more parsimonious cladograms, each based on specific assumptions

of character transformation probabilities, plus a less parsimonious but preferable

cladogram that requires fewer state changes in what are considered better characters.

The choice of a cladogram to use as a basis for classification is extremely

important if we believe our classification should respresent cladistic affinity. If a

single cladogram is clearly more parsimonious than all others, it should be adopted

as the basis for classification, with associated nomenclatural changes. But, if a

marginally more parsimonious solution portends substantial nomenclatural changes,

such as proposing new genera or dismemberment of currently recognized genera,

the consequences of adhering strictly to parsimony as an arbiter of taxonomy must

be carefully weighed. A major aim of this study is the presentation of

synapomorphies deemed likely to define monophyletic groups in the Lachnophorini.

Thus, I have chosen the preferred cladogram based on three criteria; 1 ) it is among

the more parsimonious solutions, if not the most parsimonious; 2) it does not entail

wholesale nomenclatural changes; 3) it includes synapomorphies for currently

recognized genera of Lachnophorini {i.e., lachnophorine genera are considered

monophyletic). This cladogram and attendant classification are profferred as

Quaest. Ent., 1988, 24 (1)

8

Liebherr

preliminary hypotheses to be tested when mainland taxa are analyzed.

CLADISTIC ANALYSIS OF ODACANTHITAE

Taxa Included in the Analysis

Of the initial survey of 14 genera with potential affinities to the Lachnophorini

(Fig. 2), five were excluded based on lack of synapomorphies.

The genus Pehgona Laporte was considered closely related to Lachnophorini by

Jeannel (1948). The spermatheca of Perigona (Fig. 4a) is unipartite, unlike the

bipartite spermatheca of Lachnophorini (Liebherr, 1983). The larva of Perigona

termitis Grasse and Jeannel (1941) lacks the segmented urogomphi observed in the

odacanthines Eudalia macleayi Bates and Scopodes simplex Blackburn (Moore,

1965), and the lachnophorine Calybe sallei Chevrolat (Liebherr, 1983). For these

reasons, Perigona is excluded from the analysis.

The ctenodactyline-hexagoniine taxa Hexagonia Kirby, Leptotrachelus Latreille,

Ctenodactyla Dejean, and Omphreoides Fairmaire, were also found to lack

synapomorphies uniting the rest of the taxa examined. The ctenodactyline female

reproductive tract is characterized by a tubular or sac-like spermatheca attaching

broadly near the junction of the common oviduct and bursa copulatrix (Figs. 4b,c).

The spermathecal gland duct is long and enters near the base of the spermatheca.

The male aedeagus is in the inverse position, on its left side in repose, unlike the

aedeagus of most Platynini, and all Odacanthini and Lachnophorini examined. Thus,

the ctenodactyline-hexagoniine taxa are not considered closely related to

Lachnophorini. The tubular spermatheca and multiple nematiform setae on the

apical gonocoxite (Figs. 4b, c) are states observed in Lebiini {e.g., Tecnophilus

Chaudoir, Dromius Bonelli) (unpubl. data).

The cladistic analysis includes 22 taxa (Table 1); nine species of West Indian

Lachnophorini, two non-West Indian lachnophorine species, eight species hitherto

considered Odacanthini, two species hitherto considered Pentagonicini, and

Glyptolenus chalybaeus of the Platynini.

All species included in the analysis possess female spermathecae that can be

derived from a ground-plan represented by the bipartite spermatheca of Glyptolenus

chalybaeus (Fig. 4d). Other Glyptolenus spp. possess unipartite tubular

spermathecae typical for Platynini (Liebherr, 1986a). The use of G. chalybaeus as

the outgroup for Odacanthitae is based on the assumption that the Odacanthitae is

derived from a platynine-like ancestor. Other as yet unexamined platynines may

prove more closely related than G. chalybaeus, but this species is the only platynine

known to me with a bipartite spermatheca. The two non-West Indian lachnophorine

taxa are included for different reasons. Calybe laetula LeConte is included as an

exemplar of that genus. Lachnophorus pilosus Dejean is the type species of that

genus, and its inclusion is used to represent prior subgeneric concepts of

Lachnophorus {e.g., subgenera Lachnophorus and Aretaonus of Liebke [1936]).

Odacanthitae and West Indian Lachnophorini

9

Figure 4. Female reproductive tracts, ventral view; A, Pehgona nigriceps Dejean; B, Leptotrachelus dorsalis

F.; C, Ctenodactyla drapiezi Gory; D, Glyptolenus chalybaeus Dejean, bc-bursa copulatrix, co-common

oviduct; gcl-basal gonocoxite; gcll-apical gonocoxite; sp-spermatheca; sg-spermathecal gland;

sab-spermathecal apical bulb; sbb-spermathecal basal bulb.

Quaest. Ent., 1988, 24 (1)

10

Liebherr

For the Odacanthini, single generic examplars are included, with the exception of

CoUiuris Dejean, where three species are included. Placement of these three

congeners in a monophyletic group is one means to judge resultant cladograms.

Two genera assignable to the Lachnophorini are not included; Asklepia Liebke

and Phaedrusium Liebke (Reichardt, 1974; Ball and Hilchie, 1983). Affinities of

these taxa are discussed below based on published information.

Character Polarity - Results

The character-state transformation series determined via out-group comparison

are presented in Appendix 1, with taxon X character-state data in Table 1. Using the

method of Maddison et al. (1984) with the 3-group statement of Fig. 3, 53

characters can have primitive and derived states unambiguously assigned. Primitive

states of characters 47, 53, 54, and 57 are ambiguous because the unresolved closest

out-groups all possess one state, and Glyptolemis possesses the other (Table 1). The

primitive state for these characters was assigned based on character states found

generally in other Glyptolemis and Platynus. Character 50 was coded the reverse of

that determined by out-group comparison within this data set. Throughout Platynini,

two lateral ensiform setae on the apical gonocoxite is the more generalized state.

Other Glyptolemis spp. possess only two such setae whereas G. chalybaeiis

possesses three, making the necessity to reverse the polarity of this character an

artifact of picking a single species as the secondary out-group.

The character-state transformation series were used with the Mix algorithm to

detemiine several cladograms. The preferred cladogram placed one of the closest

out-group taxa, Calophaena Klug, as the basal member of the in-group,

Lachnophorini. This change did not affect any of the prior character-state polarities.

On the contrary, inclusion of Calophaena in the in-group made polarity assessment

of characters 1, 3, and 5 more clear-cut, as it resulted in all closest out-groups and

the secondary out-group possessing the same character-state: 0 (Fig. 7, Table 1).

Cladistic Analysis - Results

If the character polarities of above are used to designate an hypothetical ancestor,

the Mix algorithm of PHYLIP determines the shortest cladogram to be one of 147

steps (Fig. 5). This cladogram results using Wagner parsimony coding for all

characters, and Camin-Sokal coding for all characters. If character polarities are not

used, and the cladogram is rooted at the outgroup Glyptolemis, the PAUP algorithm

finds 12 cladograms of 146 steps, all with the same topology (Fig. 6) but differing in

the placement of character state changes. This most parsimonious cladogram has

several undesirable attributes: 1) Odacanthini is not recognized as a monophyletic

group, and the overall structure is of sequential addition of one or two out-group

genera to the in-group; 2) the various representatives of CoUiuris are separated; 3)

this scheme necessitates evolution of the lachnophorine female reproductive tract

(Figs. 9c, d) by reversal from the highly derived female tract of Odacanthini (Figs.

Odacanthitae and West Indian Lachnophorini

11

Table 1. Character state x taxon data matrix for cladistic analysis of Odacanthitae;

0=primitive state, l=advanced state, ?=character state inapplicable or sex unknown.

Species

Glyptolenus chalybaeus Dejean

Calophaena bicincta De j . £. Bois.

Eucaerus insularis Darlington

Eucaerus haitianus Darlington

Anchonoderus j amaicensis , n. sp.

Anchonoderus humeralis Bates

Anchonoderus darlingtoni, n. sp.

Anchonoderus subtilis Bates

Anchonoderus subaeneus Reiche

Euphorticus pubescens Dejean

Calybe laetula LeConte

Lachnophorus leucopterus Chevrolat

Lachnophorus pilosus Dejean

Eudalia latipennis Macleay

Pentagonica picticornis Bates

Scopodes aeneus Macleay

Lachnothorax sp .

Dicraspeda brunnea Chaudoir

Odacantha melanura L.

Colliurus pensylvanicus L.

Colliuris sp.

Colliuris hubenthali Liebke

12345 67890

00000 00000

10101 01011

mil 11101

10001 01000

00000 01000

10101 01000

mil 11010

mil 11000

10111 11000

00000 00010

00000 00011

00000 00001

00000 01010

00000 00010

00000 01010

00000 00010

Character

mil 11112

12345 67890

00000 00000

10000 00110

11000 10000

11001 00000

10001 00000

11001 00000

11001 01100

11001 00100

10001 01000

11001 01100

00001 01000

00100 11000

00110 00000

11001 00010

00000 01010

00000 00110

00000 01111

11001 01111

00001 01011

22222 22223

12345 67890

00000 00000

00100 00001

00000 00000

00011 00000

00010 00010

00010 00000

00000 00100

01000 00000

01000 11010

01011 00010

00100 00000

10000 00000

10010 00000

00100 00000

00000 00000

00000 00001

01000 11010

01000 10010

44444 44445

12345 67890

?0000 00001

70000 01001

71000 01011

01000 01010

71100 01011

11100 01000

01100 01010

11100 01000

33333

12345

00000

00110

00100

11110

00111

11110

10010

00011

00111

00100

00110

00111

00010

00100

00110

00100

00111

00110

00100

33334

67890

00000

10111

mil

10100

00110

00100

11110

10110

11110

00000

11110

00000

01100

01101

01001

01011

01001

01011

01010

11010

01010

01111

01101

OHIO

55555 55

12345 67

00000 00

10111 01

07777 77

01101 01

Hill 11

11101 11

Hill 11

71110 11

01100 11

01100 10

01100 11

01100 10

00110 01

10110 01

00110 01

8, 9a). Objections 1 and 2 reflect taxonomic considerations, and the desire to

conserve the current nomenclature in the light of an insignificant (1 step or 0.68%)

gain in parsimony. The third objection is based on rejection of what is considered an

unlikely character transformation. Thus, the most parsimonious cladogram usefully

portraying taxonomic categories and phylogenetic affinity is considered to be the

147 step cladogram found by PHYLIP (Fig. 5).

When Camin-Sokal coding is used for the three wing cell and vein characters

(37, 38, 39), and Wagner coding is used for the balance, the result is a cladogram of

149 steps with the same topology as the 147 step cladogram (Fig. 5). The two extra

steps are necessitated by the required repeated derivations of cell and vein loss

(characters 37 and 39).

The preferred cladogram (Fig. 7) differs from the 147 and 149 step cladograms

by placing Eucaerus close to the base of the lachnophorine clade. With characters

37-39 coded as Camin-Sokal characters, the cladogram is 150 steps long, or one step

longer than the most parsimonious cladogram using such coding (Fig. 5). The more

basal placement of Eucaerus results in a single-step transformation series for

character 43: spermatheca with a gradually narrowing basal bulb (Figs. 9b, 12c, d)

evolving once to a basal bulb with apical filament (Figs. 9c, d, 13a, 14b, e, 15d).

The 147 and 149 step cladograms (Fig. 5) require an additional reversal to the

Quaest. Ent., 1988, 24 (1)

12

Liebherr

Glyptolenus chalybaeus

Eudalia latipennis

Pentagonica picticornis

Scopodes aeneus

Lachnothorax sp.

Colliuris hubenthali

Colliuris sp.

Colliuris pensylvanicus

Odacantha melanura

Dicraspeda brunnea

Calophaena bicincta

Anchonoderus darlingtoni

Anchonoderus jamaicensis

Eucaerus haitlanus

Eucaerus insularis

Euphorticus pubescens

Calybe laetula

Lachnophorus leucopterus

Lachnophorus pilosus

Anchonoderus humeralis

Anchonoderus subaeneus

Anchonoderus subtllis

Figure 5. Cladogram of both 147 steps determined using PHYLIP Wagner algorithm, and 149 steps

determined using PHYLIP Mix algorithm in which characters 37, 38, and 39 are coded using Camin-Sokal

Coding; i.e., irreversible.

primitive state for this character in the genus Eucaerus. Placement of Eucaerus

basal to Anchonoderus also results in one fewer step in characters 15, 29, 50, and

56. The presence (Figs. 13b, 16a, c) or absence (Fig. 12e) of a V-shaped sclerite on

the dorsum of the aedeagus (character 56) changes state only once on the preferred

cladogram, raising its value as an indicator of affinity under this hypothesis.

The preferred cladogram requires an extra step in characters 17 and 36, and two

extra steps in characters 49 and 54. Character 17, presence or absence of a pronotal

basal seta, changes five times on the most parsimonious cladogram (Fig. 6), and thus

Odacanthitae and West Indian Lachnophorini

13

Glyptolenus chaiybaeus

Eudalia latipennis

Pentagonica picticornis

Scopodes aeneus

Dicraspeda brunnea

Odacantha melanura

Colliuris pensylvanicus

Colliuris sp.

Colliuris hubenthali

Lachnothorax sp.

Calophaena bicincta

Anchonoderus subtilis

Anchonoderus subaeneus

Anchonoderus humeralis

Anchonoderus darlingtoni

Anchonoderus jamaicensis

Euphorticus pubescens

Eucaerus insularis

Eucaerus haitianus

Calybe laetula

Lachnophorus leucopterus

Lachnophorus pilosus

Figure 6. 146 step, most parsimonious cladogram determined using the PAUP algorithm, with Glyptolenus

the designated outgroup.

would appear a poor indicator of affinity. Character 36 is the reduction of the

oblongum cell from complete (Fig. 13c) to incomplete (Fig. 16e). This character too

is not expected to be a good indicator of phylogeny due to repeated reduction of

flight wings. The other two characters (49 and 54) change three and two times

respectively on the most parsimonious cladogram, but are considered poorer

indicators of affinity than spermathecal configuration.

The placement of Eucaerus adjacent to Calophaena results in recognition of

Anchonoderus as the monophyletic sister group to Euphorticus - Calybe -

Lachnophorus. If Eucaerus is moved basal to Anchonoderus with the other

lachnophorine relationships of Figure 5 retained, the resultant cladogram is 154

Quaest. Ent., 1988, 24 (1)

9,42,45,48,49,54

14

Liebherr

Glyptolenus chalybaeus

Calophaena bicincta

Eucaerus insularis

Eucaerus haitianus

Anchonoderus jamaicensis

Anchonoderus humeralis

Anchonoderus darlingtoni

Anchonoderus subtilis

Anchonoderus subaeneus

Euphorticus pubescens

Calybe laetula

Lachnophorus leucopterus

Lachnophorus pilosus

Eudalia latipennis

Pentagonica picticornis

Scopodes aeneus

Lachnothorax sp.

Dicraspeda brunnea

Odacantha melanura

Colliuris pensylvanicus

Colliuris sp.

Colliuris hubenthali

Figure 7. Preferred cladogram 150 steps in which Eucaerus is placed basal to Anchonoderus', character state

advances are shown by slashes, reversals to more primitive stage by X’s, character numbers correspond to

Appendix 1.

Odacanthitae and West Indian Lachnophorini

15

Figure 8. Female reproductive tracts, ventral view; A, Odacantha melanura L.; B, Eudalia latipennis

MacLeay; C, Pentagonica picticornis Bates; D, Scopodes aeneus MacLeay. sab-spermathecal apical bulb;

sbb-spermathecal basal bulb; sg-spermathecal gland.

Quaest. Ent., 1988, 24 (1)

16

Liebherr

Figure 9. Female reproductive tracts, ventral view; A, Colliuris huhenthali Liebke; B, Calophaena bicincta

Dejean and Boisduval; C, Calybe laetula LeConte; D, Lachnophorus pilosus Dejean, sab-spermathecal

apical bulb; sbb-spermathecal basal bulb; sg-spermathecal gland.

Odacanthitae and West Indian Lachnophorini

17

steps long. By the preferred 150 step cladogram (Fig. 7), the rounded gonocoxal

apices (character 51) of Anchonoderus have been derived once and have not

subsequently reversed to the ancestral state. Also, the setose body condition

(character 15) is hypothesized to have arisen once within Lachnophorini based on

this cladogram.

The taxa with setose body and setose maxillary palps {Euphorticus - Calybe -

Lachnophorus) constitute a third lineage of the Lachnophorini. Euphorticus

possesses moderately inflated apical maxillary palpomeres with sparse long setae

(Fig. 1 Ic), whereas Calybe and Lachnophorus possess more inflated, densely setose

apical palpomeres (Fig. lid). Development of inflated, densely setose palpomeres

has occurred in two lineages; once in Eucaerus (Fig. 11b) and again in Calybe and

Lachnophorus. The infrageneric variation in palpal shape exhibited between L.

pilosus and L. leucopterus suggests that this characteristic has evolved three or more

times. Anchonoderus spp. generally have fusiform palps (Fig. 11a), but those of A.

jamaicensis, are slightly inflated (Fig. 14f), suggesting that the apparently

generalized fusiform palpomere shape of most Anchonoderus spp. has evolved via

reversal from a moderately inflated palpomere (Fig. 7, character 3).

This analysis places Calophaena as the basal taxon in the lachnophorine clade.

Calophaena shares derived states of the palpi (characters 1,3,5,), antennal setation

(character 7), subgenal setation (character 1 1), and aedeagal sac spination (character

55) with other lachnophorines. It has previously been grouped with Odacanthini

(Bates, 1883, Liebke, 1938), and shares the general habitus of the genera Colliuris

and Odacantha F. But, Calophaena bicincta Dejean and Boisduval lacks the

distinctive U-shaped sclerite joining the spermatheca to the oviduct observed in

other Odacanthini (Figs. 8, 9a, b). The elongate prothorax observed in Calophaena

cannot characterize Odacanthini, as Eudalia latipennis MacLeay possesses a

quadrate prothorax. Thus, based on cladistic affinity, Calophaena is more closely

related to Lachnophorini than Odacanthini.

If one wished to classify Calophaena in Odacanthini, placing it at the base of the

odacanthine clade in a modification of Figure 7, the cladogram would be four steps

longer than the preferred cladogram. Additional derivations of characters 1, 3, 5, 7,

11, and 55 would be offset by saving one step each in characters 9 and 54. The six

characters requiring extra steps change two, two, two, three, three, and three times,

respectively, on the shorter preferred cladogram (Fig. 7). Characters 9 and 54 both

require three state changes on the modification of Figure 7, showing that the saving

in steps for these characters still leaves them as homoplasious as characters 7, 11,

and 55 on the preferred cladogram. Thus, there is no justification based on

parsimony, or character weighting to retain Calophaena in Odacanthini.

Pentagonica Schmidt-Goebel and Scopodes Erichson have previously been

recognized along with Actenonyx White as the tribe Pentagonicini. Bell (1985)

revised the West Indian species of Pentagonica. Based on cladistic analysis of adult

characters, the Pentagonicini must be combined with Odacanthini. Thus, even

Quaest. Ent., 1988, 24 (1)

18

Liebherr

though Pentagonica and Scopodes possess a characteristic pronotal shape (character

21), and fused gula and mentum (characters 13, 14), these synapomorphies occur in

a matrix of odacanthine synapomorphies that include spermathecal configuration

(characters 44, 45, 48; Figs. 8, 9a), and a derived aedeagal apex (character 54; Figs.

lOa-d). Moore (1965) noted the similarities of larvae of Scopodes simplex Blackburn

and Eudalia macleayi Bates. Moreover, the “pentagonicine” prothoracic shape is

approached in Eucaerus (Fig. 12a). Based on cladistic principles, there is no

justification for recognition of Pentagonicini.

Taxonomic Conclusions of Cladistic Analysis

Based on the preferred cladogram (Fig. 7), the supertribe Odacanthitae is

comprised of two tribes, the Odacanthini and Lachnophorini. The Odacanthini

includes those genera previously considered in the tribe (Jeannel, 1948; Darlington,

1968; Reichardt, 1977), plus those previously considered Pentagonicini

{Pentagonica, Scopodes, Actenonyx). The genus Calophaena is removed from the

Odacanthini and placed in Lachnophorini.

Lachnophorini is considered the sister group to Odacanthini. The major

synapomorphy grouping these two tribes is a bipartite spermatheca in the female.

Supporting synapomorphies of more erratic occurrence throughout the Platynini and

Lebiomorphi include segmented urogomphi and absence of a maxillary inner lobe in

the larval stages. ^

The Lachnophorini as presently constituted comprises eight genera: Calophaena,

Eucaerus, Anchonoderus, Euphorticus, Calybe, Lachnophorus, and the two genera

included by Ball and Hilchie (1983) in the eucaerine complex, Asklepia and

Phaedrusium. The tribe is distributed in the warmer regions of the Neotropical and

Nearctic regions.

TAXONOMIC TREATMENT

Tribe Lachnophorini LeConte.

Lachnophori LeConte, 1853: 370.

Anchonoderides Lacordaire, 1854: 373.

Eucaeri LeConte, 1861: 22.

Egini G. Horn, 1881: 152.

Tribal Diagnosis. — A group of disparate external appearance, as evidenced by

the numerous classificatory proposals above, but possessing the following

characteristics: (1) a patch of setae on the subgena ventrad eye (Fig. 11); (2)

antennomeres 2 and 3 setose over length (except in Phaedrusium)’, (3) body size

small, generally less than 8 mm in length; (4) elytra usually moderately truncate

apically; (5) apical palpomeres inflated and setose (Figs, llb-d), or if fusiform,

covered by very short setae (Fig. 11a); (6) spermatheca bipartite (Figs. 9b, c) or

derivable from a bipartite ground-plan (Fig. 16d).

Figure 10. A-F, Male aedeagus; A, Odacantha melanura L., median lobe dextro-ventral view; B, Colliuris

pensylvanicus L., median lobe and parameres, ventro-laevo view; C, Dicraspeda brunnea Chaudoir, median

lobe and parameres, ventro-laevo view; D, Scopodes aeneus MacLeay, median lobe and parameres,

ventro-laevo view; E, Calophaena bicincta Dejean and Boisduval, median lobe and parameres, ventro-laevo

view; F, Calophaena bicincta, median lobe, dextro-ventral view, inner sac partially everted. G, Colliuris sp.,

flight wing. lA, 2 A — 1st and 2nd anal veins; O - oblongum cell; w - wedge cell.

Way of Life. — Lachnophorines are riparian and can be found along stream banks

and on the shores of temporary and permanent ponds. Adults may be observed if

water is splashed on sandy shorelines. Some species, such as Euphorticus

pubescens, Lachnophorus leucopterus, and Calybe sallei are diurnal in activity.

Anchonoderus spp. are less often observed during daytime, except by splashing and

are most commonly taken at light at night.

Liebherr (1983) described the larval stages of Calybe {Ego) sallei. The larvae

and adults are active on the clay-sand surface during May in the southern United

Quaest. Ent., 1988,24(1)

20

Liebherr

Figure 11. SEM micrographs of lachnophorine mouthparts, lateral view; A, Anchonoderus subtilis Bates; B,

Eucaerus sp., Trinidad, Simla Biol. Sta. N Arima; C, Euphorticus pubescens Dejean; D, Lachnopliorus

leucopterus Chevrolat. Scale bar = 0. 1 mm.

States. The larvae possess long urogomphi with whitish tips which they move in

apparent imitation of antennae. Their movements are quick and erratic, much the

same as the adults.

Provisional Key to Genera of Lachnophorini

Based on the characters recorded in the West Indian taxa, in other mainland taxa

used in the cladistic analysis, and in publications, the following provisional key is

presented. The generic concepts are based on characters of the type species, and to

the degree that the herein recognized genera are natural, this key will be useful. I

have derived key characters for Asklepia and Phaedrusium from Reichardt (1974)

and Ball and Hilchie (1983).

1 Dorsal integument glabrous, except for macrosetae in standard

positions 2

T Pronotum, elytra, and head with a pelage of shorter secondary

setae in addition to standard macrosetae 5

2 (1) Pronotum elongate; neck constricted; tarsomeres 1-4 of all legs

broadly expanded laterally Calophaena Klug

Odacanthitae and West Indian Lachnophorini

21

2' Pronotum broad, wider than long; neck not strongly constricted;

tarsomeres of mid- and hindlegs not expanded 3

3 (2') Pronotum with base truncate, lacking median lobe; elytra with

striae effaced; body bicolored; microsculpture not evident at

lower magnifications (to 50X) Asklepia Liebke

3' Pronotum with basal median lobe (Figs. 12a, b); elytral striae

effaced or evident; body bicolored or concolorous;

microsculpture of elytra not evident, or strong transverse mesh,

or parallel lines 4

4 (3') Antennomeres 2-3 without vestiture of short setae, bearing long

preapical setae; male protarsomeres lacking ventral adhesive

setae Phaedrusium Liebke

4' Antennomeres 2-3 with vestiture of short setae as well as

longer preapical setae; male protarsomeres 2-3, or 2-4 with

ventral adhesive setae Eucaerus LeConte, p. 21

5 (F) Maxillary apical palpomeres fusiform or at most slightly

inflated, covered with very short setae and thus appearing

glabrous (50X); pronotum with well-developed lateral margin

(Figs. 1, 14a, d, 15a, c) Anchonoderus Reiche, p. 24

5' Maxillary apical palpomeres moderately to strongly inflated

with at least a sparse covering of long setae (Figs. 11c, d);

pronotum cylindrical with poorly developed lateral margin 6

6 (5') Maxillary apical palpomeres moderately inflated (Fig. 11c)

with sparse coating of longer setae Euphorticus Horn, p. 28

6' Maxillary apical palpomeres more strongly inflated (Fig. lid)

densely covered by setae at least in basal 0.50 7

7 (6') Elytra with secondary setation composed of stout black setae

over twice as long as fine yellow setae and with well-developed

transverse impression at basal 0.33 Calybe Laporte

T Elytra with secondary setation more concolorous, longer setae

rufous, shorter setae yellow; elytra with three impressions in

third interval that may be foveate, but transverse impression

absent Lachnophorus Dejean, p. 34

TAXONOMIC REVISION OF WEST INDIAN LACHNOPHORINI

Eucaerus LeConte

Eucaerus LeConte, 1853: 386.

Lachnaces Bates, 1872: 201.

Type species. — Of Eucaerus, E. varicornis LeConte (monotypy); of Lachnaces,

L. sericeus Bates (Ball and Hilchie, 1983).

Quaest. Ent., 1988, 24 (1)

22

Liebherr

Generic Diagnosis. — Body glabrous dorsally except for standard macrosetae;

apical palpomeres inflated and densely setose (Fig. lib); pronotum broad with

well-developed basal median lobe (Figs. 12a, b); antennae with apical four or five

segments pale, basal segments rufo-brunneus; female spermatheca bipartite, the

basal and apical lobes subequal, not bearing an apical filament (Figs. 12c, d).

Key to West Indian Species

1 Pronotum maximum width 1.5-1.6X median length; pronotal

basal bead angled between hind angles and basal median lobe

(Fig. 12a) E. haitianus Darlington

r Pronotum maximum width 1.35-1.45X median length; pronotal

basal bead nearly straight between hind angles and basal

median lobe (Fig. 12b) E. insularis Darlington

Eucaerus haitianus Darlington

Eucaerus haitianus Darlington 1935: 210.

Diagnostic Combination. — Pronotum broader than in E. insularis, hind angles

more strongly produced, median lobe more strongly developed (Fig. 12a); flight

wings either fully developed (Fig. 12f) or with reduced apex (Fig. 12g); length

3.3^. 1 mm.

Male Genitalia. — Aedeagal median lobe straight apically, apex blunt, basal bulb

open (Fig. 12e); aedeagal internal sac with apico-ventral patch of stout spines.

Female Reproductive Tract. — Gonocoxae with apical fringe of 6-8 setae on

penultimate segment, 2-3 lateral and one dorsal ensiform setae on apical segment

(Fig. 12c); spermatheca with basal bulb gradually tapering to an acuminate tip,

apical bulb broadly rounded with small apical nipple.

Type. — Holotype $ (MCZ No. 22035), Haiti, FOuest, Etang Lachaux, SW

Peninsula, under 1000 ft., 26-27-X-1934, P.J. Darlington, Jr.

Distributional Records. — A Hispaniolan endemic with the following non-type

records: Dominican republic: Samana: Sanchez, VII- 1938 (88, MCZ). HAITI: I’Ouest: Etang

Lachaux, SW Peninsula, <1(X)0' el., 26-27-IX-1934 (12, MCZ).

Eucaerus insularis Darlington

Eucaerus insularis Darlington 1934: 120.

Diagnostic Combination. — Pronotum quadrate with base constricted, hind

angles poorly developed, base evenly rounded posteriorly (Fig. 12b); flight wings

either developed (Fig. 12h) or a vestigial flap not reaching beyond metathorax;

length 3. 7-3. 9 mm.

Female Reproductive Tract. — Gonocoxae with apical fringe of 8-10 setae on

penultimate segment, 3^ lateral and one dorsal ensiform setae on apical segment

Odacanthitae and West Indian Lachnophorini

23

Figure 12. A, Eucaerus haitianus Darlington, pronotum; B, E. insularis Darlington, pronotum; C, E.

haitianus, female reproductive tract, ventral view; D, E. insularis, female reproductive tract, ventral view; E,

E. haitianus, aedeagal median lobe with everted inner sac, dextro-ventral view; F, E. haitianus, flight wing;

G, E. haitianus, micropterous flight wing; H, E. insularis, flight wing, sab-spermathecal apical bulb;

sbb-spermathecal basal bulb; sg-spermathecal gland; 2A - 2nd anal vein.

Quaest. Ent., 1988, 24 (1)

24

Liebherr

(Fig. 12d); spermatheca with basal bulb apically hooked, apical bulb globose.

Type. — Holotype $ (MCZ no. 19536), Cuba, Las Villas, Soledad, June 1929, P.J.

Darlington, Jr.

Distributional Records. — A Cuban endemic with the following non-type record:

Cuba: Las Villas: Soledad, 2-12-VIII-1934 (17, MCZ).

Anchonoderus Reiche

Anchonoderus Reiche, 1843:38.

Lachnophonis {Axylosius) Liebke, 1936:461 (NEW COMBINATION, NEW SYNONYMY).

Type species. — Of Anchonoderus, A. eximius Audouin by original designation;

of Axylosius, A. humeralis (Bates) by original designation.

Generic Diagnosis. — Body with dense pelage of secondary setae; apical

palpomeres fusiform, appearing glabrous but covered with very short setae (Fig.

11a), penultimate maxillary palpomere appearing glabrous or bearing sparse, long

setae (Fig. 14f); pronotum cordate, hind angles evident, setose, one lateral seta;

female spermatheca bipartite, basal lobe expanded basally and bearing an apical

filament, apical lobe globose and subequal or larger than basal lobe (Figs. 13a, 14b,

e, 15d); apical gonocoxites rounded at tips.

Key to West Indian Species

1 Testaceous maculae on 5th or 6th, to 8th or 9th elytral interval

behind humeri A. humeralis (Bates), p. 25

1' Elytrta concolorous piceous, , at most with sutural interval

slightly paler 2

2 (L) Apical maxillary palpomere moderately inflated, penultimate

palpomere with long setae (Fig. 14f); length 4. 3^.4 mm

A. jamaicensis, n.sp., p. 26

2' Apical maxillary palpomere fusiform, penultimate palpomere

with very short setae except for those in apical ring (Fig. 11a);

length 4.6-7. 6mm 3

3 (2') Dorsal elytral impressions of third interval evident, the anterior

impression foveate (Fig. 1); length 4. 6-5. 8 mm

A. darUngtoni, n.sp., p. 25

3' Dorsal elytral impressions inconspicuous, difficult to discern

amidst secondary elytral punctation; length 6. 3-7.6 mm 4

4 (3') Pronotum quadrate, hind angles well developed, protruding

(Fig. 15c) A. suhtilis Bates, p. 27

4' Pronotum cordate, hind angles more weakly developed, not

protruding (Fig. 15a) A. subaeneus Reiche, p. 27

Odacanthitae and West Indian Lachnophorini

25

Anchonoderus darUngtoni, n.sp.

Lachnophorus humeralis Darlington, 1953: 4.

Diagnosis. — Vertex of head shiny black, pronotum and elytra concolorous

piceous, elytra lacking maculae; pronotum with sides parallel before obtuse hind

angle (Fig. 1); dorsal elytral impressions evident, the anterior and posterior

impressions slightly foveate; size moderate among West Indian species, length

4.6-5. 8 mm.

Description. — Head. Moderately elongate, eyes convex; mandibles, labrum, and clypeal edges

rufous; maxillae and labrum testaceous; mentum tooth broad, about 0.50X length lateral lobes of mentum;

antennae with moderately robust segments, scape testaceous, segments 2-1 1 slightly darker. Prothorax.

Pronotum with base constricted; hind angles obtuse-angulate; basal seta situated before hind angle; lateral

margin parallel for at least short distance anterad hind seta; lateral pronotal seta nearly 0.75 distance toward

front angle; front angle narrowly rounded, moderately projecting; median longitudinal depression well

developed, expecially in basal 0.25, transverse wrinkles intersecting depression in basal 0.50; base of

pronotum with continuous marginal bead; very small triangular depression inside hind angles. Elytra. Sides

subparallel, humeri broad, apex subtruncate with subapical sinuation nearly obsolete; scutellum and basal

0.50 of sutural edge rufous, rest of elytra piceous; striae complete and smooth apically, more incomplete and

broken into series of fine punctures medio-basally; intervals moderately convex; 13-16 umbilicate setae in

or adjacent to eighth stria from humerus to subapical sinuation. Pterothorax and Abdomen. Metepistemum

1.67X as long as wide; flight wings (Fig. 13c) with oblongum cell complete, wedge cell and first anal vein

absent (compare to Fig. lOg); first visible abdominal segment with longitudinal wrinkles adjacent to

metacoxae, segments 3-6 densely setose. Legs. Femora testaceous with rufous apices, coxae and trochanters

rufous, tibiae with rufous longitudinal seta-bearing ridges and testaceous canaliculi, tarsomeres

rufo-testaceous; male protarsomeres moderately expanded laterally, segments 1-3 bearing four longitudinal

rows of adhesive setae. Microscidpture. Vertex of head with strong isodiametric microsculpture; pronotum

with transversely stretched isodiametric mesh; elytra with regular isodiametric mesh. Male Genitalia.

Median lobe of aedeagus nearly straight with pointed apex (Fig. 13b); internal sac bearing two patches of

spines apically; V-shaped sclerite present at base of internal sac. Female Reproductive Tract. Apical

gonocoxites bearing two lateral and one dorsal ensiform setae, their apices rounded (Fig. 13a); basal

gonocoxites with apical fringe of nine setae; spermatheca with basal bulb narrow, apical bulb much wider

than basal bulb.

Holotype: d, HAITI: Artibonite: Ennery, nr. 1000' el., 6-1 l-lX-1934, P.J. Darlington, Jr. (MCZ).

Allotype: ?, same data (MCZ).

Paratypes: DOMINICAN REPUBLIC: Puerto Plata: 25 km S by rd. of Puerto Plata, VI- 1938 (Id, MCZ).

HAITI: Artibonite: Ennery, nr. 1000' el., 6-11-IX-1934 (44dd-, 64??, MCZ; Id, 3??, BMNH). Nord; Cape

Haiten (Id, MCZ). FOuest; Carrefour, V-1908 (1?, BMNH); Port au Prince, R. Eroide, 3-X-1934 (Id,

MCZ). Sud; Camp Perrin, nr. 1000’ el. (3dd, 3??, MCZ); Massif de la Hotte, Desbarrier, 12-14-X-1934 (3dd,

MCZ), NE Foothills, 2-4000' el.,10-24-X-1934 (lOdd, 15??, MCZ). JAMAICA: St. Thomas: Com Puss Gap,

2060' el., 14-XI-1946, ex fallen tree (Id, lOJK)

Etymology. — The very large type series exemplifies the efforts Philip Darlington

made in procuring specimens from the field. The species is respectfully dedicated to

his memory.

Anchonoderus humeralis (Bates), n.comb.

Lachnophorus humeralis "QdXQS, 1883: 153; Csiki, 1931; 890.

Lachnophorus (Axylosius) humeralis, Liebke, 1936; 461.

Diagnostic Combination. — Placed in Anchonoderus based on fusiform and

glabrous apical palpomeres, setose pronotal hind angles (Fig. 14a); head and

Quaest. Ent., 1988, 24 (1)

26

Liebherr

pronotum shiny piceous with slight aeneous cast; elytra with testaceous maculae at

humeri, extending from 5th to 9th interval; dorsal elytral impressions evident,

foveate; legs testaceous with smoky femoral apices, tibial apices and tarsi dark;

length 4.6-5. 1 mm.

Male Genitalia. — Median lobe of aedeagus bluntly rounded apically, internal

sac bearing apical fringe of spines (Fig. 14c), sclerotized straps at base of internal

sac on dorsum of aedeagus (retracted position).

Female Reproductive Tract. — Apical gonocoxites with two lateral and one

dorsal ensiform setae, basal gonocoxite with apical fringe of 1 1-12 setae (Fig. 14b);

spermatheca with basal bulb columnar basally, more constricted at apical filament,

apical bulb large and globose.

Type. — LECTOTYPE d hereby designated: [Panama] Caldera, 1200 ft..

Champion; Lachnophorus humeralis Bates; B.C.A. Col. I. 1, Lachnophorus

humeralis Bates; Lectotype d, Lachnophorus humeralis Bates, By Erwin ’76

(BMNH).

Extralimital distribution. — Recorded from Guatemala and Panama (Bates,

1883). NICARAGUA: Chontales(l, BMNH).

West Indian Records. Grenada: Chamilly Est. (windward side)(12, BMNH). ST. LUCIA:

29-III-1936 (2, NMNH); Marisule, 1 l-IV-1978 (2, IREC).

Anchonoderus jamaicensis, n.sp.

Diagnosis. — Apical maxillary palpomere fusiform with acute tip (Fig. 14f),

appearing glabrous but with very short setae covering surface (125X); penultimate

maxillary palpomere setose; pronotum constricted with nearly obsolete, setose hind

angles (Fig. 14d); dorsal elytral impressions strongly foveate; upper body surface

shiny aeneo-piceous; length 4.7 mm.

Description. — Head. Vertex convex dorsally, eyes moderately convex; mandibles, labrum, clypeus

and apical palpomeres rufous; maxillae, labium, and basal palpomeres testaceous; median tooth of mentum

slender, acute, 0.6X length lateral lobes of mentum; antennae with basal four segments rufo-brunneus,

segments 5-11 darker; outer segments robust, width of segment 10 0.6X length. Prothorax. Pronotum with

narrow lateral margins, constricted base, and nearly obsolete hind angles (Fig. 14d); basal seta situated just

before obtuse-angulate hind angle; lateral margin slightly concave before hind angle; lateral pronotal seta set

0.7X distance from pronotal base to front angles; front angle angulate, not projecting forward; median

longitudinal depression fine; basal bead broad, evenly developed across pronotal base; latero-basal

depression nearly obsolete, only slight depression inside hind angle. Elytra. Humeri well developed, lateral

margin of elytra slightly constricted at about basal 0.33 of length; apex subtruncate, wider elytral margin

evident before nearly obsolete subapical sinuation; three strongly foveate dorsal depressions in third interval,

each fovea with an associated macroseta; elytral intervals convex, with surfaces roughened by secondary

setation; striae complete, though punctate and nearly evanescent near base; 14 umbilicate setae in or adjacent

to eighth stria from humerus to subapical sinuation. Pterothorax and Abdomen. Metepistemum elongate;

flight wings with oblongum cell complete, wedge cell and first anal vein absent (as in Fig. 13c); first two

visible abdominal segments nearly smooth, shiny, segments 3-6 setose. Legs. Femora, tibiae, and tarsi

piceous, coxal apices and trochanters rufo-testaceous on front two pairs, coxae concolorous with femora on

hind legs. Microsculpture. Vertex without visible microsculpture, very shiny; pronotum with very faint

transverse mesh obscured by punctations of secondary setation; elytra with weak isodiametric mesh, the

surface shiny. Female Reproductive Tract. Apical gonocoxite with two or three lateral and one dorsal

Odacanthitae and West Indian Lachnophorini

27

ensiform setae, basal gonocoxite with apical fringe of seven to eight setae (Fig. 14e); spermatheca with basal

bulb wide basally, strongly constricted at apical filament, outer bulb longer than basal bulb plus filament.

Holotype: ?, JAMAICA: St. Andrew: Kingston, 20-11-1936, Sta. 184, Chapin and Blackwelder (MCZ). The

type also bears the label: 25527, MCZ Paratype, jamaicensis.

Nomenciatural Note. — . This specimen was labelled a paratype and entered into

the MCZ type book as “25527, Lachnophorus jamaicensis Dari., 1 para-Carabidae.”

There is no evidence of a manuscript description of this species, nor is there any

evidence of a specimen labelled as holotype either in the MCZ or NMNH (T.L.

Erwin, pers. comm.). As such, I retain Darlington’s specific epithet, validly describe

the species, and place the taxon in Anchonoderus based on the generic diagnosis.

Anchonoderus subaeneus Reiche

Anchonoderus subaeneus Reiche, 1843: 40; Bates, 1883: 153; Csiki, 1931: 888.

Diagnostic Combination. — Head elongate, eyes strongly convex; pronotum

narrow (Fig. 15a), basal width 0.4X width at humeral angles; dorsal elytral

impressions obsolete, difficult to discern from punctations of secondary setation;

elytral striae with strong punctures in basal 0.50; 17-18 umbilicate punctures in or

adjacent to eighth stria anterad obsolete subapical sinuation; elytral apex rounded;

length 6. 8-7. 6 mm.

Male Genitalia. — Aedeagal median lobe straight, narrow, with a rounded knob

apically (Fig. 15b); internal sac globose, with fine spicules apically; V-shaped

sclerite on base of membranous dorsum of median lobe.

Type. — Type not found in MNHP. Bates’ determined material was examined in

BMNH, and this serves as basis for species interpretation.

Extralimital Distribution. — Recorded from Guatemala and Colombia (Bates,

1883). ECUADOR (5, BMNH). FRENCH GUIANA: Cayenne (1, BMNH). GUATEMALA: Guatemala

City (1, BMNH). PANAMA: San Feliz (1, BMNH).

West Indian Records. GUADELOUPE: Deux Mamelles, 24-11-1973 (4, IREC); Duclos (Htrs.), 22-V-1978

(LIREC).

Anchonoderus subtilis Bates

Anchonoderus subtilis Bates, 1871: 33; Bates, 1883: 153; Csiki, 1931; 889; Darlington, 1934; 98;

Darlington, 1941; 13.

Diagnostic Combination. — Head moderately elongate, eyes strongly convex;

pronotum broad (Fig. 15c), basal width >0.50X width at humeral angle; dorsal

elytral impressions obsolete, difficult to discern amidst secondary setation; elytral

striae smooth, weakly punctate in basal 0.50; 17-18 umbilicate punctures in or

adjacent to eighth stria anterad obsolete subapical sinuation; elytral apex angulate,

margin laterad apex straight; length 6.6-1. 1 mm.

Male Genitalia. — Median lobe of aedeagus straight, broadly rounded at apex

(Fig. 15e); internal sac with expansion just apicad apex of median lobe, two rows of

large spines on ventral surface about 0.50 distance to apex, and broad field of fine

Quaest. Ent., 1988, 24 (1)

28

Liebherr

spicules; V-shaped sclerite at base of membranous dorsum of median lobe.

Female Reproductive Tract. — Apical gonocoxite with two lateral and one dorsal

ensiform setae (Fig. 15d); basal gonocoxite with apical fringe of 11-12 setae;

spermatheca with small basal bulb bearing long apical filament, apical bulb larger

than basal.

Type. — LECTOTYPE ? hereby designated; Guatemala; Anchonod. subtilis

Bates; H.W. Bates, Biol. Cent. Amer.; LECTOTYPE (purple border);

LECTOTYPE, Anchonoderus subtilis Bates, det. George E. Ball, 1972 (MNHP).

Extralimital Distribution. — Cordova and Paro del Macho, Veracruz, Mexico,

and Guatemala (Bates, 1883), north to Nuevo Leon, vicinity of Linares (unpubl.

data).

West Indian Records. CUBA: Las Villas: Soledad, Cienfuegos, VI-1929 (1, MCZ), 2-12-VIII-1934

(29, MCZ), IV- 1936 (39, MCZ), V-1936 (2 MCZ); Trinidad Mtns., San Bias and vie., 9-V-1936 (3, MCZ).

Oriente: Aguadores, 6-VI-1936 (1, MCZ); Maisi, 17-VII-1936 (2, MCZ); coast below Pico Turquino,

26-30-V1-1936 (3, MCZ); Sierra Maestra, VI-1965 (1, MCZ); Upper Ovando R., 1-2000' el.,

17-20-VII-1936 (7, MCZ). Pinar del Rio: Rangel Mtns., -1500' el., 24-VIII-1936 (4, MCZ). JAMAICA:

Portland: Comfort Castle, Rio Grande R., 170 m el., riparian gravel, 7-9-V-1985 (2, UASM), 16-V-1985 (1,

UASM). St. Catherine: Rio Cobre 5 mi. above Spanishtown, 29-VIII-1934 (39, MCZ). St. Thomas: Bath

Fountain, 18-VII-1961 (l,IOJK).

Euphorticus G. Horn

Euphorticus Horn, 1881: 144.

Type species. — Laefmophorus pubescens Dejean by original designation.

Generic Diagnosis. — Apical maxillary palpomere fusiform, with sparse long

setae (Fig. 11c), penultimate palpomere setose; body with pelage of erect golden

secondary setae; pronotum lacking basal seta, one large lateral seta, and four to five

smaller lateral setae set in lateral margin, all setae stouter than secondary setation;

elytra lacking dorsal impressions associated with third interval; elytral striae

strongly punctate and discontinuous basally, smooth and complete apically.

Euphorticus pubescens (Dejean)

Lachnophorus pubescens Dejean, 1831: 30.

Euphorticus pubescens, Horn, 1881: 144; Bates, 1883: 156; Blatchley, 1910: 137; Csiki, 1931: 891;

Darlington, 1941: 13.

Lachnophorus niger Gory, 1833: 245.

Lachnophorus laevicollis Reiche 1843: 180.

Lachnophorus pubescens var. aeneolus Bates, 1883: 156.

Diagnostic Combination. — A shiny black ant-like lachnophorine, length 4. 1^.9

mm; antennomeres 2-4 rufo-testaceous, scape and apical segments darker with a

smoky cast; microsculpture of dorsum greatly reduced, surface shiny; leg bases

rufo-testaceous, femora piceous.

Male Genitalia. — Median lobe of aedeagus with short rounded apex, shaft

convex ventrally (Fig. 16a); internal sac globose, constricted basally, lacking spines

Odacanthitae and West Indian Lachnophorini

29

Figure 13. Anchonoderus darlingtoni, new species; A; female reproductive tract, ventral view; B, male

aedeagal median lobe with everted inner sac, dextro-ventral view; C, flight wing, sab-spermathecal apical

bulb; sbb-spermathecal basal bulb; sg-spermathecal gland; 2A - 2nd anal vein; O - oblongum cell;

v-vee-shaped sclerite of median lobe dorsal membrane.

or spicules, with two strap-like dorsal sclerites.

Female Reproductive Tract. — Apical gonocoxites pointed apically, bearing two

lateral and one dorsal ensiform setae (Fig. 16b); basal gonocoxite with apical fringe

of six setae; spermatheca with smaller basal bulb bearing apical filament, and longer

gradually widening outer bulb, which is bluntly rounded at apex.

Type. — I could not find Dejean specimens of this species in the Chaudoir

collection (MNHP). My interpretation of this widespread species is based on

specimens in North American collections, including those below.

Habits. — E. pubescens occurs on peaty shores of waterways in the southeastern

United States. At a pond 4.1 miles west of Bonifay, Washington Co., Florida, it was

taken with Calybe sallei and the fire ant, Solenopsis geminata (F.) (ant

determination by W.L. Brown, Jr.).

Extralimital Distribution. — Specimens examined from the southeastern United

States, eastern Mexico, Honduras, Colombia, and Trinidad (CUIC and MCZ

Quaest. Ent., 1988, 24 (1)

30

Liebherr

Figure 14. A, Anchonoderus humeralis Bates, pronotum; B, A. humeralis, female reproductive tract, ventral

view; C, A. humeralis, male aedeagal median lobe and parameres with everted inner sac, dextro- ventral

view; D, Anchonoderus jamaicensis, new species, pronotum; E, A. jamaicensis, female reproductive tract,

ventral view; F, A. jamaicensis, maxillary palp, ventral view, sab-spermathecal apical bulb;

sbb-spermathecal basal bulb; sg-spermathecal gland; v-vee-shaped sclerite of aedegal median lobe dorsal

membrane.

Odacanthitae and West Indian Lachnophorini

31

Figure 15. A, Anchonoderus subaeneus Reiche, pronotum; B, A. subaeneus, male aedeagal median lobe with

everted inner sac, dextro-ventral view; C, A. subtilis Bates, pronotum; D, A. subtilis, female reproductive

tract, ventral view; E, A. subtilis, male aedeagal median lobe with everted inner sac, dextro-ventral view,

sab-spermathecal apical bulb; sbb-spermathecal basal bulb; sg-spermathecal gland; v-vee-shaped sclerite of

aedeagal median lobe dorsal membrane.

Quaest. Ent., 1988, 24 (1)

32

Liebherr

Figure 16. A, Euphorticus pubescens Dejean, male aedeagal median lobe with everted inner sac,

dextro-ventral view; B, E. pubescens, female reproductive tract, ventral view; C, Lachnophorus leucopterus

Chevrolat, male aedeagal median lobe with everted inner sac, dextro-ventral view; D, L. leucopterus, female

reproductive tract, ventral view; E, L. leucopterus, flight wing, o - oblongum cell; sab-spermathecal apical

bulb; sbb-spermathecal basal bulb; sab-spermathecal gland; v-vee-shaped sclerite of aedeagal median lobe

dorsal membrane.

Odacanthitae and West Indian Lachnophorini

33

Quaest. Ent., 1988, 24 (1)

Figure 17. Distribution records of Euphorticus pubescens Dejean for which specimens have been examined.

34

Liebherr

records, Fig. 17). Reported from Guatemala, and the Amazon south to Santa

Catarina, Brazil (Bates, 1883), and Colombia and French Guiana (Csiki, 1931).

West Indian Records. CUBA: Camaguey: Baragua, lO-XI-1926 (2, MCZ), 16-X-1927 (1, MCZ).

Habana: Santiago de las Vegas, 2-VII-1930 (1, MCZ). Isla de Pinos: Nueva Gerona, 18-1-1967 (1, IZPW).

JAMAICA: Clarendon: nr. Kellits, Mason R. Field Sta., 700 m el., treading Typha marsh, 30-V-1985 (3,

UASM); Mason R., 16-11-1984 (3, lOJK). Portland: Hope Bay W Port Antonio, 3 m el., treading Typha

marsh, 18-V-1985 (1, UASM). St. Ann: Ocho Rios, 20-24-VIII-1934 (18, MCZ). St. Elizabeth: Frenchman

at Black R., O m el., Typha marsh, 2-VI-1985 (9, UASM); Maggotty, 7-III-1976 (1, lOJK).

Lachnophorus Dejean

Lachnophorus Dejean, 1831: 28.

Lachnophorus (Aretaonus) Liebke, 1936: 461.

Type Species. — Of Lachnophorus, L. pilosus Dejean by Chenu (1851); of L.

{Aretaonus), L. elegantulus Mannerheim by original designation.

Nomenclatural Note. — Csiki (1931) places Stigmaphorus Motschulsky as a

junior synonym of Lachnophorus. The description of Stigmaphorus tesselatus

Motschulsky (1864) directly follows a key to Lachnophorus species, but based on

Motschulsky ’s (1862) diagnosis of Stigmaphorus, this genus should not be

considered the same as Lachnophorus.

Generic Diagnosis. — Apical maxillary palpomere fusiform to strongly inflated,

subulate, densely setose basally to entirely setose (Fig. 1 Id), penultimate palpomere

setose; body with secondary setation of two setal types - abundant anteriorly

recurved setae, and intermittent erect setae; pronotum lacking basal seta, bearing

one lateral seta; elytral dorsal impressions of third interval foveate, associated setae

indiscernible from secondary setation; elytral striae continuous, punctate basally,

smooth apically.

Lachnophorus leucopterus Chevrolat

Lachnophorus leucopterus Chevrolat , 1863: 198; Csiki, 1931: 890; Darlington, 1934: 98; Darlington, 1941:

13.

Lachnophorus (Aretaonus) leucopterus, Liebke, 1936: 463.

Diagnostic Combination. — Vertex of head with longitudinal wrinkles, area

between wrinkles with strong isodiametric microsculpture; scape testaceous

ventrally, smoky metallic green dorsally, antennomeres 2-A rufo-testaceous, outer

segments piceous; head and pronotum metallic cupreous-green; elytra pale creamy

white with white humeral macula, white discal macula at basal 0.33 on 6th interval,

and two oblique sequences of white maculae extending from 0.66 length at margin,

to 0.90 length at suture, dorsal elytral impressions at lateral umbilicate punctures

darker; length 4. 1-4.9 mm.

Male Genitalia. — Median lobe of aedeagus bluntly rounded apically (Fig. 16c),

internal sac globose, not spinose; two strap-like sclerites at base of dorsal

membranous area.

Odacanthitae and West Indian Lachnophorini

35

Female Reproductive Tract. — Apical gonocoxite pointed apically, bearing two

lateral and one dorsal ensiform setae (Fig. 16d); basal gonocoxite with apical fringe

of seven to nine setae; spermatheca with a sclerotized base bearing the gland duct,

and a single bulb that possesses a short broad apical filament.

Type. — Lectotype 9 hereby designated: [Cuba] Sto. Dgo., Salle; Lachnophorus,

leucopterus, Chev. 1817; Chevrolat, Carabidae, Fr. V.d. Poll, Pres. 1909, E., B.P.

Boulton; Type Col: 150 1/2, Lachnophorus, leucopterus Chevr., Hope Dept. Oxford;

Lectotype, Lachnophorus, leucopterus, Chevrolat, J.K. Liebherr 1987.

Habits. — L. leucopterus is found along the gravel shores of creeks. At Rio Cain

near San German, Puerto Rico, it occurred on bare gravel banks after recent

flooding, where it was the only carabid species observed. On the Rio Guanajibo near

Sabana Grande, Puerto Rico, it was taken along with beetles of the genus

Bemhidion. It is active in bright sunlight.

Distributional Records. — Endemic to the West Indies, with the following

records.

CUBA: Las Villas: Soledad, Cienfuegos, Arimao R„ 19-X-1926 (12, MCZ), 31-X-1926 (3, MCZ),

3-XI1-1926 (4, MCZ), 13-XII-1926 (2, MCZ), VI-1929 (1, MCZ), 2-12-VIII-1934 (10, MCZ), IV-1936 (3,

MCZ); Trinidad Mtns., 600-2000’ el., VI-1929 (2, MCZ), Hanabanillo Falls, 30-IV-1936 (1, MCZ). Oriente:

Aguadores, 6-VI-1936 (2, MCZ); Baracoa, 12-VII-1936 (4, MCZ); Cauto el Cristo (Cauto R.), 12-VIII-1936

(4, MCZ); Daiquiri, 30-V-1936 (3, MCZ); Hongolosongo, 7-VII-1936 (1, MCZ); Imias, 24-VII-1936 (5,

MCZ); Maisi, 17-VII-1936 (1, MCZ); Marcata, Rio Bayamo, 1913 (5, MCZ); Upper Ovando R., 1-2000' el.,

17- 20-VII-1936 (1, MCZ). Pinar del Rio: San Diego, 18-V-1931 (3, MCZ); Sierra de Rangel, XII- 1933 (4,

MCZ).

DOMINICAN REPUBLIC: Distrito Nacional: San Cristobal, 18-IX-1938 (1, MCZ), 5-IX-1966 (15,

TAMU); Villa Altagracia, VII-1938 (1, MCZ). La Vega: Constanza to Jarabacoa, 2^000' el., VIII-1938 (4,

MCZ). Puerto Plata: 25 km by road S Puerto Plata, VI- 1938 (7, MCZ). Santiago: San Jose de las Matas,

1000’-2000' el., VI-1938 (3, MCZ); S of Santiago, fthills Cord. Cent., VI-1938 (2, MCZ).

HAITI: Artibonite: Ennery, nr. 1000' el., 6-1 l-IX-1934 (23, MCZ). He de la Gonave: La Source, 15-11-1983

(7, RLDC). rOuest: Port au Prince, R. Eroide, 3-X-1934 (2, MCZ). Sud: Camp Perrin, 1-I1I-1983 (14,

RLDC), nr. 1000' el., 8-27-X-1934 (2, MCZ), Grand R. du Sud, 20-11-1983 (51,RLDC); Massif de la Hotte

NE, fthills 2^000' el., 10-24-X-1934 (9, MCZ).

JAMAICA: Wag Water, 6-III-1958 (3, BMNH). Clarendon: Rock R. 2 mi. NE Chapelton, 23-XII-1946 (2,

lOJK). Portland: Blue Mtns., nr. 4500'el., 13-20-VIII-1934 (27, MCZ); Comfort Castle, Rio Grande, 170 m

el., riparian gravel, 7-9-V-1985 (1, UASM), 16-V-1985 (2, UASM); nr. Windsor, Rio Grande, 40 m el.,

18- V-1985 (7, UASM). St. Andrew: Castleton Bot. Gard. (2, MCZ); Hope R., 26-V-1908 (28, BMNH), nr.

Gordontown, 213 m el., 7-XI1-1985 (9, UASM); Kingston, 14-11-1928 (4, MCZ), 26-IV-1937 (1, UMMZ);

St. Peter’s Yallah’s R., 850 m el., gravel, 21-V-1985 (21, UASM). St. Catherine: Rio Cobre 5 mi. above

Spanishtown, 29-VIII-1934 (3, MCZ); Spanishtown, 3-II (2, MCZ); nr. Zion Hill along Rio Pedro,

21-111-1954(1, CNC).

PUERTO RICO: Caguas, 28-29-V-1915 (7, AMNH); Coamo Spgs., Il-II (9, AMNH), 17-19-VII-1914 (13,

AMNH); Carite St. For. nr. Guavate, 650 m el., l-V-1985 (7, CUIC); Las Vegas, l-VI-1938 (2, MCZ);

Maricao, 30-V-1938 (3, MCZ); Mayaguez, 21-23-VI-1915 (2, AMNH), 28-11-1931 (3, MCZ), 19-1-1936

(20, MCZ), 27-III-1936 (7, MCZ); Rio Cain at Hwys. 2 and 360, 3-V-1985 (12, CUIC); Rio Guanajibo at

Sabana Grande, 1(X) m el., 4-V-1985 (33, CUIC); San German, XII-1933 (1, MCZ); Vinoles, Rio S. Vicente,

15-111-1937(1, MCZ).

Quaest. Ent., 1988, 24 (1)

36

Liebherr

ACKNOWLEDGMENTS

The research collection amassed by Philip Darlington formed the basis for the

cladistic analysis and species revision. I thank the Museum of Comparative Zoology

and James M. Carpenter for access to that material. Additional specimens used in

the analysis were provided by Charles Griswold and Jan Washburn. The Dean’s

Travelling Fellowship of the College of Agriculture and Life Sciences, Cornell

University, provided travel support for visits to European collections. I thank Rick

Hoebeke for taking the SEM micrographs, and Frances L. Fawcett for the

frontispiece. Quentin Wheeler graciously offered his computing facilities for the

PAUP analysis. Velvet Saunders typed the manuscript. Steve Nichols and Quentin

Wheeler provided critical reviews of the manuscript. This research was supported by

Hatch project NY(C) 139406.

REFERENCES

Ball, G.E. and G.J. Hilchie. 1983. Cymindine Lebiini of authors: redefinition and

reclassification of genera (ColeopteraiCarabidae). Quaestiones Entomologicae,

19: 93-216.

Bates, H.W. 1871. Notes on Carabidae, and descriptions of new species. (No. 5).

Entomologist’s Monthly Magazine, 8: 29-34.

Bates, H.W. 1872. Notes on Carabidae, and descriptions of new species (No. 12).

Ibid, 8 : 199-202.

Bates, H.W. 1881. Biologia Central! -Americana, Coleoptera, Carabidae, 1(1):

19-40.

Bates, H.W. 1882. Ibid: 41-152.

Bates, H.W. 1883. Ibid: 153-256.

Bates, H.W. 1884. Ibid: 257-296.

Bell, R.T. 1985. Pentagonica of the West Indies (Coleoptera: Carabidae).

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Blatchley, W.S. 1910. The Coleoptera of Indiana. Bulletin of the Indiana

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Camin, J.H. and R.R. Sokal. 1965. A method for deducing branching sequences in

phylogeny. Evolution, 19: 311-326.

Chenu, J.C. 1851. Encyclopedie d’Histoire Naturelle, 1: 1-312.

Chevrolat, A. 1863. Coleopteres de File de Cuba (1). notes, synonymies et

descriptions d’especes nouvelles. Annales de la Societe Entomologique de

France (Ser. 4), 3: 183-210.

Csiki, E. 1931. Coleopterorum Catalogus (Carabidae, Harpalinae V), 115:

739-1022.

Darlington, P.J., Jr. 1934. New West Indian Carabidae, with a list of the Cuban

species. Psyche, 41: 66-131.

Odacanthitae and West Indian Lachnophorini

37

Darlington, P.J., Jr. 1935. West Indian Carabidae II: itinerary of 1934; forests of

Haiti; new species; and a new key to Colpodes. Ibid, 42: 167-215.

Darlington, P.J., Jr. 1936. Variation and atrophy of flying wings of some carabid

beetles. Annals of the Entomological Society of America, 29: 136-176 + 3 pis.

Darlington, P.J., Jr. 1941. West Indian Carabidae VI. the Jamaican species and their

wings. Psyche, 48: 10-15.

Darlington, P.J., Jr. 1953. West Indian Carabidae IX. more about the Jamaican

species. Occasional Papers of the Museum of the Institute of Jamaica No. 8: 14

pp.

Darlington, P.J., Jr. 1968. The carabid beetles of New Guinea part III. Harpalinae

(continued): Perigonini to Pseudomorphini. Bulletin of the Museum of

Comparative Zoology, 137: 1-253.

Dejean, P.F.M.A. 1831. Species general des Coleopteres, 5: 1-883.

Erwin, T.L., D.R. Whitehead, and G.E. Ball. 1977. Checklist of the beetles of

Canada, United States, Mexico, Central America, and the West Indies, Family 4.

Carabidae, the Ground Beetles. World Digest Publ., Kinderhook, NY. 68 pp.

Erwin, T.L., and L.L. Sims. 1984. Carabid beetles of the West Indies

(Insecta:Coleoptera): a synopsis of the genera, and checklists of the tribes of

Caraboidea and of the West Indian species. Quaestiones Entomologicae, 20:

351^66.

Farris, J.S. 1970. Methods for computing Wagner trees. Systematic Zoology, 19:

83-92.

Gory, H.L. 1833. Centurie de carabiques nouveaux. Annales de la Societe

Entomologique de France, 2: 168-247.

Grasse, P.P., and R. Jeannel. 1941. Un carabique termitophile nouveau de TAfrique

tropicale. Revue Fran9aise d’Entomologie, 8: 135-146.

Horn, G.H. 1881. On the genera of Carabidae with special reference to the fauna of

boreal America. Transactions of the American Entomological Society, 9:

91-196.

Jeannel, R. 1948. Coleopteres carabiques de la region Malgache (part 2). Faune de

I’Empire Fran^aise, 10: 373-765.

Kryzhanovskiy, O.L. 1976. An attempt at a revised classification of the Carabidae

(Coleoptera). Entomological Review, 55(1): 56-64. (Transl. from Russian).

Lacordaire, T. 1854. Histoire Naturelle des Insectes. Genera des Coleopteres. Vol. 1,

486 pp. Roret, Paris.

Laporte, F.L. de. 1835. Etudes Entomologiques, Part I. Mequignon-Marvis, Paris.

94 pp.

LeConte, J.L. 1853. Notes on the classification of Carabidae of the United States.

Transactions of the American Philosophical Society, 10: 363^03.

LeConte, J.L. 1861. Classification of the Coleoptera of North America, Part 1.

Smithsonian Miscellaneous Collections, 3: 1-208.

Liebherr, J.K. 1983. Larval description of Calybe (Ega) sallei (Chevrolat) with a

Quaest. Ent., 1988, 24 (1)

38

Liebherr

preliminary assessment of lachnophorine affinities (CarabidaeiLachnophorini).

Coleopterists Bulletin, 37: 254-260.

Liebherr, J.K. 1986a. Cladistic analysis of North American Platynini and revision of

the Agonum extensicolle species group (Coleoptera:Carabidae). University of

California Publications in Entomology, 106: 1-198.

Liebherr, J.K. 1986b. A taxonomic revision of the West Indian Platynus beetles

(Coleoptera:Carabidae). Transactions of the American Entomological Society,

112: 289-368.

Liebke, M. 1936. Die Gattung Lachnophorus Dejean (Col. Carabidae). Revista de

Entomologia, 6: 461^68.

Liebke, M. 1938. Denkschrift iiber die Carabiden-Tribus Colliurini. Festschrift zum

60. Geburtstage von Professor Dr. Embrik Strand, 4(1937): 37-141.

Maddison, W.P., M.J. Donoghue, and D.R. Maddison. 1984. Outgroup analysis and

parsimony. Systematic Zoology, 33: 83-103.

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Licininae, Odacanthinae, and Pentagonicinae (Coleoptera). Proceedings of the

Linnean Society of New South Wales, 90: 157-163.

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Motschulsky, V. 1864. Enumeration des nouvelles especes de Coleopteres. Bulletin

de la Societe Imperiale des Naturalistes de Moscou, 37 (part 2): 171-241.

Reichardt, H. 1974. The systematic position of Asklepia Liebke, 1938, with the

description of a new species (Coleoptera, Carabidae). Coleopterists Bulletin, 28:

177-179.

Reichardt, H. 1977. A synopsis of the genera of Neotropical Carabidae (Insecta:

Coleoptera). Quaestiones Entomologicae, 13: 346-493.

Reiche, L. 1843. Coleoptera Colombiana, etc. Revue Zoologique, 6: 37-41.

Sober, E. 1986. Parsimony and character weighting. Cladistics, 2: 28-42.

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Illinois Natural History Survey, Champaign.

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Zoology, 35: 102-109.

Odacanthitae and West Indian Lachnophorini

39

APPENDIX 1

The primitive (0) and derived (1) states of the 57 characters used in the cladistic

analysis of Odacanthitae are described below.

Maxillary Palps

1. 0- apical maxillary palpomere fusiform (Fig. 1 la).

1- apical maxillary palpomere inflated (Figs. 1 Ib-d).

2. 0- apical maxillary palpomere moderately inflated (Fig. 1 Ic).

1- apical maxillary palpomere strongly inflated (Figs. 1 lb, d)

3. 0- apical maxillary palpomere glabrous or with very short setae (Fig. 1 la).

1- apical maxillary palpomere with at least some long setae (Figs. 1 Ib-d).

4. 0- apical maxillary palpomere with sparse long setae (Fig. 1 Ic).

1- apical maxillary palpomere with dense covering of longer setae (Figs.

11b, d).

5. 0- penultimate maxillary palpomere glabrous except for apical ring of setae

(Fig. 11a).

1- penultimate maxillary palpomere setose over length (Figs. 1 Ib-d).

6. 0- setae sparse over length of penultimate maxillary palpomere (Fig. 1 Ic).

1- setae dense over length of penultimate maxillary palpomere (Figs. 11b,

d).

Head

7. 0- antennomeres 2-3 glabrous except for ring of apical setae

1- antennomeres 2-3 setose over entire length.

8. 0- apical 4— antennomeres concolorous with basal segments.

1- apical 4—5 antennomeres pale, basal segments contrasting, darker.

9. 0- neck not constricted.

1- neck with distinct constriction, head pedunculate.

10. 0- mentum medial tooth present.

1- mentum medial tooth absent.

11. 0- subgenal setae absent.

1- subgenal setae present (Fig. 1 1).

12. 0- five or fewer subgenal setae each side of head.

1- six or more subgenal setae each side of head.

13. 0- suture between mentum and gula fully formed and evident.

1- suture between mentum and gula reduced to indistinct depression.

14. 0- suture between mentum and gula reduced but traceable.

1- suture between mentum and gula absent.

Quaest. Ent., 1988, 24 (1)

40

Liebherr

Body Surface

15. 0- integument glabrous except for standard macrosetae.

1- integument with pelage of short setae in addition to standard macrosetae

(Fig. 1).

Pronotum

pronotal base straight (Figs. 1, 14a, d)

pronotal base recurved with a median lobe (Figs. 12a, b).

laterobasal seta present.

laterobasal seta absent.

a single lateral pronotal seta.

more than one seta along medio-lateral margin of pronotum.

pronotal shape quadrate, median length less than or equal to width.

pronotal median length greater than maximum width.

pronotum longer than wide.

pronotum two or more times as long as wide.

pronotal shape cylindrical or quadrate.

pronotum with angulate side, outline hexagonal.

three setae in or adjacent to third elytral interval (Fig. 1).

dorsal elytral setae absent.

three setae in or adjacent to third elytral interval.

more than three dorsal elytral setae.

dorsal elytral impressions small, indistinct.

dorsal elytral impressions foveate (Fig. 1).

dorsal elytral impressions moderately foveate (Fig. 1).

dorsal elytral impressions strongly foveate, very evident.

elytra without transverse depression at basal 0.33.

elytra with transverse depression at basal 0.33.

basal transverse elytral depression weak.

basal transverse elytral depression strongly developed, broad and deep,

elytra with 11-16 macrosetae in 8th interval from base to subapical

sinuation (Fig. 1).

elytra with 17-18 macrosetae in 8th interval from base to subapical

sinuation.

elytra concolorous, or with only subtle differences in color across surface,

elytra with distinct testaceous maculae on a darker ground color,

elytra concolorous or nearly so.

Odacanthitae and West Indian Lachnophorini

41

Flight Wings

36. 0- oblongum cell of flight wing present and complete (Figs. lOg, 13c).

1- oblongum cell with incomplete venation, or absent (Fig. 16e).

37. 0- oblongum cell with incomplete venation (Fig. 16e).

1- oblongum cell absent (Figs. 12f-h).

38. 0- wedge cell of flight wing present (Fig. lOg).

1- wedge cell absent (Figs. 12f-h, 13c, 16e).

39. 0- first anal vein of flight wing present (Fig. lOg).

1- first A vein absent (Figs. 12f-h, 13c, 16e).

40. 0- flight wings constantly macropterous.

1- wings of some individuals brachypterous.

Male Protarsomeres

41. 0- tarsomeres 1-3 moderately broad.

1- tarsomeres 1-3 broader, more robust, mediodorsal surface broadly

flattened.

Pygidial Glands

42. 0- dorsal lobe of defense gland present.

1- dorsal lobe absent.

Female Reproductive Tract

43. 0- basal bulb of spermatheca gradually narrowing to apex (Figs. 9b, 12c, d).

1- basal bulb of spermatheca with apical filament narrower than base of bulb

(Figs. 9c, d, 13a, 14b, e, 15d, 16b).

44. 0- basal bulb of spermatheca wide basally (e.g., Figs. 8a, 13a).

Quaest. Ent., 1988, 24 (1)

42

Liebherr

1- basal bulb of spermatheca with narrow basal duct and apical reservoir

(e.g., Figs. 8c-d, 9a).

45. 0- apical bulb of spermatheca large (e.g., Figs. 4d, 9c, 13a).

1- apical bulb of spermatheca reduced (e.g., Figs. 8, 9a, d, 16d).

46. 0- apical bulb of spermatheca reduced but evident (Fig. 9d).

1- apical bulb of spermatheca absent (Fig. 16d).

47. 0- junction of spermathecal duct and common oviduct lacking distinct

sclerite (Fig. 4d).

1- junction of spermathecal duct and common oviduct with elongate sclerite

(e.g.. Figs. 8, 9, 12c, d, 13a).

48. 0- sclerite at spermathecal duct base present (e.g.. Figs. 12c, d, 13a).

1- sclerite at spermathecal duct base well-developed, U-shaped (e.g.. Figs. 8,

9a).

49. 0- more than 10 setae on penultimate gonocoxite (e.g.. Figs. 14b, 15d).

1- from 5-9 setae on penultimate gonocoxite (e.g.. Figs. 13a, 14e).

50. 0- apical gonocoxite with two lateral ensiform setae (e.g.. Figs. 13a, 16b).

1- apical gonocoxite with at least three lateral ensiform setae unilaterally or

three or four on both sides (e.g.. Figs. 4d, 9b, 14e).

51. 0- apical gonocoxite pointed at tip (e.g., Figs. 9c, d).

1- apical gonocoxite rounded at tip (e.g.. Figs. 13a, 14b, e).

Male Genitalia

52. 0- aedeagal basal bulb complete, base closed (e.g.. Figs. lOa-f).

1- basal bulb of aedeagus incomplete, base open (e.g.. Figs. 12e, 13b, 16a,

c).

53. 0- aedeagal median lobe with curved shaft.

1- aedeagal median lobe with straight shaft.

54. 0- aedeagal median lobe apex simple, acutely rounded (e.g.. Figs. 12e, 13b).

1- aedeagal median lobe apex modified as a knob-like or hook-like

projection (Figs. lOa-f, 15b).

55. 0- aedeagal inner sac without spines or tubercles (e.g., Figs. 10a, 16a, c).

1- aedeagal sac with sclerotized spines (e.g.. Figs. lOf, 15e).

56. 0- aedeagal median lobe shaft cylindrical (e.g., Figs. lOa-f).

1- aedeagal median lobe shaft membranous dorsally with V-shaped sclerite

(e.g.. Figs. 13b, 15b, 16a).

57. 0- parameres subequal.

1- ventral paramere reduced (e.g.. Figs. lOb-e).

Book Notice

43

BOOK NOTICE

LINDROTH, CARL The Carabidae of Fennoscandia and Denmark. Fauna

Entomologica Scandinavica, 15 (1): 1-225 (1985), and 15 (2): 233^95, with an

appendix on the Family Rhysodidae, pp. 496-497 (1986). E. J. Brill, Publishers,

Scandinavian Science Press Ltd., P.O. Box 9000, 2300 PA, The Netherlands. Price:

$30.25 (U.S.), each part.

This two-parted volume was completed some seven years after the death of the

author, Carl H. Lindroth, that remarkably talented Swedish entomologist, who

contributed so extensively to the literature of Holarctic taxonomy, ecology, and

biogeography. The manuscript for the present volume, incomplete at the time of

Lindroth’s death, was completed by an international group of his colleagues,

acknowledged on the title page: F. Bangsholt, R. Baranowski, T. L. Erwin, P.

J0rum, B.-O. Landin, D. Refseth, and H. Silfverberg. Their contributions were

appreciable, as is generally true of those who accept the task of completing the work

begun by another.

The fine series, of which this volume is a part, provides faunistic treatments

intended primarily for species identification, but additionally as something of an

introduction to the treated taxon, so that a novice reader could use it as the basis for

more extensive studies. Thus, in the first 23 pages of this volume are found in

addition to advice about “collecting”, “killing and mounting”, “labelling”, “notes on

identification” and “distribution and state of knowledge” (of the

Fennoscandian-Danish [i.e., Nordic] species), a general discussion of natural history

of carabids, notes about larval characters, the system of carbid classification in

general, comments about the principles and practice of nomenclature, and a review

of the literature about Nordic Carabidae that will serve as a context for more

extended reading. There is also a “new classification of Carabidae” (after Erwin and

Sims) that introduces a user to a thoroughly up to date arrangement of higher taxa of

Carabidae. The extensive bibliography, which ends the text in Part 2, is also a

valuable source of information for more extended reading about carabids.

Five hundred and fourteen excellent black and white illustrations grace the pages

of this volume. For most genera, a habitus illustation is provided of a typical adult of

the taxon. Diagnostic features of most taxa are illustrated, including details of form

of head, various thoracic and abdominal sclerites, elytra, legs, and genitalia. In

addition, eight plates of excellent photographs illustrate not only habitus but also

color and luster of the dorsal surfaces of adults of 130 species. Most of the line

drawings came from the pen of the gifted author, but many of the habitus drawings

are labelled “After Victor Hansen”.

A conservative traditional classification serves as the basis for arrangement of

taxa, beginning with tiger beetles and ending with bombardier beetles. Supraspecific

categories used throughout are subfamily, tribe and genus. The five subfamilies and

23 tribes of the Nordic fauna are characterized very succinctly in terms of structural

Quaest. Ent., 1988, 24 (1)

44

Book Notice

features, and with generalized statments about geographical distribution for some of

these taxa, as well. The 63 genera are more fully treated, including notes about life

history features common to the included species.

The subgeneric category is used for eight genera, five of which are markedly

diverse, with 20 or more species in the Nordic area. For the more diverse genera, the

utility of using a lower ranking supraspecific category is clear. It is less clear why

the author used that category for some of the less diverse genera. Thus, subgenera

hardly seem necessary in a work of this scope for Calathus (seven species), Badister

(nine species), and Dromius (14 species).

Most supraspecific taxa recognized are in accord with Lindroth’s previous

publications. In the Pterostichini, however, I noted that Dolichus Bonelli is given

generic status, though it was previously combined with Calathus Bonelli— correctly

so, I believe.

Each of the 400 species is designated consecutively by number. For each, a brief

description is provided, featuring color and easily observed structural details that

characterize the adults and that are of maximum use in identification. For many

species of Bembidion and of difficult groups in other genera, diagnostic details of

the male genitalia are also recorded. Short accounts of geographical distribution,

both within and beyond the Nordic area, and of “biology”, complement the

morphological description, to provide in total a very clear and compact

characterization of each species.

Folllowing the taxonomic portion of the text in each Part is a matrix giving the

Nordic distribution of each species included, in terms of country and region within

each country. Location of each of the regions is clearly indicated on maps printed on

the front and back endpapers of each Part.

The text is in English, throughout. The print, on good quality paper, is easy to

read. The binding, of stiff cardboard coated with emerald green paper seems

excellent and designed for long use.

This book must be on the shelves of all Nordic coleopterists, ought to be owned

by carabid specialists wherever they may live, and biological bibliophiles will find it

a welcome addition to their collections.

My memories of Carl Lindroth are still so vivid that I find it hard to believe he is

no longer alive. The appearance of this excellent volume, written in his style and

containing many of the figures that he drew makes it no easier for me to accept the

fact that we will never see him again. Though he is gone physically, his

many-splendored life continues to reward and inspire those of us fortunate enough

to have known him. His published works, including the one reviewed here, provide

evidence, to those who came too late to know him personally, of his remarkable

qualities as a systematic entomologist.

George E. Ball

45

COMMENTARY

“Commentary” is a new section of Quaest. Ent. that will appear from time to

time, and will contain expressions of opinions about general items, controversial or

otherwise, that ought to be of interest to many of our readers. These contributions

will not be refereed because they are intended to be free expressions of opinion.

Changes by the Editor might be made to the form of presentation, but not to its

substance. Remarks that are deliberately abusive or insulting will not be published.

Rebuttals to previously expressed views will be considered, but the journal is under

no obligation to publish them.

The paper inagurating “Commentary” addresses an issue that is becoming

controversial in cladistic systematics. The author is a distinguished Norwegian

dipterist, known principally for his extensive work on and publications about

chironomid flies.

The Editor

ON THE LIMITATIONS OF PARSIMONY IN PHYLOGENETIC

ANALYSIS

Ole A. Saether

Museum of Zoology

University of Bergen

N-5000 Bergen

Norway Quaestiones Entomologicae

24:45-50 1988

All sciences unselfconsciously use parsimony as a criterion of hypothesis choice,

i.e., as a way of measuring support of alternative hypotheses against a body of

evidence. In phylogenetic reasoning “parsimony interprets synapomorphy as

evidence for phylogenetic relationship, but denies that symplesiomorphy has this

significance” (Sober, 1986:28). Parsimony, however, (again citing Sober) does not

itself suffice to solve the problem of incongruent data; character weighting must

play an indispensable role. “Parsimony is not a device that tells biologists how to

weigh characters; rather parsimony requires that the characters already should be

weighted.” Parsimony, however, also enters into the process of character weighting.

And character weighting carries with it substantive assumptions about the

evolutionary processes.

The question is, then, will the use of parsimony in choosing among alternative

hypotheses of explanations of single character distributions, lead to unparsimonious

Quest. Ent., 1988, 24(1)

46

Saether

results when reconstructing genealogy according to the methods of quantitative

phyletics and transformed cladism? My contention is that it will. To show this was

the main aim of my recent papers (Saether 1983, 1986) and not, as Farris (1986)

maintains, to “bolster a contention that postulated homoiologies can be used as

evidence on phylogenetic relationships.” I believe that when the monophyly of

different groups has been established by “true” synapomorphies, an underlying

synapomorphy, but only when appearing as a unique inside parallelism (Brundin

1976) showing principal deviation, may assert monophyly for a collective taxon.

Whether or not underlying synapomorphies are operationally recognizable is beside

the point, which is that they occur and apparently commonly.

There are four main categories of explanations for the distribution of alternative

character transformations of an apomorphic trend within a monophyletic group

(Saether 1983). The apomorphy may be explained as: (1), uniquely derived, an

“objective” synapomorphy; (2), as caused by parallel selection or convergence (both

giving the same distribution of character alternatives); (3), secondary reduction,

reversal or change; or (4), underlying synapomorphy i.e.„ inherited factors causing

incomplete synapomorphy. I agree fully with Farris (1986:15) in summarizing

parsimony. Genealogical hypotheses are indeed “potentially able to explain

observed points of similarity among organisms as the result of inheritance from a

common ancestor.” “An observation is said to provide evidence favoring a first

hypothesis over a second when the first is better able to explain the observation.” “A

genealogy that is consistent with a single origin of some trait is able to account for

all similarities in that trait as inheritance.” “Each additional requirement for a

separate origin of a feature reduces the explanatory power of a theory of

phylogenetic relationship.”

To find the hypothesis best able to explain the observed similarities, then, it is

necessary to minimize the requirements for independent origins, the most

parsimonious explanation for the distribution of alternative character

transformations. I have shown (Saether, 1983) that for very many types of character

distributions underlying synapomorphy is the most parsimonious solution, and

except for uniquely derived “objective” synapomorphy, it always is as good as any

alternative explanation. The more times the apomorphic alternative appears

separately in the different branches of a monophyletic group, the more all other

explanations require additional separate origins, while underlying synapomorphy

still is able to account for all similarities in the trait as inheritance with a single

origin.

Farris (1986) rejects underlying synapomorphies since they lead to

unparsimonious results in his method of reconstructing phytogeny. One of my

objections to neocladistic methods is that they disregard the most parsimonious

explanations of character distributions. I agree with Farris (1985:196) that “any

theory that implies both parsimony and a non-parsimonious method must be

self-contradictory.” In a qualitative method all characters, including underlying

Parsimony in Phylogenetic Analysis

47

synapomorphy, can be properly judged and weighted in such a way that the search

for sister groups can continue in a parsimonious manner. Farris (1986) refers to me

as claiming that in quantitative phyletics “the most parsimonious tree is taken as the

truth I stated that there is no biological evidence that minimum length trees are

most in accordance with the “tree”. If Farris is not interested in finding the

genealogical hypothesis most likely to be in accordance with the “true” tree, we

have different purposes in erecting synapomorphy diagrams.

If underlying synapomorphy always is at least as good an explanation as parallel

selection or secondary change why is it not used as the prime explanation? Farris

touches on this in his contention that I omit an explanation for underlying

synapomorphy. An explanation of the possible mechanisms is given in Saether

(1983). The potential information classes of Wiley and Brooks (1982:4) are

descriptions of underlying synapomorphy. However, the mechanisms in question

apparently cause the expression or non-expression to function like an on-off switch.

There seldom are intermediates. Nonetheless, secondary reduction and parallel

selection often can be followed through several steps often corresponding with

environmental changes and obvious functional adaptations. Usually, they are easy to

spot, with sufficient knowledge of the group studied. Knowledge and judgment, or

what Humphries and Camus (1986:95) label intuition, are alternative necessary

elements in choosing between hypotheses of character distribution. To repeat the

citation from Hennig (1966:120) given in my paper: “Characters cannot be

considered in isolation, even in regard to transformation series of other characters.”

According to Farris, one of the main grounds offered by me for using

homoiology should be that Hennig recommended it. I stated that Hennig in his

theoretical works did not recommend it. Hennig (1966), as also stated by Farris,

dismissed parallelism as equivalent to convergence for purposes of phylogenetic

analysis, as he dismissed multiple derivations. In his practical work, however, he

used both. In the citation by Farris, Hennig (1966:121) discussed the interpretation

of character distributions. The first alternative for choosing between two

contradictory characters with respect to phylogenetic relations is that “(1) It was

erroneous to interpret one or both characters as plesiomorphous or apomorphous”;

i.e.„ they are not synapomorphous. Multiple derivations of a synapomorphy thus are

included in (2) parallelism. Yet, as shown by me and repeated by Farris, Hennig, in

actual practice, used underlying synapomorphies as if they were “true”

synapomorphies.

The first paper in which I used the term underlying synapomorphy (Saether,

1977) was refereed by Hennig. He did not raise any objection to the term. Schlee

was a pupil and close collaborator of Hennig. His opinions of Hennig ’s theoretical

viewpoints therefore carry particular weight. While Schlee (1968) in his excellent

revision of the Corynoneura-group follows Hennig ’s practical applications, in his

paper of 1971 and particularly 1975 (a) he noted the consequence of Hennig ’s

theoretical writings. Here, Schlee recognizes uniquely derived synapomorphies, i.e.,

Qaest. Ent., 1988, 24 (1)

48

S aether

“objective” synapomorphies, as the only basis for reconstructing genealogy since all

other alternatives, including subjective synapomorphy, would cause loss of

explanatory power in the reconstructed genealogy. He is strongly critical of Brundin

(1966) as well as of his own work from 1968. That Farris finds Schlee’s paper

unobjectionable and undogmatic is a result of reference to the wrong paper. (Schlee,

[1975 b] a critique of numerical phyletics.) It is, however, clear that if using the

criteria of Schlee for asserting synapomorphy, i.e.„ using “objective” synapomorphy

only, both neocladistic methods and transformed cladism would be workable, but

superfluous.

A complete set of “objective” synapomorphies, is, by definition, the “true”

phylogenetic tree and is easily transformed into a dichotomous diagram without the

help of calculators or computers (Schlee 1975 b). The number of “objective”

synapomorphies, however, usually is sufficiently large only when comparing higher

taxa (Saether 1986). Clique methods, transformed cladism, and dogmatic cladism all

either recognize “objective” synapomorphy, only, or give such features nearly

exclusive weight. It is thus not a deception, as maintained by Farris, to state that the

same critique is valid against all of those three systems of analysis even when the

exponents of each object to the other’s methods.

Farris, in his chapter on homoiology includes parallelism {i.e., parallel selection,

homoiology and, by implication, multiple derivations) with convergence in

homoplasy. He equates homoplasy with nonhomology, as is common practice. As

shown by Saether (1983), if characters are defined as including different character

states, parallel selection will be included in definitions of homoplasy, but not in

nonhomology. Underlying synapomorphies are homologies, the common character

consisting in the common capacities to develop the same feature. They can be

regarded both as homoplasious or as non-homoplasious depending on whether the

character itself or the capacity to develop the character is considered. When I

maintained that homoplasy is “nearly universal” I was referring to the wider

meaning of homoplasy as used by Farris. When homoplasy is involved in 275 of

291 trends (Saether 1986, table 1), and only two of the 291 trends are “objective”

synapomorphies showing principal deviations, homoplasy must be regarded as

“nearly universal”. This also clearly demonstrates the need for using all available

characters in estimating genealogy. If Farris dismisses the “necessity argument”

(Farris 1986:16) why does he recommend the use of characters showing outside

parallelism, i.e.„ multiple derivations?

It is of little importance whether one of the methods of Felsenstein, Estabrook or

Farris are better than the other for estimating genealogy. My intention was to show

that while they all are quantitative methods and fail to take all types of characters

into account, they lead to different results and are based on different philosophies. (I

did not, as stated by Farris, cite Patton and Avis (1983), but Dupuis (1984) as

advocating qualitative over quantitative techniques.)

Parsimony in Phylogenetic Analysis

49

Quantitative phyletics, however, is well-suited for estimation of anagenetic

levels (Saether 1979). Farris significantly neglects to address the important

distinction between cladogenetic and anagenetic trends and analyses. Cladogenetic

analysis should be qualitative, while anagenetic analyses are quantitative. The

search for sister groups, i.e.„ cladogenetic analysis, involves choosing theoretical

ancestors for later species. Starting with the lowest taxa, for instance species, to be

included in the analysis the scheme of argumentation is gradually built up to include

more inclusive groups. It is what Sober (1986) calls thinking backward. In order to

include a species in a synapomorphy diagram, it is in principle sufficient to know

that one single character it shares with its sister species is uniquely derived.

Anagenetic analysis, on the other hand, evaluates the attributes of the theoretical

progenitor of the monophyletic group under study and estimates deviations from the

ancestor in its proposed descendants. In principle, to find the correct anagenetic

levels for all included taxa, every single trend evaluated needs to be scored as

apomorphous or plesiomorphous relative to the progenitor. This is what Sober calls

forward thinking. The difference in treatment will be clear for instance from Saether

(1976) which forms the base for table 1 in Saether (1986). It can be noted that

underlying synapomorphies were not used as evidence of phylogenetic relationships.

In the synapomorphy diagrams forming the base for tables 2 and 3 (Saether, 1986)

only a few underlying synapomorphies showing unique inside parallelism of

principal deviations were used.

There is nothing wrong with a posteriori postulation of natural processes from

patterns in the distribution of characters, testing these patterns in other schemes of

argumentation, and than using the theories in the building of genealogical

hypotheses. As stated by Ridley (1986), the argument proceeds not in a circle, but

by “successive approximation”. The sequence - hypothesis, test, further hypothesis -

is not circular. The chain of inductive-deductive or hypothetico-deductive reasoning

is a common scientific method. What transformed cladists, numerical taxonomists,

and, apparently Farris, object to is not circular reasoning, but theoretical ideas.

Theories, in science, if they are interesting, are always taken beyond evidence, and

assumed in further tests (Ridley [1986]). Hennig, as shown by the last citation in my

paper, recomended that this be done.

LITERATURE CITED

Brundin, L. 1966. Transantarctic relationships and their significance, as evidenced

by chironomid midges, with a monograph of the subfamilies Podonominae and

Aphroteniinae and the austral Heptagyiae. Kungl. Svenska Vetenskapakedmiens

Handllingar, (4) 11: 1-472.

Brundin, L. 1976. A Neocomian chironomid and Podonominae-Aphroteniinae

(Diptera) in the light of phylogenetics and biogeography. Zoologica Scripta 5:

139-160.

Qaest. Ent., 1988, 24 (1)

50

S aether

Dupuis, C. 1984. Willi Hennig’s impact on taxonomic thought. Annual Review of

Ecology and Systematics 15: 1-24.

Farris, J. 1985. The pattern of cladistics. Cladistics 1: 190-201.

Farris, J. 1986. On the boundaries of phylogenetic systematics. Ibid. 2: 14-27.

Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana.

Humphries, J.C. and J.M. Camus. 1986. Contemporary issues in systematics.

Cladistics 2: 85-99.

Patton, J.C., and J.C. Avise. 1983. An empirical evaluation of qualitative Hennigian

analysis of protein electrophoretical data. Journal of Molecular Evolution 19:

244-254.

Ridley, M. 1986. Evolution and classification. The reformation of cladism.

Longman Inc., New York.

Saether, O.A. 1976. Revision of Hydrobaenus, Trissocladius, Zalutschia,

Paratrissocladius, and some related genera (Diptera: Chironomidae). Bulletin of

the Fisheries Research Board of Canada 195: 1-287.

Saether, O.A. 1977. Female genitalia in Chironomidae and other Nematocera:

morphology, phylogenies, keys. Ibid. 197: 1-211.

Saether, O.A. 1983. The canalized evolutionary potential: Inconsistencies in

phylogenetic reasoning. Systematic Zoology 32: 343-359.

Saether, O.A. 1986. The myth of objectivity - post-Hennigian deviations. Cladistics

2: 1-13.

Schlee, D. 1968. Vergleichende Merkmalsanalyse zur Morphologie und

Phylogenetic der Corynoneura-Gm^^t. (Diptera, Chironomidae). Zugleich eine

allgemeine Morphologie der Chironomiden-Imago (o). Stuttgarter Beitrdge zur

Naturkunde aus dem staatlichen Museum fur Naturkunde im Stuttgart 180:

1-150.

Schlee, D. 1971. Die Rekonstruktion der Phylogenese mit Hennig’s Prinzip.

Aufsdtze und Reden der senckbergischen Naturforschenden Gessellschaft 20:

1^2.

Schlee, D. 1975 a. Das Problem der Podonominae-Monophylie (Diptera).

Entomologica Germanica 1:31 6-35 1 .

Schlee, De. 1975 b. Numerical phyletics: An analysis from the viewpoint of

phylogenetic systematics. Entomologica Scandinavica 6: 193-208.

Sober, E. 1986. Parsimony and character weighting. Cladistics 2: 28^2.

Wiley, E. O. and D.R. Brooks. 1982. Victims of history. A nonequilibrium approach

to evolution. Systematic Zoology 31: 1-24.

THE CICINDELA SYLVATICA GROUP: GEOGRAPHICAL VARIATION

AND CLASSIFICATION OF THE NEARCTIC TAXA, AND

RECONSTRUCTED PHYLOGENY AND GEOGRAPHICAL HISTORY OF

THE SPECIES (COLEOPTERA: CICINDELIDAE).*

T. G. Spanton

Department of Entomology

The University of Alberta

Edmonton, Alberta T6G 2E3 Quaestiones Entomologicae

24:51-161 1988

ABSTRACT

The Cicindela sylvatica species group as defined by Rivalier (1950, 1954)

includes the North American species C. longilabris Say and C. nebraskana Casey, as

well as the Palearctic species C. sylvatica Linnaeus, C. granulata Gebler, C. japana

Motschulsky, C. gemmata Ealdermann, C. soluta Dejean and C. lacteola Pallas.

Discriminant analysis of morphometric data, univariate analysis of qualitative

characters, and study of the distribution and soil associations of adult specimens of

Cicindela longilabris Say and C. nebraskana Casey show that C. longilabris is a

boreal and montane species living on Podzolic soils in eastern North America, and

Luvisolic and Brunisolic soils of coniferous forests in western North America, as

well as in boreal forest-grassland transition areas. Its three subspecies are: (1), C.

longilabris longilabris Say, across the boreal zone from Newfoundland and New

England to Alaska; (2), C. 1. laurentii Schaupp, in the Rocky Mountain region of the

United States, including isolated populations of northern New Mexico, eastern

Arizona, northern Arizona and southwestern Utah; and (3), C. I. perviridis Schaupp,

living in the Sierra Nevada and Cascade Mountains of California, Oregon and

Washington. An area of hybridization includes southwestern Alberta, southeastern

British Columbia, eastern Washington, northern Idaho and northwestern Montana

where the three subspecies converge geographically. Cicindela nebraskana is a

monobasic species which lives on Chernozemic soils of prairie grasslands and

grassland-forest transition zones of western North America.

The following new synonymies are presented; Cicindela longilabris longilabris

Say f=Cicindela longilabris novaterrae Leng); C. 1. laurentii Schaupp (=C. 1. oslari

Leng); C. 1. perviridis Schaupp (=C. 1. ostenta Casey); C. nebraskana Casey (=C.

nebraskana chamberlaini Knaus).

^Modified from a thesis submitted to Lakehead University, Thunder Bay, Ontario, in partial fulfdlment of

the requirements for the degree of Master of Science.

52

Spanton

Individuals of C. longilabris live for three years: two winters are passed in the

larval stage and one winter in the adult stage before mating and oviposition.

A male neotype specimen is designated for the species C. longilabris Say, from

Ontario, Thunder Bay District, Sibley Provincial Park, 1 km W of Silver Islet.

Phylogenetic analysis shows three pairs of sibling species: C. soluta-C.

gemmata, C. sylvatica-C, granulata and C. longilabris-C. nebraskana, with C. japana

most closely related to the soluta-gemmata pair. Cicindela lacteola was either

derived earlier in the evolution of the group; or in the absence of a synapotypy for

C. lacteola and the other species of the group, it is possible that this species should

not be classified in this species group.

RESUME

Le groupe-espece Cicindela sylvatica definis par Rivalier (1950, 1954} inclus I’especes Nord

Americaines Cicindela longilabris Say, et Cicindela nebraskana Casey, de mime que les especes paleartiques

C. sylvatica Linnaeus, C. granulata Gebler, C. japana Motschulsky, C. gemmata Faldermann, C. soluta

Dejean et C. lacteola Pallas.

L’ analyse discriminante des donnees morphometriques, I' analyse univariee des characteres qualitatifs,

ainsi que I association entre le type de sol et les adultes de Cicindela longilabris Say et de C. nebraskana

Casey demontrent que C. longilabris est une espece boreale et de montagne habitant sur les sols Podzoliques

de I' est de I’Amerique du Nord, et sur les sols Luvisoliques et Brunisoliques des forets de coniferes de

I’ouest du continent, ainsi que dans les regions de transition entre prairie et for it boreale. Ses trois

sous-especes sont: (1), C. longilabris longilabris Say (zone Boreale entre Terre-Neuve, la Nouvelle

Angleterre et 1’ Alaska); (2), C. 1. laurentii Schaupp, (regions des Montagnes Rocheuses des etas-Unis,

incluant des populations isolees du nord du Nouveau Mexique, de Lest et du nord de I’ Arizona, et du

sud-ouest de I’Utah); et (3) C. 1. perviridis Schaupp, (Sierra Nevada et les montagnes Cascades de la

Californie, Oregon et Washington). Une region d’ hybridation inclus le sud-ouest de I’ Alberta, le sud-est de

la Colombie Britannique, Lest de I’etat de Washington, le nord de 1’ Idaho et le nord-ouest du Montana, ou

les trois sous-especes convergent geographiquement. Cicindela nebraskana est une espece monotypique qui

habite les sols Chernozemiques des prairies et des regions de transition prairie - forit boreale de I’ouest de

I’Amerique du Nord.

Les nouvelles synonymies suivantes sont presentees: Cicindela longilabris longilabris Say (=C. 1.

novaterrae Leng); C. 1. laurentii Schaupp (=C. 1. oslari Leng); C. 1. perviridis Schaupp (=C. 1. ostenta

Casey); C. nebraskana Casey (=C. n. chamberlaini Knaus).

Les individus de C. longilabris ont une longevite de trois ans: Us vivent deux hivers au stade larvaire, et

le troisieme au stade adulte avant d’ engager la copulation et L oviposition le printemps suivant.

Un specimen neotype, male est designe pour 1’ espece C. longilabris de I’ Ontario, district de Thunder

Bay, Parc Provincial de Sibley, 1 km ouest de Silver Islet.

L’ analyse phylogenique demontre qu’il y a trois paires d’ especes-soeurs: C. longilabris-C. nebraskana,

C. sylvatica-C. granulata, et C. soluta-C. gemmata, avec C. japana s’ apparentant le plus a la paire C.

soluta-C. gemmata. Cicindela lacteola c’est developpee, soit tres tot au cours de revolution du groupe, soit

en I’ absence d’ une synapotypie entre C. lacteola et les autres especes du groupe, il est possible que cette

espece ne devrait pas etre classifiee dans ce groupe-espece.

TABLE OF CONTENTS

Introduction 53

Materials and Methods 54

Material 54

The Cicindela sylvatica Group

53

Methods 56

Structural and Ecological Features of the Nearctic Taxa of the Cicindela sylvatica

Group 72

Analysis of Adult Features 81

Larvae and Life History Data 99

Soil Associations 1 12

Classification of the Nearctic taxa of the Cicindela sylvatica Group 123

Cicindela longilabris Say 123

Cicindela nebraskana Casey 131

Genitalia and Evolutionary Aspects of the Cicindela sylvatica Group 134

Genitalia of the C. sylvatica Group 134

Reconstructed Phytogeny 144

Geographical History 150

Concluding Remarks 153

Acknowledgements 153

Literature Cited 154

INTRODUCTION

The adults of most taxa of tiger beetles in the Nearctic and Palearctic regions

have been described and named for some time, but only in the past 20 years have

extensive studies of intraspecific variation, and classification of species groups of

Cicindela been conducted. For example, Freitag (1965) studied the geographic

variation and distribution and revised the North American species of the Cicindela

maritima group. Willis (1967) studied geographic variation and ecology of a diverse

group of Cicindela species living in saline habitats in the central United States,

Gaumer (1977) studied intraspecific variation and taxonomy of adults and larvae of

C.formosa, Murray (1980) studied geographic variation of C. rufiventris Dejean, C.

sedecimpunctata Klug, and C. flohri Bates, and Kaulbars (1982) studied the

morphological and ecological variation of the species of the C. sexguttata species

group.

Wallis (1961) confused many of the intraspecific taxa of C. longilabris and C.

nebraskana but indicated that his understanding of these species was based on few

specimens and suggested that this group required additional study. Leffler and

Pearson (1976) indicated that revisionary study of C. nebraskana and C. longilabris

was required to establish the taxa and to correctly apply the available names.

The immature stages and life histories of a number of tiger beetle species have

been studied (Shelford, 1908; Griddle, 1907, 1910; Hamilton, 1925; Willis, 1967,

among others) but the life histories of C. longilabris and C. nebraskana are largely

unknown. The larvae of C. longilabris are known only from one third instar

specimen and three exuviae (Leffler, 1979).

Quaest. Ent., 1988, 24 (1)

54

Spanton

This study began as an investigation of the patterns of geographic variation in the

Cicindela longilabris-C . nebraskana complex. For this reason a detailed treatment

of the Nearctic taxa of the C. sylvatica group is presented. After investigating

genitalic structures of the members of the C. longilabris-C. nebraskana complex,

similar genitalic dissections were made of the other species in the sylvatica group to

determine what degree of structural difference might occur among the species of the

group. As some synapotypies became apparent in genitalic structures, I decided to

carry out a cladistic analysis to hypothesize' phylogenetic relationships among the

species of the group. Detailed study of the pattern of geographical variation of the

Palearctic species was beyond the scope of this study.

The objectives of this study were: (1), to determine if C. longilabris and C.

nebraskana are conspecific, or distinct species; (2), to investigate the pattern of

intraspecific variation in these species; (3), to establish the North American taxa and

correctly apply the available names; (4), to investigate the life history of C.

longilabris and describe immature life stages, as possible; and (5), to carry out a

phylogenetic analysis of the species of the C. sylvatica group to hypothesize

relationships among the taxa.

MATERIALS AND METHODS

Material: Adult Specimens and Loaning Institutions

More than 6,210 adult specimens were examined. Most specimens were obtained

on loan from the following institutions and private collections. I have used standard

codens for collections of insects as proposed by Heppner and Lamas (1982),

wherever possible. For private collectors, initials were used as codens. Curators,

and/or staff members with whom I corresponded, are named following their

respective institutional addresses.

AAM Alan and Anne Morgan, Departments of Earth Sciences and Biology

respectively. University of Waterloo, Waterloo, Ontario N2L 3G1

AMNH American Museum of Natural History, New York, New York 10024; L.

H. Herman

BGSU Bowling Green State University, Bowling Green, Ohio 43403; R. C.

Graves

CAS California Academy of Sciences, San Francisco, California 94118; D. H.

Kavanaugh

CDF Clifford D. Ferris, P. O. Box 3351, University Station, Laramie,

Wyoming 82071

CMP Carnegie Museum of Natural History, Pittsburgh, Pa. 15213 ; R. L.

Davidson

CNC Canadian National Collection of Insects, Biosystematics Research Centre,

Ottawa, Ontario KIA 0C6; J. E. H. Martin

CSU Colorado State University, Fort Collins, Colorado 80523; H. E. Evans

The Cicindela sylvatica Group

55

CU Cornell University, Ithaca, New York 14853; L. L. Pechuman, Q. D.

Wheeler

ISU Iowa State University, Ames, Iowa 50011; R. E. Lewis

KSU Kansas State University, Manhattan, Kansas 66506; H. D. Blocker

MPM Milwaukee Public Museum, Milwaukee, Wisconsin 53233; G. R. Noonan

MSU Montana State University, Boseman, Montana 59717; S. Rose

MUN Memorial University of Newfoundland, St. John’s, Newfoundland AIB

3X9; D. J. Larson

NAU Northern Arizona University, Flagstaff, Arizona 86001; C. D. Johnson

NCSR North Carolina State University, Raleigh, NC 27650; C. Parron

NDSU North Dakota State University, Fargo, ND 58105; E. U. Balsbaugh

OKS Oklahoma State University Natural and Cultural History Museum,

Stillwater, OK; W. A. Drew

PSU Pennsylvania State University, University Park, PA 16802; D. L. Pearson

PUL Purdue University, Lafayette, Indiana 47907; A. Provonsha

REA Robert E. Acciavatti, 2111 Cherry Street, Morgantown, West Virginia

26505

SMEK Snow Museum of Entomology, University of Kansas, Lawrence, KS

66045; G. W. Byers

UAE University of Alberta, Edmonton, Alberta T6G 2E3; G. E. Ball, D.

Shpeley

UAE University of Arkansas, Fayetteville, Arkansas 72701; R. Chenowith, C.

Carleton

UBC University of British Columbia, Vancouver, B.C.; S. G. Cannings

UIM University of Idaho, Moscow, Idaho 83843; W. F. Barr

UMAA University of Michigan, Ann Arbor, Michigan 48109; T. E. Moore, M. F.

O’Brien

UMW University of Manitoba, Winnipeg, Manitoba R3T 2N2; T. D. Galloway

UNM University of New Mexico, Albuquerque, NM 87131; C. S. Crawford

UOG University of Guelph, Guelph, Ontario NIG 2W1; D. Pengelly, S. A.

Marshall

USNM National Museum of Natural History, Smithsonian Institution,

Washington, DC 20560; T. L. Erwin

USU Utah State University, Logan, Utah 84322; W. J. Hanson

UV University of Vermont, Burlington, Vermont; R. T. Bell

UWM University of Wisconsin, Madison, Wisconsin 53706; S. Krauth

WJ Walter Johnson, 2917 16th Avenue South, Minneapolis MN 55407

WSU Washington State University, Pullman. WA 99164; R. Zack

Additional specimens were acquired by field collecting. During a ten week

collecting trip in the summer of 1981, specimens were collected in situ and

observations were made of the habitats of these beetles over much of the geographic

Quaest. Ent., 1988, 24 (1)

56

Spanton

range of C. longilahris and C. nebraskana in Canada and the United States.

Specimens were also collected in the summer of 1982 in western Ontario and

northern Minnesota.

Methods

Characters and measurements. — A number of measurements were taken to

investigate differences in size and body proportions among populations of species of

the C. longilahris-C. nebraskana complex.

Characters of the labrum of the adult stage were examined because the large size

of the labrum in C. longilabris and C. nebraskana has been used to distinguish

between these and other North American tiger beetle species (Willis, 1968), and

because the color of the labrum is dark in many specimens of C. nebraskana,

especially in females, whereas it is light tan in most specimens of C. longilabris

(Leffler, 1979).

The number and pattern of setae on the antennal scape has been used to

distinguish among species of Cicindela (Willis, 1968).

Color and pattern of markings have figured prominently in descriptions of tiger

beetles. In many groups of tiger beetles, and especially in the C. longilabris-C.

nebraskana complex, many specific and subspecific names have been applied to

individual variants, and to variant populations, based solely on differences in color

and pattern of markings. I attempted to elucidate the pattern of variation in

characters of color and color pattern, across the range of these two species. The

following characters were used in either the numerical analyses, or qualitative

character analyses of this study. The alphanumeric characters in brackets following

each character listed below are abbreviations used in this paper.

1. Total head width across the widest point on the eyes (hw)(Fig. 1)

2. Length of labrum including the median tooth (ll)(Fig. 1)

3. Width of labrum (lw)(Fig. 1)

4. Ratio: length of labrum/width of labrum (11/lw)

5. Color of labrum (Icol). I arbitrarily assigned three states for this character.

1 . uniformly pale in color, or pale except for a darkened apical margin.

2. apical margin and midrib broadly darkened or mottled.

3. uniformly dark brown or black.

The setal pattern on the labrum was used as a set of characters (6-9).The

number of setae in each of four locations on the frontal surface of the labrum

was indicated (Fig. 2). Each of these loci was treated separately because the

number of setae was, in many instances seen to vary from one side of the

animal to the other (i.e. positions 1 and 4; positions 2 and 3).

6. Number of setae in position 1 (Isl)

7. Number of setae in position 2 (ls2)

8. Number of setae in position 3 (ls3)

Quaest. Ent., 1988, 24 (1)

58

Spanton

1mm

Figs. 4-5. 4. Adult prothorax, dorsal aspect: pw, pronotal width; pi, pronotal length. 5. Left mesothoracic

leg, anterior aspect: fl, femur length; tl, tibial length.

The Cicindela sylvatica Group

59

m /

m b

el

al

1

m m

e w

Fig. 6. Adult elytra, dorsal aspect: el, elytral length; ew, elytral width; hi, humeral lunule; mb, middle band;

ml, marginal line; al, apical lunule.

Quaest. Ent., 1988, 24 (1)

Spanton

60

7

D

E

F

2mm

Fig. 7. Percent of elytral surface covered by maculations: A, one per cent; B, two per cent; C, five per cent;

D, ten per cent; E, twenty per cent; F, thirty per cent.

The Cicindela sylvatica Group

61

0

1

2

2mm

5

Fig. 8. Humeral lunule character states. The number at the lower left of each drawing indicates arbitrarily

assigned values.

Quaest. Ent., 1988,24(1)

62

Spanton

2mm

J

Fig. 9. Middle band character states. The number at the lower left of each drawing indicates arbitrarily

assigned values. A zero was assigned if the middle band was absent.

I

2mm

Fig. 10. Apical lunule character states. The number at the lower left of each drawing indicates an arbitrarily

assigned value.

Quaest. Ent., 1988, 24 (1)

64

Spanton

TABLE 1. Designated states for color of the dorsal surface of elytra of C.

longilabris and C. nebraskana.

*Kelly and Judd, 1965.

TABLE 2. Designated states for color of the proepistemum of C. longilabris and C.

nebraskana.

Kelly and Judd, 1965.

The Cicindela sylvatica Group

65

TABLE 3. Designated states for color of the venter of the abdomen of C. longilabris

and C. nebraskana.

*Kelly and Judd, 1965.

9. Number of setae in position 4 (ls4)

10 Number of sensory setae on the first antennomere (scape) of the left antenna

(ssl) (Fig. 3)

1 1 Number of sensory setae on the scape of the right antenna (ssr)

12 Number of other setae on the scape of the left antenna (osl)

13 Number of other setae on the scape of the right antenna (osr)

In characters 10 to 13 the corresponding number of setae on left and right sides

were treated separately because this was seen to vary in some instances from

left to right antenna of the same individual.

14 Pronotal width (pw)(Fig. 4)

15 Pronotal length (pl)(Fig. 4)

16 Ratio: pronotal width/pronotal length (pw/pl)

17 Mesothoracic femur length (fl)(Fig. 5) The left mesothoracic leg was chosen

preferentially. Where the left was missing, the same measurement was taken

from the right leg. The mesothoracic leg was chosen because prothoracic and

metathoracic legs were more frequently missing from pinned specimens.

18 Mesothoracic tibia length (tl)(Fig. 5)

19 Ratio: mesofemur length/mesotibia length (fl/tl)

20 Length of left elytron (el)(Fig. 6). This was measured from the apex of the

scutellum along the suture to its apex.

21 Width of left elytron at its widest point (ew)(Fig. 6). This was measured with

Quaest. Ent., 1988, 24 (1)

66

Spanton

the beetle held in a horizontal plane.

22 Ratio: head width/pronotal width (hw/pw)

23 Ratio: elytral width/elytral length (ew/el)

24 Ratio: femur length/elytral length (fl/el)

25 Per cent of elytral surface covered by light markings or maculations. This was

estimated in a manner very similar to that used by Gaumer (1977). A series of

specimens representing the range of variation present in C. longilabris and C.

nebraskana was selected. A drawing was made of the left elytron of each

specimen. A polar planimeter was used to determine the percentage of each

elytron covered by the maculations. Subsequently, these drawings were used as

standards of comparison for estimating the percentage to the nearest one of six

categories: 1%, 2%, 5%, 10%, 20%, and 30% (Fig. 7).

26 The configuration of the humeral lunule (hi). I recognized the following six

states of this character (Fig. 8):

0. humeral lunule absent.

1 . one humeral dot present at shoulder of elytron.

2. one subhumeral dot present.

3. both humeral dots present.

4. humeral lunule complete, or nearly so.

5. humeral lunule complete and connected to marginal line.

27 The configuration of the middle band (mb). Specimens were categorized as

being closest to one of the following states of this character (Fig. 9):

0. middle band completely absent.

1 . middle band barely discernible, or in two pieces.

2. middle band present, with angle of bend greater than 45 degrees, and not

touching lateral margin of elytron.

3. band complete, touching lateral margin of elytron and angle of bend

greater than 45 degrees.

4. band present, not touching lateral margin of elytron and elbow less than

45 degrees.

5. middle band complete, touching lateral margin and elbow less than 45

degrees.

6. band complete, touching lateral margin, and marginal line well

developed.

28 Apical lunule character states (al) The following states were recognized in the

degree of development of the apical lunule (Fig. 10):

0. apical lunule entirely absent.

1 . apical lunule consists of a small subapical dot.

2. apical lunule consists of a large subapical dot.

3. apical lunule complete.

4. lunule complete, with dot expanded anteriorly.

5. apical lunule complete and continuous with marginal line

The Cicindela sylvatica Group

67

anteriorly.

30 Color of dorsal surface of elytra (ec)(Table 1)

31 Color of proepistemum (pc)(Table 2)

32 Color of ventral surface of abdomen (vc)(Table 3)

Tables 1-3 contain a vernacular designation of a color condition, the character

state number and corresponding name and number from the ISCC-National Bureau

of Standards Color Charts (Kelly and Judd, 1965). For each of characters 29, 30, and

3 1 a small series of specimens was chosen to represent the range of variation found

in the C. longilabris species complex. These standard specimens were compared

with the ISCC-National Bureau of Standards Color Charts and the corresponding

color name and number were noted. If a specimen did not closely match any one

color, the two or three colors closest to it were noted. Subsequently, each studied

specimen was compared against standard specimens and designated as being closest

to one of the representative color categories.

One difficulty with this method is that many of the colors of tiger beetles are

structural (Shelford, 1917) and have a metallic sheen, whereas those of the standard

color charts are opaque. The color of the dorsal surface of elytra was designated a

single color state. For color of the proepistemum and venter of the abdomen, each

character state was, in many specimens, a mosaic of more than one color, which

added to the problem.

Measurements and character states were taken from adult specimens from 60

localities across the range of the species complex: 12 population samples of C.

nebraskana and 48 of C. longilabris (Fig. 11, Table 4). An effort was made to

choose larger population samples. While only small samples were available from

many localities, they were analyzed with the knowledge that they may have been

atypical because of biased sampling by collectors. Collectors may take

disproportionately large numbers of unusual color morphs in preference to common

morphs, especially of the common species.

Numerical analysis of morphometric data. — Sexual dimorphism was examined

by comparing males and females for each of the variables using a one-way analysis

of variance (ANOVA) procedure as described by Kim and Kohout (1975). With the

probability of a type one error set at 0.05, significant sexual dimorphism was found

in both C. nebraskana and C. longilabris . Females of both groups showed

significantly larger measurements in head width, labrum width, labrum length,

pronotal width and length, and elytral width and length, suggesting that females of

both species are significantly larger in overall size. In all subsequent analyses of

morphometric data males and females were treated separately. All analyses were

performed with the use of a Vax 11/780 computer.

Discriminant analyses based on the above series of measurements were

performed to investigate differences among population samples. The discriminant

procedure used was described by Klecka (1975) and was taken from the Statistical

Quaest. Ent., 1988, 24 (1)

68

Spanton

Package for the Social Sciences for Vax.VMS, Version M, Release 8.1, May 1,

1981.

A linear discriminant function is a combination of character scores which

discriminates between groups much better than one character taken singly (Sneath

and Sokal, 1973). For this reason significance levels of 0.05 or 0.01 are not useful in

this type of analysis, since they identify taxonomically insignificant differences.

Some neighbouring populations which appeared to be very similar were

significantly different beyond the 0.01 level in these analyses. A significance level

of 0.001 was selected as the minimum requirement indicative of a taxon. However,

no taxonomic decisions were based entirely on the evidence of discriminant

analysis.

For seemingly important population groupings established by discriminant

analyses, additional one-way analyses of variance were used (Kim and Kohout,

1975) to investigate the statistical differences in those variables which scored

highest in the discriminant functions.

Analysis of color and pattern of markings. — Color variation was demonstrated

with the use of pie graphs on distribution maps (Figs. 12-19) in a manner similar to

that used by Freitag (1965), Willis (1967), and Gaumer (1977). Mean states of

additional characters for each population sample are summarized in Table 1 1 and

are indicated with symbols on maps in a way similar to that used by Goulet and

Baum (1981, 1982).

Dissections of male and female genitalia. — Using standard methods, genitalia of

male and female specimens of many populations of C. nebraskana and C.

longilabris were examined for structures of taxonomic importance. Males and

females of six Palearctic species (not more than two or three specimens of each

gender, per species) were dissected to aid in determining relationships among the

species of the C. sylvatica group. In all 170 genitalic dissections were performed.

Soil associations. — Collecting localities from specimen label data were located

on national scale and state soil maps to seek relationships between the distribution

of dominant soil types and the distribution of the different forms of the C. sylvatica

group in North America. Conversions between the United States and Canadian

systems of soil classification were made with tables provided in Clayton et al.

(1977). Descriptions of soil types were followed in Clayton et al. (1977) for the

Canadian classification and in Soil Survey Staff (1960, 1967) for the American

system.

Dates of collection were used to plot histograms of frequency of capture versus

date to investigate seasonality in adults of C. longilabris and C. nebraskana, and

other label data were used to compile distribution lists and to plot distribution maps

of the two North American species.

Field methods. — Adult specimens were collected with an insect net, killed in an

ethyl acetate jar and either pinned the same day or transferred to 70% ethanol for

temporary storage. Larvae were collected in one of two ways. The “lie in wait”

The Cicindela sylvatica Group

69

method involved waiting near the mouth of an open burrow until the larva appeared

near the surface, at which time a small shovel was driven at an angle under the larva,

cutting off its escape route. The other method was to dig out the larva with a hand

trowel. As larvae of C. longilabris were found at depths to 60 cm, it was helpful to

insert a long piece of flexible grass into the burrow until the larva was felt at the

bottom. A hole was then dug beside the stalk of grass until the larva was

encountered. Larvae were either preserved directly in 70% ethanol or were placed

live in a glass vial with a small amount of soil for transport to the laboratory.

Samples of soil were collected from larval sites and transported to the laboratory

where they were thoroughly dried in an oven and if necessary, rolled gently with a

rolling pin to break up any aggregations which formed during drying. Each sample

was then shaken through a standard sieve series with mesh sizes of 2.0 mm, 0.50

mm, 0.25 mm, 0.125 mm, 0.063 mm and 0.037 mm to determine the distribution of

soil particle sizes.

Study sites. — One site, located in the Thunder Bay district of Ontario near the

east bay of Dog Lake, 1-5 km W of highway 527 and 50 km N of highway 17, was

used primarily as a source of specimens, both for mating experiments and for

rearing larvae. The species C. longilabris was found along logging roads that extend

through second growth forest in an area of sandy soil where the dominant trees were

Trembling Aspen (Populus tremuloides Michx.) and Jack Pine (Finns banksiana

Lamb.) and through an area of slightly more gravelly soil with Trembling Aspen,

Jack Pine and Spruce (Picea sp.). Ground cover varied from absent on the road

surface to patchily distributed mosses and lichens, grasses, wild strawberry

(Fragaria virginiana Duch.) and leaf litter.

The other study site, at Stanley Hill Cemetery on highway 17, 16 km W of

Thunder Bay, Ontario, included some of the grounds of the cemetery, a small sandy

area along the edge of forested land across the highway from the cemetery, and part

of a pasture bordering the cemetery on the east side. This was an area of sandy soil

with vegetation cover ranging from mixed forest of predominantly Trembling Aspen

and Jack Pine to old field habitat and bare soil.

A bare road surface extended along the edge of the field. The south end was in

close proximity to Jack Pine trees, and the north end extended into an open field

habitat. This road was marked at intervals of approximately 5 meters for a distance

of 800 meters.

A multiple mark and recapture study was conducted at this site through the

summer of 1982 to investigate mobility and relative abundance of the adult beetles.

Beetles were captured with a net. Marks were placed on the elytra in the form of

small dots of enamel model paint, which has been used successfully in mark and

recapture studies of tiger beetles (Willis, 1974; Palmer, 1976; Kaulbars, 1982), and

numerous other insects (Southwood, 1978). Using six locations on each elytron

where spots could be placed, and six colors of paint, 468 different combinations of

marks were possible with no individual bearing more than two spots. Each captured

Quaest. Ent., 1988, 24 (1)

70

Spanton

beetle was marked and released at the point of capture and its sex, location and date

of capture were recorded. Each capture session consisted of one survey from one

end of the 800 meters of marked road surface to the other, and back again. On days

when few beetles were captured this took approximately two hours; when beetles

were numerous a capture session was limited to three hours. Capture/recapture

sessions throughout the summer were conducted mainly on sunny, warm days.

The chronology of larval development was studied at the Stanley Hill site by

marking burrows. A golf tee numbered with a waterproof ink marker was placed 2

cm north of each burrow, and the developmental stage inferrred from the size of the

head and pronotum and the diameter of the burrow. Both the size of head and

pronotum and the diameter of the burrow show three discrete size categories

corresponding to the three larval instars. Burrows were checked at intervals of a few

days to a week throughout the summer. Newly found burrows were marked and

each burrow was noted as open or closed, and if open the instar was recorded.

In the middle to latter half of the summer, 1st instar larval burrows appeared in

numbers too large for all to be marked with golf tees. At this time visual counts

were made of open burrows in each stage of development at intervals of a few days

to a week to gather information on the seasonality of the larval stages.

Rearing techniques. — Live adults were kept in glass terraria approximately 15

cm X 40 cm X 25 cm in size, the bottom of which were covered to a depth of

approximately three cm at one end to six cm at the other end with soil taken from

the site where the beetles were captured. A petri dish filled to the level of the rim

with soil was placed in the shallow end of each terrarium and periodically filled to

overflowing with water. In this way soil moisture available to the beetles ranged

from wet in and around the petri dish at one end of their enclosure to dry at the other

end. Two to three adults of both sexes were placed in each terrarium. Initially,

mortality from cannibalism was high until clumps of mosses, grasses and leaf litter

from the beetles’ natural habitat were added to the terraria. The tiger beetles

immediately used the leaf litter for cover or dug shallow burrows under the clumps

of grass or moss. Aggressive encounters and cannibalism were greatly reduced after

these modifications. The beetles were fed primarily flour beetles {Tribolium spp.)

supplemented occasionally with assorted arthropods collected with a sweep net.

First instar larvae which appeared in the terraria subsequent to mating and

oviposition, and other larvae dug from the field were reared in glass tubes

approximately 2 cm in diameter by 30 cm long in a manner similar to that described

by Palmer (1979). The rearing tubes were plugged at the bottom with wet cotton

balls or crumpled paper towelling and filled to a depth of 20 - 25 cm with soil from

the site where the larvae were collected, or for those produced in the laboratory,

where their parents were collected. The rearing tubes were placed on end in a plastic

bucket and the soil kept slightly moist with water added to the bucket and

occasionally applied to the surface with a plant sprayer. Soil moisture was regulated

to minimize mould growth. In a few instances mould developed and specimens were

The Cicindela sylvatica Group

71

lost. First instar larvae of Cicindela were fed early instar larvae of Tribolium spp.

and second and third instar tiger beetle larvae were fed late instar larvae, pupae and

adult Tribolium .

In the laboratory the ambient temperature was approximately 20°C, with

fluorescent lights which were generally on during the day and off at night. No

attempt was made to approximate naturally occurring photoperiod, temperature, or

humidity. Palmer (1979) noted that fecundity of some species of tiger beetles is

reduced under laboratory conditions and suggested that temperature may be

important in egg production. Reproductive success may have been increased if

laboratory conditions more closely approximated the warm daytime temperatures,

cool nights and long photoperiod typical of summer in their natural environment.

Criteria for species and subspecies. — Species concepts have been discussed by

Simpson (1961), Mayr (1969) and Wiley (1981), among many others. The use of a

subspecific category has in the past been controversial among zoologists, with

workers such as Edwards (1954), Parkes (1955), and Smith and White (1965)

arguing in favour of its use, and Wilson and Brown (1953), Gosline (1954), Hubble

(1954) and Owen (1963), among others, opposed to its use. More recently, it seems

that many ornithologists (Mayr, 1982; Parkes, 1982; Gill, 1982; Storer, 1982;

Barrowclough, 1982; Lanyon, 1982; Johnson, 1982; Zusi, 1982; Monroe, 1982;

O’Neill, 1982; Phillips, 1982) feel that the subspecific category should be retained

in zoological nomenclature and that trinomina are at least of practical value in some

instances. Virtually all of these more recent essays express some reservations

regarding the use of subspecific names. Taken to extremes, the formal naming of

minutely different populations has little meaning biologically, and merely confuses

the nomenclature. This has certainly been a problem in tiger beetle taxonomy when

in the past, many subspecies and species names have been formally proposed, based

on one or very few variant specimens. Other concerns regarding subspecies include,

(1), the tendency for different characters to show discordant patterns of geographic

variation; (2), the occurrence of similar or phenotypically indistinguishable

populations in geographically separated areas (the “polytopic subspecies” of Mayr,

1969); (3), that an artificial compartmentalization of our concept of the pattern of

variation in a species tends to obscure geographic variation within subspecies; (4),

the subjectivity in the degree of distinction required by different workers to justify

the application of a formal name.

Advantages of the use of a subspecies category are many. Subspecies names are

important sorting devices for curators of collections, even if this is strictly a clerical

convenience. Trinomina are useful in information retrieval, as much information

regarding genetics, population dynamics, feeding behavior, and other aspects of an

organism’s biology have been published on certain subspecies of polytypic species.

Ironically, as Mayr (1982) points out, searching of collections for possible new

subspecies has had a heuristic benefit in elucidating patterns of geographic variation

which might have otherwise gone unnoticed. It is now largely believed, as opined by

Quaest. Ent., 1988, 24 (1)

72

Spanton

Barrowclough (1982), that looking for subspecies to describe is an innappropriate

type of taxonomic work, and subspecies should be only the occasional biproduct of

studies of geographic variation. Many ecologists have been alerted by subspecific

names to subjects for study of adaptation to local selection pressures in widespread

species. Such workers frequently find it convenient to have formal names to apply to

the subjects of their investigations. Subspecies can be of interest in the context of

historical biogeography in that they can (although they need not necessarily) be

units of evolution. Some geographically isolated subspecies may represent

“incipient species”: should the selection pressures which brought about their

subspecies-level differentiation continue for a sufficient period of time, speciation

may occur.

As Monroe (1982) points out, Mayr’s (1969) definition that a “subspecies is an

aggregate of phenotypically similar populations of a species, inhabiting a

geographic subdivision of the range of a species, and differing taxonomically from

other populations of the species”, is less than precise. The phrase “differing

taxonomically” implies a degree of subjectivity, but, I believe it is still a good

definition. After all, there are many instances where the biological species concept

breaks down and must be applied in an arbitrary manner, yet no one would seriously

suggest that we scrap the species concept! Subjectivity cannot be completely

removed from taxonomic work. The judicious application of subspecies names can

be appropriate where studies of geographical variation show reasonable grounds for

recognizing subsets of a species.

In the absence of direct breeding evidence, relationships between phena in this

study were inferred, based on holomorphological evidence with emphasis on adult

structure and supplemented with some ecological and distributional data. Sympatric

forms which show little or no intergradation in at least one character are considered

specifically distinct. Allopatric forms which intergrade clinally over a fairly wide

zone of contact are considered subspecies if the forms are sufficiently different

structurally. Allopatric populations which are completely isolated geographically are

considered subspecies if they differ only in color or color pattern.

STRUCTURAL AND ECOLOGICAL FEATURES OF THE NEARCTIC TAXA

OF THE C. SYLVATICA GROUP

This section includes: (1), an analysis of adult features, including discriminant

analyses of morphometric data, and characterization of the pattern of variation of

qualitative characters of color and color pattern; (2), a description of immature

stages and life history of C. longilabris", and (3), a treatment of soil associations of

the two species.

The Cicindela sylvatica Group

73

Quaest. Ent., 1988, 24 (1)

Fig. 11. Population samples used in numerical and color analysis (Table 4).

74

Spanton

TABLE 4. Population samples of C. longilabhs and C. nebraskana used in

numerical and color analyses (Fig. 1 1).

N

The Cicindela sylvatica Group

75

Table 4 (continued)

Species Code Locality Males

(continued on next page)

Females

19

3

7

34

12

5

7

24

10

22

6

7

6

10

16

23

6

11

13

10

11

15

11

10

1

6

Quaest. Ent., 1988, 24 (1)

76

Spanton

Table 4

Species

(continued)

N

TOTAL

647

710

TABLE 5. Canonical discriminant functions evaluated at group centroids for discriminant analyses of C. longilabris and C. nebraskana.

The Cicindela sylvatica Group

77

|Jh

Uh

Quaest. Ent., 1988, 24(1)

(continued on next page)

Test Sex Group Function 1 Sign. Function 2 Sign. Function 3 Sign.

78

Spanton

w tii

§ §

■i I

g 2

I §

-i I

g 2

-fe ■?

G G ^

§ -s

■s I

^ -S

c: c!o

-ci s;

o

§ -s

-s ■§

C Oo

-Cs C

c —

G ^ G ^

Uh

PU

VO

The Cicindela sylvatica Group

79

TABLE 6. Test 1, F statistics* and associated significance levels^ between groups of

males of C. longilabris from NFl, NF2, QB2 and QB3 populations.

Each F statistic has 1 1 and 5 1 degrees of freedom.

TABLE 7. Test 2, F statistics* and associated significance levels^ between groups of

females of C. longilabris from NFl, NF2, QB2 and QB3 populations.

Each F statistic has 1 1 and 52 degrees of freedom.

Quaest. Ent., 1988, 24(1)

80

Spanton

TABLE 8. Test 3, F statistics* and associated significance levels^ between groups of

males from the combined population samples of C. longilabris from the Rocky

Mountain States, Pacific Region and the Boreal Region.

Each F statistic has 12 and 431 degrees of freedom.

GROUP

Pacific Region

Rocky Mtn. Region

BOREAL ZONE

18.885*

0.0000+

12.662

0.0000

PACIFIC REGION

10.140

0.0000

TABLE 9. Test 4, F statistics* and associated significance levels^ between groups of

females from the combined population samples of C. longilabris from the Rocky

Mountain States, Pacific Region and the Boreal Region.

Each F statistic has 12 and 464 degrees of freedom.

GROUP BOREAL ZONE PACIFIC REGION

Pacific Region 22.872*

0.0000+

Rocky Mtn. Region 15.807

0.0000

16.323

0.0000

The Cicindela sylvatica Group

81

Analysis of adult features

Discriminant analysis. — Figure 1 1 and Table 4 indicate the population samples

used in these analyses. Table 5 indicates the canonical discriminant functions

derived in each of the tests and Tables 6 to 9 present F statistics and associated

significance levels for comparisons among groups. Table 10 indicates variables

selected by the discriminant program in each of these tests.

Discriminant analysis tests 1 and 2 (Tables 5, 6 and 7) were conducted to

investigate variation among population samples from Gander, Nfld. (NFl), Harmon

Field, Nfld.(NF2), Thunder River, Quebec (QB2) and Mont Albert, Quebec (QB3)

(Fig. 11). This was done to seek significant metric differences between

Newfoundland and mainland populations, to corroborate Leng’s (1918) designation

of the formal name C. /. novaterrae, applied to the island populations based on an

increased frequency of green colored individuals. In test 1 for males, two

discriminant functions were derived which dealt with a significant amount of

variation (Table 5). Function 1 scored the two Quebec populations close together,

with NF2 having an intermediate value and NFl scored farthest from the Quebec

populations. Function 2 scored NFl and QB3 closer together near the mid range of

the scale, with QB3 and NF2 farthest apart at either end of the range of values. In

both functions, measurements of the labrum and pronotum contributed most to the

functions (Table 10). Males from Newfoundland were found to have a larger labrum

and a larger pronotum. These differences seem to reflect a difference in body size,

those specimens on the island being larger, on average. It is a commonly occurring

phenomenon, for island populations of animals to be different in size. Freitag (1965)

showed that specimens of Cicindela oregona on the Queen Charlotte Islands are

larger, on average than mainland specimens, and Lindroth (1963: 88) indicated that

a number of species of carabid beetles are noticeably larger in body size on the

island of Newfoundland than are their mainland populations.

The F statistics and associated significance levels between groups (Table 6)

indicate a discordant pattern of variation. Males from NFl are different from QB2

and QB3 at a significance level beyond 0.0001, whereas NF2 males are not

significantly different from QB3 males at a 0.01 level. NFl males and NF2 males

differ from each other significantly beyond the 0.0001 level. This greater statistical

difference between populations from Newfoundland than between NF2 and QB3

seem to refute subspecific status for Newfoundland populations, based on

morphometric characters.

Test 2 (Table 5), with female specimens also produced two functions with a

significant amount of variation (at or beyond the 0.0001 level). Function 1 separates

most strongly between NFl and QB2 at opposite ends of the range of values, with

NF2 and QB3 having very similar scores, intermediate on the scale. Function 2

scores QB2, NFl and NF2 close together with QB3 distinctly separate from the

other three. Femur length and tibia length are the variables contributing the greatest

amount of variation to Functions 1 and 2 for females of these four populations. The

Quaest. Ent., 1988, 24 (1)

82

Spanton

multivariate F statistics and associated significance levels for test 2 (Table 7)

indicate that females of NF2 do not differ from those of QB2 and QB3 significantly

at the 0.001 level. The F statistics and significance level between NFl and NF2 is

comparable to that between the NF2 and Quebec populations. These differences

based on morphometric characters do not warrant subspecific distinction between

the populations on Newfoundland and those of the mainland.

Discriminant tests 3 and 4 compared three large groupings of population samples

of C. longilabris: (1), grouped populations of the Rocky Mountain states from

Montana to Arizona and New Mexico (MT2, SDl, WY3, COl, C02, UT2, UT3,

AZl, AZ2, NMl and NM2)(Table 4, Fig. 11); (2), grouped populations of the

Cascade and Sierra Nevada Mountains from southern British Columbia to east

central California (BC3, WA2, WA3, ORl, OR2, CA2, CA3, CA4)(Table 4, Fig.

11); and (3), the nominal form from across the boreal zone in its broadest sense in

the northern part of the continent (all remaining population samples. Table 4, Fig.

11).

In test 3, comparing male specimens, two functions were derived with a

significant amount of variation (significant beyond the 0.0001 level. Table 5). Table

8 for test 3 indicates multivariate F figures between any two of these groups

significant beyond the 0.0001 level. Test 4, comparing female specimens, produced

very similar results. Two highly significant discriminant functions were produced

(Table 5). One function set the Pacific form apart from the other two and the second

function separated the Rocky Mountain group from the Pacific and Boreal forms.

Table 9 shows that any two-group comparison among these three, has an F statistic

that is significant beyond the 0.0001 level. Both male and female specimens show

highly significant statistical differences among these groups, beyond the minimum

significance level (0.0001) here chosen for taxonomic purposes.

The variables contributing most to the discrimination among these three groups

were pronotal width (pw), pronotal length (pi), per cent of elytral surface covered

with maculations (prct), head width (hw), length of mesothoracic tibia (tl) and

elytral width (ew)(Table 10). A one-way analysis of variance was performed to

investigate the statistical differences in each of these measurements. For males, all

three groups were statistically different from each other in pw (F=27.958, p=0.0001,

sign, at 0.05 by Scheffe’s procedure) and hw (F=25.710, p=0.001, sign, at 0.05 by

Scheffe’s procedure). For both of these variables males of the Boreal group

measured the largest and the Pacific group measured smallest. Similarly for females,

a one-way analysis of variance (ANOVA) showed that each of the groups differs

from the others based on the variables pw (F=57.793, p=0.0001, sign, at 0.05 by

Scheffe’s procedure) and hw (F=52.201, p=0.0001, sign, at 0.05 by Scheffe’s

procedure). For both variables, females of the Boreal form had the largest average

measurement and the Pacific form was the smallest of the three. These statistics

probably reflect the overall size differences, with the nominate form largest in body

size, the Rocky Mountain form smaller, and the Pacific form smallest.

The Cicindela sylvatica Group

83

The Boreal form has a proportionately shorter, wider pronotum than the other

two groups evidenced by the statistical difference in the variable pl/pw (ANOVA,

Males: F=42.165, p=0.0001, sign, at 0.05 by Scheffe’s procedure; Females:

F=36.908, p=0.0001, sign, at 0.005 by Scheffe’s procedure).

Per cent of the elytral surface covered with maculations was another

discriminating variable which shows statistical differences among the groups. An

ANOVA of male specimens confirmed that the Boreal form is less maculate than

the other two groups (F=43.175, p=0.0001, sign, at 0.05 by Scheffe’s procedure).

The same test with female specimens also confirmed that the Boreal form is least

maculate, the Rocky Mountain group more so, and the Pacific group most maculate,

on average (F=55.577, p=0.0001, sign, at 0.05 by Scheffe’s procedure).

Discriminant tests 5 and 6 compared population samples of C. nebraskana from

east of the Rocky Mountains with those west of the continental divide. Leffler

(1979) in studying tiger beetles of the Pacific Northwest recognized two subspecies,

C. n. nebraskana from east of the divide and C. n. chamberlaini from west of the

divide, based primarily on a statistical difference in head width: the eastern

populations having narrowest heads with intermediate populations occurring near

the divide in eastern Idaho and northwestern Wyoming. Test 5 (Table 5) for male

specimens, produced a discriminant function which separated between the eastern

and western populations significant beyond the 0.0001 level. The variables

contributing most to the discriminant function were, in order of importance, fl and el

(Table 10). Hw was not selected in this analysis. Test 6 using female specimens also

separated between the groups, with a multivariate F ratio of 14.166, significant

beyond the 0.0001 level. In this discriminant function the variables contributing

most to the variation were, in order of importance, fl and hw.

Using an ANOVA, with the probability of a type one error set at 0.01, for both

males and females, the western group of C. nebraskana had significantly longer

mesofemora, on average and larger elytra than the eastern populations. No

differences in other measurements significant at the 0.05 level were found between

the eastern and western populations of C. nebraskana.

Discriminant tests 7 and 8 compared C. longilabris with C. nebraskana. As

Table 5 indicates, for test 7, comparing male specimens, a discriminant function was

derived which separated the two groups very well. The multivariate F ratio was

39.988, significant beyond the 0.0001 level. The variables which contributed most

to the discriminant functions were 11, Iw, 11/lw, prct, hw/pw, Icol, and ls2 (Table 10).

The discriminant function derived in test 8 between females of the two species,

was also highly significant (Table 5). The multivariate F ratio was 50.403,

significant beyond the 0.0001 level. The variables most effective in separating the

females of the forms were prct, Iw, Icol, 11/lw, pw, ls2, and osr (Table 10).

In summary, using an ANOVA with probability of a type one error at 0.001,

males and females of C. longilabris, as compared to those of C. nebraskana, have

significantly longer and wider labra, have proportionately longer labra relative to

Quaest.Ent., 1988, 24(1)

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labral width, are more maculate, and have proportionately wider heads, in relation to

pronotal width. Males and females of C. nebraskana have significantly more

non-sensory setae on the scape of the antenna (Fig. 3) than C. longilabris, but

individuals could not be identified on that basis.

Color and pattern of markings. — Figure 12 presents the frequency of occurrence

of three categories of labrum color for females of each population of C. longilabris.

In most samples, all females had light colored labra, in some populations a

significant fraction of females exhibited labra intermediate in color, mottled, or

darkened along the outside edge and midrib. In a small number of population

samples a small fraction of females had dark colored labra. A geographic pattern to

the variation in this character in C. longilabris was not identified.

The frequency of occurrence of color character states of the female labrum is

highly variable among populations of C. nebraskana (Fig. 13). In samples from

southern Alberta (ABl, AB2)(Table 4, Fig. 11), and eastern California (CAl), the

dark colored labrum predominates. In the Wyoming populations (WYl, WY2) most

females have light colored labra, and in other populations studied the three character

states are represented in varied frequencies between the two extremes. There

appears to be no pattern to the variation in this character in C. nebraskana. Overall,

among C. nebraskana females 44.8% had light colored labra, 30.6% were

intermediate in color, and 24.6 % were black (total, n=134). In contrast, among C.

longilabris females, labra were light in 90.3%, intermediate in character in 8.2% and

black in color in 1.5% of specimens (n=586).

Among male specimens of C. nebraskana, 82.6% had light labra, 15.6%

intermediate, and 1.8% black (n=109), and among C. longilabris males the

corresponding figures were 97.6% light labra, 1.8% intermediate, and 0.6% dark

(n=538).

Figure 14 shows the frequency of occurrence of colors of the proepistemum of

C. longilabris. This character varies across the range of the species, but in most

populations, specimens with green and black, green, green and blue, or bronze and

green colored proepistema (character states 2-7, Table 2), make up the largest

fraction of the samples. Figure 15 shows the frequency of proepistemal color states

in population samples of C. nebraskana. In this species the proepistema in most of

the populations are black in color, with notable exceptions being the Wyoming

populations (WYl and WY2) and the northern Utah population (UTl), in each of

which specimens with bronze colored proepistema (character states 6, 7, Fig. 15)

make up a large fraction of the sample.

Figure 16 presents frequency of occurrence of colors on the ventral surface of the

abdomen in C. longilabris This character varies throughout the species, but metallic

greens and mixtures of metallic greens, blues and bronze color (character states 3, 4,

6, Table 3) predominate in most populations.

In contrast. Figure 17 shows the frequency of occurrence of various colors of the

venter of the abdomen in specimens of C. nebraskana. In most population samples.

TABLE 10. Variables (as described in the Materials and Methods section) selected in discriminant

analysis of Cioindela longilabris/nebraskana. The 'X* indicates variables that failed

The Cicindela sylvatica Group

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TABLE 11. Weighted mean values of characters prct, hi, mb, al for samples of C.

nebraskana and C. longilabris. Characters are defined in the Materials and

Methods section and indicated in Figs. 4, 5, 6, and 7.

(continued on next page)

The Cicindela sylvatica Group 97

Table 1 1 (continued)

black (character state 1, Table 3) is the ventral color in the majority of specimens,

with blue and purple (character state 5) and bronze (states 6, 7), comprising large

fractions of some of the samples. The UTl sample is peculiar in having a large

number of specimens with black and green abdominal coloring (character state 2,

Fig. 17).

Figure 18 shows the frequency of occurrence of colors of the elytra of C.

longilabris. This character is highly variable throughout the range of the species.

The blue-green and dark blue color (states 6 and 7, Table 1) occurred so rarely that

they did not appear as a fraction of any of the sampled populations.

Green specimens predominate in the Newfoundland populations of C. longilabris

(Fig. 18). The mainland specimens across eastern North America are almost all

black or dark brown in dorsal elytral coloration. In the western part of the continent

elytral color is more variable. In the Rocky Mountains of the western United States

dark brown, bronze, and olive green (character states 2, 3, and 4) specimens

predominate and in the Pacific region from California to Washington State, bright

Quaest. Ent., 1988, 24 (1)

98

Spanton

green colored specimens are most numerous. In C. nebraskana (Fig. 19), black

colored specimens are in the overwhelming majority with dark brown and bronze

elytra occurring in small fractions of the populations.

Figures 20 and 21 show the average percentage of the elytral surface covered

with maculations (Fig. 7), in C. longilabris and C. nebraskana, respectively. In the

former species (Fig. 20), most of the specimens of the Boreal zone have an average

of between 1 and 5 per cent of their elytral surface covered with maculations. This

percentage increases greatly in specimens of some populations of the Pacific and

Rocky Mountain regions of the United States.

In C. nebraskana (Fig. 21), most of the specimens have an average of less than

one per cent of the elytral surface area covered with maculations. The exceptions are

the IDl and WY2 samples which have an average of 1.2 and 1.6 per cent,

respectively, of their elytral surfaces covered with light markings.

Figures 22-27 and Table 1 1 present weighted mean values of characters of the

elytral markings for population samples of C. longilabris and C. nebraskana. Figure

22 presents weighted mean character values of the humeral lunule (Figs. 6, 8) for

population samples of C. longilabris. In most populations of this species the

humeral lunule is present as one or two distinct dots or as a complete lunule (Fig. 8).

Figure 23 presents the weighted mean character values of the humeral lunule for

samples of C. nebraskana. In all samples of this species, the humeral lunule is

lacking from the majority of specimens.

Figure 24 presents the weighted mean character values of the middle band (Figs.

6, 8) for C. longilabris. In most population samples from the Boreal zone, the

middle band is either indistinctly present (state 1, Fig. 9) or complete, but quite thin

(state 2, Fig. 9). The middle band is more developed in the southern Rocky

Mountain region and in the Pacific coast states. Figure 25 presents weighted mean

character values of the middle band (Fig. 9) for samples of C. nebraskana. In a

majority of the populations the middle band is lacking completely in most

specimens. In BCI, WAl, IDl, WY2 and MBl (Fig. 11) the middle band is either

incompletely present or is present as a thin line (states 1 and 2, Fig. 9) in most

specimens.

Figure 26 presents weighted mean character values of the apical lunule of

samples of C. longilabris. In some populations in the Atlantic region of Canada

(QBl, NF2, NBl, NSl, Fig. 11), the apical lunule is typically a dot of varying size

(states 1-2, Fig. 10). In most of the samples from across the Boreal zone the apical

lunule is missing (state 0, Fig. 10), or is present as a dot of varying size (states 1, 2,

Fig. 10). The apical lunule is more developed in many populations of the southern

Rocky Mountains and Pacific regions, but is quite variable between populations.

Figure 27 shows weighted mean character values for the apical lunule of samples

of C. nebraskana In all sampled populations of this species the apical lunule is

lacking from the majority of specimens but present as a small dot (character state 1 ,

Fig. 1 1) in a few specimens.

V.

The Cicindela sylvatica Group

99

Larvae and Life History Data

Larvae of Cicindela longilabris. — Leffler (1979) described the third instar larva

of C. nebraskana and a single second instar larval specimen of C. longilabris from

an area of hybridization between the Rocky Mountain form, C. /. laurentii and the

Pacific Coast form, C. 1. perviridis.

As descriptions of larvae have in the past been based on the third instar

(Hamilton, 1925; Willis, 1967), and some characters in these descriptions vary

between instars of a species, the following description is based on the third instar,

with differences among the three instars noted, where applicable. The format used

for the descriptions of the larvae follows that of Hamilton (1925) and Willis (1967),

to facilitate comparisons among species.

Material. — Twelve third instar specimens, twelve second instar specimens, and

five first instar specimens were reared from mating of captive adults of C. /.

longilabris collected near Thunder Bay, Ontario.

Description. — Colon Head, pronotum and clypeus bronze-black with some metallic reflections

varying from green to cupreous to bronzy in some specimens; labrum black or very nearly so in second and

third instars, bronze in first instars; mandibles rufous basally to black in distal portion in second and third

instars, in first instars mandibles are light brown proximally to dark brown distally; antennae dark brown to

black; maxillae rufous to medium brown in second and third instars, light brown in first instars, with apical

segments of palpus and galea dark brown-black in many individuals of all instars; genae rufous posteriorly

to dark brown or black anteriorly; mesonotum grayish brown anteriorly to light brown posteriorly;

metanotum slightly lighter brown than mesonotum; legs light brown in some individuals to dark brown in

others; sclerotized areas on abdomen yellowish brown in second and third instars, almost transparent in first

instars; setae on head and pronotal surface brown, lighter brown in first instars, some white setae around

lateral margins of pronotum; setae on remainder of body brown in second and third instars, very light brown

in first instars. Head: Diameter of stemma 1 approximately equal to diameter of stemma II and

approximately equal to interstemmatal distance (slightly variable); frontoclypeolabral area approximately as

wide as long, or very nearly so (somewhat variable); U-shaped ridge on caudal part of frons with two setae;

antennomere 4 approximately 0.7 times length of antennomere 3, antennomere 3 about 0.6 times as long as

antennomere 2; antennomere 1 approximately 0.8 times length of antennomere 2; antennomere 1 with eight

setae in third instar, five or six setae in second instar, zero setae in first instar; antennomere 2 with eight or

nine setae in third instar, five to seven setae in second instar, two setae in first instar; antennomere 3 with

two setae; antennomere 4 with two or three obvious setae and one to three minute setae in all instars; mesal

edge of maxillary galeomere 1 with one seta in first instar, two setae in second instar and three setae in third

instar; galeomere 2 with five setae; maxillary palpi three segmented, palpomere 2 with two setae, basal and

distal palpomeres lacking setae; labial palpomere 1 with three small ventrodistal spines flanked on each side

by one or two setae in second and third instars, flanking setae lacking in first instars; labial palpomere 2 with

one seta; ligula usually with four setae, five in some individuals; labio-stipites with two long distinct setae

and two minute setae (Figs 28, 29). Pronotum: Cephalolateral angles projecting as far anteriorly as mesal

edge; lateral margins slightly carinate; disc with six to eight setae (seven in most individuals). (Fig. 28).

Abdomen: Sclerotized areas distinct; secondary setae numerous, less than one half as long as primary setae;

ventral elevations of abdominal sternum 9 usually with four distinct setae (occasionally three large and one

smaller); pygopod bearing variable number of primary setae, 16-20 in second and third instars, 12-14 in

first instar; median hooks of segment 5 with two to four setae (three in most individuals) in third instar, two

setae in second instar, one seta in first instar; inner hooks with central spine approximately one third length

of entire hook in third instar, one half of entire length in second instar, and two-thirds length of entire hook

in first instar; two setae at shoulder of inner hook twice as long as spine (Figs. 30-32).

Measurements. — Third instar; Total body length, 12-20 mm (likely to vary

greatly with nutritional state of individual and length of time since last moult);

diameter of stemma I, 0.28-0.30 mm; diameter of stemma II, 0.25-0.29 mm;

Quaest. Ent., 1988, 24 (1)

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The Cicindela sylvatica Group

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1mm

Figs. 28-32. Structures of the third instar larva of C. longilabris longilabris Say. 28, head and pronotum,

dorsal aspect. 29, head, ventral aspect. 30, ninth abdominal sternum. 3 1 , pygopod, dorsal aspect. 32, dorsum

of fifth abdominal segment. Legend: an, antenna; fcla, frontoclypeolabral area; ga, galea; ge, genae; ih, inner

hook; li, ligula; Ip, labial palpus; Ir, labrum; Is, labio-stipites; mb, mandible; mh, median hook; mx, maxilla;

mxp, maxillary palpus; pn, pronotum; stl, stemma one; st2, stemma two; st3, stemma three.

The Cicindela sylvatica Group

107

Quaest. Ent., 1988, 24 (1)

duration larval stages varies geographically and from year to year.

108

Spanton

distance between stemma I and II, 0.26-0.33mm; length of frontoclypeolabral area

1.8-1. 9 mm; length of pronotum, 2.00-2.22 mm; width of pronotum 3.44-3.61 mm.

Second instar: Total length of body, 10-14 mm; length of pronotum, 1.26-1.40 mm;

width of pronotum, 2.04-2.36 mm. First instar: Total length 4.7-7. 1 mm; length of

pronotum, 0.78-0.84 mm; width of pronotum 1.34—1.46 mm.

The three larval instars differ from each other in the following:

1 Size: The size of head and pronotum appears to occur in distinct size classes, as

is evident for many other soil inhabiting tiger beetles in the work of Shelford

(1908), Hamilton (1925), Willis (1967), Palmer (1978), Palmer and Gorrick

(1979). The overall body size or abdominal size probably varies with nutritional

state and the length of time the larva has had to grow since its most recent

moult, which has been shown for C. repanda (Palmer and Gorrick, 1979), and

for C . japonica (Hori, 1982).

2 Color of labrum: Because the metallic lustres are structural colors, they vary

somewhat with the age of the individual as was shown by Shelford (1917).

These colors also vary with different preserving fluids used.

3 Number of setae on the median hooks of the 5th abdominal segment differ

between instars. First instars have one, 2nd instars have two, and most 3rd

instars have three but number varies between two and four.

4 Number of setae on the mesal margin of galeomere 1 is one in 1st instars, two

in 2nd instars, and three in 3rd instars, as noted by Leffler (1979).

5 Fewer setae are evident on each of the antennal segments in the earlier instars

as indicated in the above descriptions.

A third instar larva of C. longilabris should run to couplet 18 in Hamilton’s

(1925) key, with a possible ambiguity at couplet 8 where variablility in number of

setae on the median hook of abdominal segment 5 could cause uncertainty. As stated

above, most 3rd instar larvae have three setae on the median hook, but some third

and all earlier instar larvae have a smaller number. From couplet 18 of Hamilton’s

key, the larvae of C. longilabris can be separated from those of C. tranquebarica

and C. silvicola by the following couplets:

1 Pronotal surface with numerous (less than 25) small secondary

setae in addition to primary setae C. tranquebarica

r Pronotal surface with one or two secondary setae 2

2 (T) Pronotal surface with two setae on each side in addition to the

six primary setae; distance between stemmata I and II greater

than diameter of stemma II; Palearctic Region C. silvicola

2' Pronotal surface with one seta on each half in addition to the

six primary setae; distance between stemmata I and II

approximately equal to diameter of stemmata I and II; Nearctic

The Cicindela sylvatica Group

109

Region C. longilabris

Life history. — In studying Cicindela species in the vicinity of Chicago, Shelford

(1908) identified three life cycle patterns (Fig. 33). In a one year cycle such as that

of C. punctulata, eggs are laid in mid-summer and larvae emerge and attain third

instar by fall, hibernate as third instars, pupate the following June, adults emerge in

early July and quickly become sexually mature, mate, oviposit and die within two

months. In a two year cycle such as that typical of C. lepida, the eggs are laid in

mid-summer, attain second instar by fall, hibernate as second instars, moult to third

instars the second summer, hibernate a second winter as third instars, pupate the

following spring, adults emerge early in the third summer, quickly become sexually

mature, mate and die in two or three months. In an alternate two year pattern, such

as that of C. purpurea, the eggs are laid in June, larvae attain third stage by fall,

hibernate the first winter and pupate the following summer. Adults emerge late in

the second summer, hibernate the second winter, emerge the next spring, mate, and

die. In this last pattern larval life lasts approximately 13 months, and the adult stage

lasts 12 to 13 months.

Shelford (1908) indicated that temperature, moisture, and food influence the

duration of the larval stadia . Some of the species, which Shelford had found to have

a two year cycle, as in the third pattern mentioned above, were found by Griddle

(1910) to have a three year cycle farther north in Manitoba, where the larval life was

prolonged over another winter. This is probably caused by shorter summer seasons

which limit total food intake and delay progress through the larval stages. The work

of Palmer and Gorrick (1979) and Hori (1982) indicates that the larvae of tiger

beetles must attain a threshold body mass in each instar before moulting to the next

stage can occur.

No studies of the chronology of C. longilabris or C. nebraskana through their

immature stages have been published and information regarding adult seasonality of

these species is limited. Leffler’s (1979) observations of the unworn appearance of

the elytra of adult C. longilabris perviridis collected in late summer, suggest that the

adults emerge late in summer, overwinter and become sexually mature the following

spring and, based on adult specimens collected in September which appeared

teneral, he notes the same pattern of adult seasonality for C. nebraskana. Dunn

(1978) provided a frequency histogram of C. longilabris specimens captured in New

Hampshire which showed that adults were most frequently collected during the

month of August. My data concerning the adult seasonality of these two species is

consistent with the findings of Dunn (1978) and Leffler (1979).

Figure 34 indicates the total number of adult C. longilabris taken in each capture

session at the Stanley Hill study site in 1982. Absolute population size was not

estimated, but relative abundance at different times of the season was observed. June

5, 15, and September 2 were cool, overcast and rainy days when few beetles were

active. The overall curve is bimodal. The number of specimens captured in the

Quaest. Ent., 1988, 24 (1)

110

Spanton

August 24 session does not adequately represent the abundance of C. longilabris on

that date. The technique used to mark and record the location and date of capture of

each individual required a handling time which limited the total number of

specimens which could be processed in one capture session. Specimens were so

numerous on that day, that with each step eight or 10 adult specimens of C.

longilabris could be seen to fly in many directions, and a few times two or three

specimens were captured in one drop of the net. At other times of the season when

beetles were not so numerous the upper limit on the number of individuals which

could be captured, marked, recorded, and released was not approached.

Seventy-six per cent of the individuals captured on August 10 were teneral, 59

per cent of those captured on August 17th were teneral, and 66 per cent of those

captured on August 24th were teneral, suggesting that the majority of specimens

collected at that time of year had recently emerged from their pupae. The fact that

very few of the specimens collected in May appeared teneral supports the hypothesis

that most adults of this species emerge in late summer and overwinter in the imago

stage.

The sex ratio of males to females of all adults of C. longilabris captured in May

1982 was 1.74:1 (n = 85). In June this ratio was 1.22:1 (n = 60), in July 1.57:1 (n =

18), in August 1.17:1 (n = 117) and in September 0.81:1 (n = 38). Freitag (1965) in

examining seasonal changes in sex ratios of C. oregona and C. decimguttata found

that females outnumbered males early in the season with males more numerous than

females late in the season. The above data suggest that the reverse is true in C.

longilabris. However, these ratios may not be indicative of the actual sex ratio in the

population at any point in time. Kaulbars (1982) indicated that males of C. denikei

were more numerous in prime open foraging areas of his study sites during breeding

season, and at any given time a percentage of females were ovipositing in sites

outside of foraging areas. As most of the oviposition occurs in early summer in C.

longilabris, the apparent sex ratios skewed in favour of males at that time of year,

could also be an artifact of such behavioural differences between the genders.

Few mating pairs of adult C. longilabris were observed at the Stanley Hill site

during June and July. No mating pairs were seen after July 19th or any time during

August or September. This is consistent with the hypothesis that adults emerging in

late summer do not mate until the following spring.

Figure 35 illustrates frequency of capture of adult specimens of C. longilabris by

date of collection from the label data of borrowed specimens collected in Canada. A

bimodal pattern is weakly evident with population peaks in June and late August.

There is probably a bias in these figures because insect collectors are usually more

active during the summer months, in temperate climates. This may account for the

relatively large numbers indicated for the month of July at a time of year when the

relative abundance of C. longilabris was seen to decline at the Stanley Hill site (Fig.

34).

The Cicindela sylvatica Group

111

Figure 36 illustrates frequency of capture of adult C. longilabris from label data

of borrowed specimens collected in the Pacific region states of Washington, Oregon

and California and suggests a bimodal pattern of adult seasonal abundance. The

large peak in the first half of July probably reflects the time of year when collectors

are most active, and the second peak in September, at a time of year when collecting

activity is greatly reduced, is probably indicative of a late summer emergence of

adults from the pupal stage. Figure 37 presents the same type of histogram based on

specimens from Utah, Colorado, Arizona and New Mexico. A bimodal frequency

distribution is presented with greatest numbers occurring in early August. The

pattern of seasonality of adults of C. longilabris is similar throughout its range, with

some differences in the timing of late season emergence. In Canada the late season

peak in numbers occurs in the latter half of August (Figs. 34, 35), in the southern

Rocky Mountain region this seems to occur in the early half of August (Fig. 37), and

in the Pacific coast states in early September (Fig. 36).

Figure 38 presents the frequency of capture of C. nebraskana adults by date from

label data of pinned specimens. This species appears to be less abundant during

mid-summer and experiences a population peak in late summer. Thus C. nebraskana

probably has a life cycle in which adults emerge in late summer and overwinter

before mating the following spring and early summer. This hypothesis, of course,

awaits confirmation from field studies of the species.

The marking of larval burrows and subsequent observations of development in

size and condition of the burrows at the Stanley Hill site in the summer of 1982

yielded three patterns of seasonality.

Third instar burrows marked in May and early June all disappeared after

approximately mid June. Specimens either died, had their burrows disturbed, or

closed their burrows, pupated and later emerged as adults. In some instances the

burrows and golf tees marking them were disturbed by off-road motorcycles and

lawn mowers.

Seven third instar larvae dug from burrows and placed in rearing tubes in May

pupated in June and emerged as adults of C. longilabris in late July. These burrows

were closed between the active 3rd instar stage and emergence of the adult for an

average of 46 days, ranging from 39 to 62 days. The specimens were probably not in

the pupal stage all of that time. Actual duration of the pupal stage is not known as it

was not observed directly.

The majority of specimens marked as 2nd instars at the Stanley Hill site in May

and June were not successfully followed through the season, presumably from

mortality. Those followed through the season were found to have attained 3rd instar

by early June. They continued to feed for most of the summer and were still in the

third instar in the third week of September. They presumably would have

overwintered in this stage.

First instar larvae appeared on the second week of July and were found in

increasingly large numbers throughout July. The majority were not followed

Quaest. Ent., 1988, 24 (1)

112

Spanton

through the duration of the summer, probably because of a high mortality rate.

Those which were successfully followed through the summer had attained the 2nd

instar by the first or second week of September and presumably overwintered in the

2nd stage.

One first instar larva found in early May at the Stanley Hill site had probably

overwintered. It was probably an individual which had not acquired sufficient body

mass to moult to 2nd instar before its first winter. Such variant individuals are to be

expected.

These data indicate that the complete life cycle of C. longilabris is three years in

duration. Figure 33 presents the chronology of the three year cycle of C. longilabris

compared with the cycles of other species studied by previous workers (Shelford,

1908; Huie, 1915).

If the chronology of the life cycle were rigid for all individuals, C. longilabris

would consist of three distinct populations each genetically and temporally isolated

from the other two. Many workers (Shelford, 1908; Willis, 1967; Palmer, 1976)

have indicated some variablility in timing of appearance of the life stages in many

species of tiger beetles. Hori (1982), in studying C. japonica in which the

seasonality of the adult stage is similar to that of C. longilabris, indicated that few

adults emerging late in summer achieve sexual maturity quickly and a small amount

of oviposition occurs in late summer and early fall. An intensive study of the

population dynamics of C. longilabris would probably reveal a similar amount of

variability in the timing of life stages. Slow developing larvae as mentioned above,

and a small amount of late season mating would maintain a genetic connection

between year classes.

It is unlikely that the life cycle of C. longilabris is less than three years in

duration anywhere in the geographical range of the species. Even in the southern

part of its range in Arizona, New Mexico and Colorado, the same pattern of adult

seasonality seems to occur (Fig. 37). In such areas these beetles inhabit montane and

subalpine forests at high elevations where the duration and temperature ranges of the

seasons roughly approximate that of the boreal forest zone father north. Possibly the

cycle may take a year longer in the more northern part of its range. Just as some

species of Cicindela which have two year cycles in the Chicago vicinity (Shelford,

1908) were found to have three year cycles in Manitoba (Criddle, 1910), the larvae

of C. longilabris may take an extra year to develop in the northern part of its range

where it approaches tree line and the southern limit of continuous permafrost (Fig.

39), and where the summer season is appreciably shorter. This is, of course,

speculative and field work would have to be carried out in various parts of the range

of the species to test the hypothesis.

Soil Associations

Leffler (1979) contended that edaphic factors are among the most important

limits in defining the habitats of ground-dwelling tiger beetles. The female chooses

TABLE 12. Frequency, and relative frequency (%) , of occurrence of locality records of C. longilabvis and C. nebraskana on

dominant soil types as indicated by soil maps (see text for details).

C. longilabvis C. nebrashma

Eastern North Prairie North-western Pacific Cordilleran Summary Surrenary

America Provinces North America States States All Areas

The Cicindela sylvatica Group

113

Quaest. Ent., 1988, 24 (1)

Borderline Cases 9 (2.8) 3 (3-9) ^ (9-5) ^ (3. A) 10 (5) 30 {k) 18 (5.8)

TABLE 13. Characters and character states classified phy logenet i ca 11 y , and their distribution

among the species of the C. sylvatioa group.

Species and character states

No. Character nebr. long. sylv. gran, jap. gemm. solu.

114

Spanton

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The Cicindela sylvatica Group

115

CO

NUMBER OF INDIVIDUALS CAPTURED

Quaest. Ent., 1988, 24 (1)

Fig. 34. Total catch of Cicindela longilabris Say, Stanley Hill study site, Thunder Bay District, Ontario, in the summer of 1982. A, males; B, females.

NUMBER OF SPECIMENS (TOTAL = 909)

116

Spanton

35

Fig. 35. Collection dates of borrowed adult specimens of Cicindela longilabris Say , collected in Canada.

16 -30

NUMBER OF SPECIMENS (TOTAL = 507)

The Cicindela sylvatica Group

1

36

Washington, Oregon and California.

Quaest. Ent., 1988, 24(1)

16-30

118

Spanton

37

Fig. 37. Collection dates of borrowed adult specimens of Cicindela longilabris Say, collected in Colorado,

Utah, Arizona and New Mexico.

16-30

NUMBER OF SPECIMENS (TOTAL = 1722)

The Cicindela sylvatica Group

119

38

Qc >- z ^ O a.

Q. < 3 3 Z) UJ

< S “5 "9 <

Fig. 38. Collection dates of borrowed adult specimens of Cicindela nebraskana Casey.

Quaest. Ent., 1988, 24 (1)

OCT 1-15

120

Spanton

the oviposition site (Shelford, 1907) and the entire larval stage is spent in a burrow

in the soil. Many species are restricted to soils of particular types such as clay, sand,

or alkali-encrusted, among others. Kaulbars (1982) reported that soil type seemed to

be a limiting factor in the distribution of some forms of the C. sexguttata species

group. Leffler (1979) noted a correlation between the length of the second

gonapophyses of the female genitalic armature used in oviposition and the particle

size of soil inhabited, among 15 species of Cicindela. He concluded that species

with long narrow styli are in soils with a high proportion of sand and those with

short stout styli inhabit soils with a high clay content, adding a cautionary note that

recently eclosed individuals should be examined in such a test, as digging activity

can greatly wear down the length of the styli making them much shorter and more

blunt in older individuals. Depending upon how this wearing of the second

gonapophyses affects the sensory abilities of the female to detect differences in soil

qualities such as texture, it raises the question of whether eggs laid by an older

female might be laid in soil of a slightly different texture or at a different depth than

those laid earlier in the life of a given female, and how these factors in turn might

affect the survival chances of the eggs. If such differential survivorship occurs it

could be a strong selective force favouring individuals laid at a certain stage in the

life of an adult female, thus tending to stabilize a given life cycle pattern for a given

species. Intensive field studies of oviposition site selection by females and

subsequent larval survivorship would have to be undertaken to answer these

questions.

Leffler (1979) also concluded that the mean ratio of breadth/length of the styli of

the second gonapophyses is not statistically significantly different between C.

longilabhs (X = 56.75) and C. nebraskana (X = 56.95). According to Leffler’s

(1979) hypothesized relationship between the dimensions of the second

gonapophyses and soil texture, one would not expect a significant difference in the

distribution of particle size of the soils inhabited by these two species. Leffler

(1979) however, indicated that there is such a difference. He stated that five samples

of soil from collecting sites for C. longilabris are largely clay soils, with a mean of

28.6% sand, in contrast with five samples of soil from collecting sites for C.

nebraskana which are all sandy clay with a mean of 50% sand.

My data are more consistent with Leffler’s (1979) indicated similarities between

the dimensions of the 2nd gonapophyses of these two species. Using 0.63 mm

diameter as the border between sand and gravel (>0.63 mm dia.), and silt and clay

(<0.63 mm dia.), samples taken from widely distributed collecting sites for C.

longilabris averaged 82% sand and gravel and three samples from sites for C.

nebraskana averaged 83% sand and gravel. In each sample gravel (particles >2.0

mm diameter) made up a very small fraction. Although the sample sizes are too

small to be conclusive, these data suggest that both species occur on sandy soils in

which silt and clay are minor components.

The Cicindela sylvatica Group

121

Edaphic factors other than particle size, such as pH, electroconductivity,

moisture and temperature of soil, amount of organic matter present, and chemical

make-up could be important in the distribution of the two species.

By plotting collecting localities from specimen label data on soil maps,

comparisons were made between the distributions of the different forms of C.

longilabris and C. nebraskana and those of the major dominant soil types at the

order and great group level of classification. Table 12 shows the percentages, and

absolute number of locality records occurring on dominant soil types for C.

longilabris and C. nebraskana. Some general trends are apparent.

C. longilabris occurs predominantly on soil types which develop under the

influence of forest vegetation. In eastern North America (from northern Ontario and

Minnesota east) the great majority of locality records of C. longilabris are on map

units indicating Humo-Ferric Podzols as the dominant soil type (Soil Conservation

Service, 1967; Soils of Canada, 1972). Soils of the podzolic order, in the Canadian

classification are well to imperfectly drained mineral soils which typically develop

under coniferous or mixed forest or heath vegetation and characteristically have

complexes of soluble organic matter and compounds of aluminum and iron leached

from the surface layers and deposited in the B or subsurface horizon (Clayton et al.,

1977). The Humo-Ferric great group is widely distributed in Canada, being the

dominant soil type in all podzolic map units indicated on the Soils of Canada (1972)

map. Humo-Ferric Podzols occur mainly on well drained sites and are characterized

by B horizons in which accumulations of aluminum and iron colloids are considered

most significant to their properties, and the organic matter content of the B horizon

is less than 10% (Clayton et al, 1977).

In the prairie provinces a number of soil types are indicated, with Gray Luvisols

(25%) and Black Chernozems (25%) accounting for the greatest percentage of the

locality data for C. longilabris (Table 12). Gray Luvisols are well to imperfectly

drained mineral soils that have developed under the influence of the decomposition

of forest vegetation in mild to cold climates and have silicate clays as the major

accumulation product in the B or subsurface horizon (Clayton et al., 1977). Black

Chernozems are typical of the Aspen Parkland or Fescue Prairie transition zone

between the treeless grasslands of the Mixed Prairie, and the true Boreal Forest

(Clayton et al, 1977).

In the northwestern part of the continent (Alaska, Northwest Territories, Yukon,

and British Columbia), most localities seem to be on Eutric Brunisols and

Humo-Ferrric Podzols. Brunisols are defined by Clayton et al. (1977) as a broad

grouping of imperfectly to well drained soils which have developed under the

influence of forest, alpine, or tundra vegetation, under climatic conditions varying

from Mesic to Arctic, and widely varying moisture regimes, in which the processes

of leaching and weathering are weakly developed, so they tend to reflect the

chemical characteristics of the parent material. Eutric Brunisols are base saturated

soils developed under forest or alpine vegetation, and usually occur on basic or

Quaest. Ent., 1988, 24 (1)

122

Spanton

calcareous parent materials.

In the Rocky Mountain states (which for the sake of this discussion include

Idaho, Montana, Wyoming, the Black Hills area of South Dakota, Colorado,

Arizona, New Mexico, and parts of Nevada and Utah) C. longilabris appears to

occur primarily on montane, or Canadian Zone forest on associations of dominantly

Luvisolic and subdominant Podzolic soils (Table 12). In the Pacific Coast States

most of the localities seem to occur on associations of Luvisols and Brunisols (Table

12).

Soil associations of Cicindela nebraskana are notably different from C.

longilabris. Most collecting records of C. nebraskana were on map units where

Chemozemic soils are dominant (Table 12). In the Canadian classification (Clayton

et al., 1977) Chemozemic soils are well to imperfectly drained soils of good

structure with dark colored virgin or cultivated A horizons, overlying subsurface

horizons of high base saturation. These soils develop within areas of cool to cold

continental climates, their humus enriched A horizons are developed and maintained

by cyclic growth and decay of xerophytic to mesophytic grasses and forbs typical of

grasslands and grassland-forest transition communities. They are typical of the

Canadian prairies and rangelands of interior British Columbia (Clayton et al., 1977),

which are the areas in Canada where C. nebraskana occurs.

In summary, throughout most of the ranges of C. longilabris and C. nebraskana

a separation between species is apparent at the level of soil orders. The former

occurs almost exclusively in areas of Podzolic soils in the eastern half of this

continent, and on Luvisols, Bmnisols and Podzols, soils of montane and subalpine

forest and alpine and arctic meadows in the west. In contrast, C. nebraskana occurs

primarily on Chemozemic soils of grasslands and grassland-forest transition zones.

Across the Canadian prairies the ranges of the two species are sympatric in a broad

zone occurring in the area where Black Chernozems predominate (see maps in

Clayton et al., 1977), which approximates the Aspen-Oak and Aspen Grove

transition zone between prairie grassland and Boreal forest (Rowe, 1972).

More field study of these species is required to elucidate the nature of the

ecological separation which seems apparent from the above comparisons of beetle

distributions and soil maps. For example, in the above mentioned Aspen Grove

regions of Alberta and Saskatchewan more intensive study in the field would likely

show that the two species, though sympatric in a coarse geographic sense, are

separate on a local scale, and exhibit different soil preferences. Similarly in some

areas of the Pacific States and southern British Columbia adults of the two species

have been collected at the same sites (Leffler and Pearson, 1976; Norman Rumpp,

pers. comm.). Additional field study may show that these are merely points of local

parapatry. Leffler and Pearson (1976) have indicated that the form they referred to

as C. longilabris ostenta occurs in the Hudsonian and Alpine-Arctic life zones of the

Olympic Peninsula and Cascade Mountains in Washington State, whereas C.

nebraskana approximately follows the distribution of the Ponderosa

The Cicindela sylvatica Group

123

Pine-Bunchgrass vegetation association in that state.

CLASSIFICATION OF THE NEARCTIC TAXA OF THE CICINDELA

SYLVATICA GROUP

Cicindela longilabris Say

Cicindela longilabris longilabris Say 1824:268. Neotype, New designation (MCZ). Type locality: Silver

Islet, Sibley Prov. Pk., Ontario, here designated. LeConte, 1848:178. 1860:33. 1861:338. Leng,

1902:119. Harris, 1911:20. Horn, 1915:377. 1928:11. 1930:82. Rivalier, 1954:252. Lindroth, 1955:19.

1963:93. Wallis, 1961:46. Graves, 1963:501. 1965:67. Leffler, 1979:467. Boyd and Associates, 1982:6.

Cicindela albilabris Kirby, 1837:12. Type locality: “Taken in Lat. 64 degrees and also in Canada...”.

Emmons, 1854:36.

Cicindela longilabris albilabris; htConit, 1848:178. 1860:33. Casey, 1913:17.

Cicindela longilabris novaterrae Leng, 1918:141. Type locality: Bay St. George, Newfoundland.

Wallis, 1961:46. Boyd and Associates, 1982:6. New Synonymy.

Cicindela oslari terracensis Casey, 1924:13. Type locality: Terrace, British Columbia.

Cicindela var. nebraskana; Horn, 1928:1 1.

Cicindela longilabris laurentii Schaupp, 1884:87. Type locality: Colorado. Leng, 1902:121. Harris, 1911:20.

Casey, 1913:20. Leffler, 1979:474. Boyd and Associates, 1982:6.

Cicindela longilabris oslari Leng, 1902:121. Type locality: San Francisco, San Miguel Mountains,

Colorado. Harris, 191 1:20. Boyd and Associates, 1982:6. New Synonymy.

Cicindela longilabris vestalia Leng, 1902:121. Type locality: Maiden, Montana.

Cicindela oslari densissima Casey, 1924:12. Locality unrecorded, “probably Colorado”.

Cicindela oslari estesiana Casey 1924:13. Type locality: Colorado.

Cicindela laurenti; Casey, 1924:13. Tanner, 1929:82. Dahl, 1941:189.

Cicindela montana laurenti; Wallis, 1961:plate 3.

Cicindela longilabris perviridis Schaupp, 1884:87. Type locality: Sierra and Placer Counties, California

(restricted by Leng, 1902). Harris, 1911:20. Horn, 1915:377. 1930:82. Leffler, 1979:477. Boyd and

Associates, 1982:6.

Cicindela perviridis-, Leng, 1902:122.

Cicindela ostenta Casey, 1913:17. Type locality:Califomia.

Cicindela perviridis placerensis Casey, 1913:18. Type locality: Placer Co., California.

Cicindela ostenta Columbiana Casey, 1924:13. Type locality: British Columbia.

Cicindela montana perviridis-, Wallis, 1961:50.

Cicindela longilabris ostenta Leffler and Pearson, 1976:29. Boyd and Associates, 1982:6. New

Synonymy.

Cicindela longilabris laurentii X Cicindela longilabris perviridis

Cicindela montana oslari-, Wallis, 1961:50.

Cicindela montana laurenti-, Wallis, 1961:50.

Cicindela longilabris oslari-, Leffler and Pearson, 1976:29.

Recognition. — The convex and elongate labrum, bald, broadly excavated head

and non-serrate elytral apices are sufficient to distinguish adult specimens of C.

longilabris and C. nebraskana from those of all other North American species of

Cicindela.

Adults of C. longilabris are distinguished from those of C. nebraskana by the

more coarse, frequently confluent, granulate punctations on the elytral surface,

usually metallic green abdomen, at least in part, or a combination of metallic green,

green and blue, or green and bronze, and by a pattern of maculations on the elytra

including, in most specimens, a humeral and/or post humeral spot or complete

Quaest. Ent., 1988, 24 (1)

124

Spanton

lunule, a sloping, elbowed middle band, and an apical lunule varying from an apical

spot to an entire lunule.

Notes about synonymy and taxonomic history. — There has been much confusion

between C. longilabris and C. nebraskana because of the similarities between

individuals of the latter species and less maculate specimens of the nominate form

of C. longilabris. The fact that populations of the two species can in some places be

found in the same locality where their habitats are immediately adjacent, increases

the confusion.

I have shown evidence from discriminant analysis, statistical differences in size

and proportion of labrum and ratio of head width to pronotal width, differences in

qualitative characters such as female labrum color, color of ventral abdominal

surface, elytral surface and proepistemum, differences in patterns and extent of

markings, as well as an apparent ecological separation based on differences in

distribution relative to major soil types, why two species should be recognized. This

view is supported by the observations of Rumpp (pers. comm.) and Leffler and

Pearson (1976), that no apparent hybridization occurs where sympatric populations

of C. longilabris perviridis and C. nebraskana occur in California and Washington

State.

Kirby’s (1837) description of C. albilabris is sufficient to establish its identity as

the same as Say’s (1824) C. longilabris. Lindroth (1953) examined Kirby’s type

specimen in the British Museum (Natural History), a male from Nova Scotia, and

concluded that it was the same as the nominate form of C. /. longilabris.

Green specimens from the island of Newfoundland were given the subspecific

name C. /. novaterrae by Leng (1902). I believe that, because of the discordant

pattern of variation between Newfoundland and mainland samples shown by the

discriminant analyses (Tests 1 and 2, Tables 5, 6 and 7), and the fact that large

numbers of brown and black specimens are on the eastern part of the island

(Lindroth, 1963), and green specimens can be found on the mainland of Quebec and

Labrador, the formal naming of the Newfoundland populations is inappropriate.

A single specimen from Terrace, British Columbia, which Casey (1924)

described as C. oslari terracensis is black, with the body size, ventral coloration and

elytral markings typical of C. longilabris longilabris Say.

Horn (1928) incorrectly applied the name Cicindela var. nebraskana to

specimens of C. longilabris from Minnesota, which exhibited reduced maculation.

The name C. /. laurentii has historically been applied to olive green, heavily

maculate specimens from the Rocky Mountain region of the United States, and the

name C. /. oslari has been applied to specimens from the same region, which exhibit

less extensive maculation and dorsal coloration varying from green to bronze to

brown.

Leng (1902) described C. /. vestalia based on immaculate specimens from

Maiden, Montana which are, in all other characters, within the range of variation

found in C. /. laurentii throughout the Rocky Mountain region of the United States.

The Cicindela sylvatica Group

125

Specimens described by Casey (1924) as C. oslari densissima and C. oslari

estesiana from Colorado were based on slight individual variations in coloration and

maculation which are within the range of variation to be found in populations of C.

1. laurentii in Colorado.

Wallis (1961) confused the relationships between the forms in this group,

classifying C. m. laurentii {sic), C. m. oslari and C. m. perviridis as subspecies

under the species name C. montana. Wallis applied the names C. montana oslari

and C. montana laurenti (sic) to populations in south central British Columbia

which are in a hybrid zone where the Pacific Coast, Rocky Mountain and boreal

forms of C. longilabris converge and produce populations with a high degree of

individual variation. Specimens from eastern Washington state referred to as C.

longilabris oslari by Leffler and Pearson (1976) are also from the area of

hybridization of the Pacific coast. Rocky Mountain and boreal forms of C.

longilabris

Schaupp (1884) originally described C. /. perviridis as occurring in California,

Oregon, Utah and Newfoundland. Leng (1902) narrowed the type area of this form

to Sierra and Placer counties in California, distinguishing C. /. perviridis from any

Rocky Mountain or Boreal forms which also exhibit a green coloration.

Notes on designation of a neotype for C. longilabris Say. — Thomas Say’s

private collection was entirely destroyed after his death (LeConte, 1859 p. vi). It

would seem to be in the interest of stability of nomenclature, in light of the

confusion which has occurred in the identification and interpretation of the species

C. longilabris and C. nebraskana to designate a type specimen for the former.

Lindroth and Freitag (1969) expressed the desirability of designating neotypes to

stabilize Thomas Say’s names, and Leffler (1979) and Huber (pers. comm.) have

opined that a neotype should be designated for C. longilabris.

Say had distributed specimens from his collection to Dejean in France (Lindroth

and Freitag, 1969). Dejean (1826) indicated that he had received specimens from

Say and he redescribed many of Say’s species (Dejean, 1825-1831). However, C.

longilabris was not among them. Dejean (1837) did not list C. longilabris in the

catalogue of his collection. It appears that none of Say’s specimens of C. longilabris

exist from which to choose a type.

Say (1824) indicated nothing more exact than “Northwest Territory” for a type

locality. Wallis (1961) indicated the approximate route of the second expedition of

Major Long to the source of the St. Peter’s River in 1823, the trip on which Thomas

Say would have collected this species. It started in Philadelphia, from where the

party travelled to Chicago, Minneapolis, and to Big Stone Lake on the border

between Minnesota and the northeast comer of South Dakota. From there the route

led down the St. Peter’s River (now called the Red River) to Lake Winnipeg, and

from there east to Lake of the Woods, around the north shore of Lake Superior and

southward toward Lake Ontario. Wallis (1961) correctly points out that C.

nebraskana does not occur throughout most of the route, but C. longilabris could be

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Spanton

collected anywhere from the Lake Winnipeg area, past the north shore of Lake

Superior and southward much of the way toward Lake Ontario.

LeConte (1859) was very familiar with the work of Say, and was probably

referring to Say’s material when he redescribed C. longilabris (1860). LeConte’s

redescription, though in Latin, is almost identical to Say’s (1824) original

description, including mention of a variant female specimen with reduced or absent

maculation from the north shore of Lake Superior. LeConte (1860) also stated the

range of the species as “New Hampshire, Canada, Mackinaw, Lake Superior...”,

mentioning no other areas. If Say had collected this species farther west than Lake

Superior it would probably have been reflected in LeConte’s statement of the known

range at that time. If Say had collected his specimens very far east and south of Lake

Superior, his statement of the range of the species would perhaps have been Canada,

or Upper Canada, rather than “Northwest Territory”. It seems probable that Say’s

type locality for C. longilabris was somewhere along the north shore of Lake

Superior. Say’s (1824) description of blackish color, head and thorax tinged with

green, white labrum “nearly as long as broad”, elytra with rather large, dense

punctures, humeral and posthumeral spots, reclivate, nearly transverse middle band

and subapical spot, are all characters typical of C. longilabris in northwestern

Ontario .

A male specimen from Silver Islet, Sibley Provincial Park, Ontario, is designated

as neotype and that place as type locality. The neotype specimen’s label reads “Ont.:

Thunder Bay Distr.: Sibley Prov. Pk. 1 km W of Silver Islet on Perry Bay.” The

neotype and three paraneotype specimens, one male and two females, are deposited

in the Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts.

Geographic variation and subspecies. — Adults of C. longilabris are uniform in

most characters among populations across the Acadian, Great Lakes-St. Lawrence

and Boreal regions and the northern part of the montane forest region in western

Canada (Fig. 39). This form typically exhibits a white labrum (Fig. 12),

proepistemum varying between metallic black and green, green, or green and bronze

(Fig. 14), ventral abdominal color of metallic green and blue (Fig. 16), humeral

lunule consisting of a humeral and/or posthumeral spot, occasionally connected

(Fig. 22), a thin, elbowed middle band (Fig. 24), apical lunule consisting of one

subapical spot (Fig. 26), and most specimens have between one and five per cent of

the elytral surface covered by these markings (Fig. 20). This is the nominate

subspecies C. longilabris longilabris Say.

Notable variant populations of this form are those from the island of

Newfoundland, and to a lesser extent the adjacent mainland of Quebec and Labrador

which contain specimens with a green colored dorsum (Fig. 18). As shown in the

discriminant analysis section, specimens from the island of Newfoundland are, on

average, slightly larger in body size than are those from the mainland. I do not

recognize these populations as taxonomically distinct from the nominate form.

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127

Fig. 39. Known distribution of Cicindela longilabris Say.

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Spanton

There is considerable morphometric variation among populations of the Pacific

region from southern British Columbia to central California, and in the Rocky

Mountain region from southwestern Alberta to Arizona and New Mexico. There is

also considerable variation in qualitative characters.

In the Pacific coast states of Washington, Oregon and California most specimens

have a metallic green proepistemum (Fig. 14) and abdomen (Fig. 16) and higher

frequency of green dorsal coloration (Fig. 18) than specimens of the nominate

subspecies. A notable exception is the OR2 population which is more variable in all

three characters than other populations in the region (Figs. 14, 16, 18). The green

elytral surface occurs in all specimens of the Washington state samples (Fig. 18) and

becomes slightly less frequent southwards through Oregon to California.

The pattern of markings on the elytra is expanded in specimens of the Pacific

region over that of typical C. /. longilabris (Fig. 20). This is due to an increase in the

extent of all three of the lunules (Figs. 22, 24, 26). Notable exceptions are specimens

of the Oregon populations, ORl being similar to the other Pacific coast populations

in the middle band (Fig. 24) and apical lunule (Fig. 26) character states, while OR2

is similar to the nominate form of the species in characters of the elytral markings.

In populations of the Rocky Mountain region of the United States there is a

notable increase in bronze coloration of the proepistemum (Character states 6 and 7,

Fig. 14) and an increase in the frequency of brown, bronze and olive green

coloration of the dorsal elytral surface (character state 2, 3, and 4, Fig. 18). The

maculations on the elytral surface are increased in Rocky Mountain populations as

compared to those of the nominate form in the Boreal zone (Fig. 20, 22, 24, 26). The

bronze proepistemal color, brown-bronze-olive green elytral color and development

of the pattern of markings of the elytra reach their maximum frequency in COl,

C02, UT2 and UT3 populations. These characters are variable within this region,

possibly because of geographical fragmentation of the montane forest habitat of

these beetles. As indicated in the discriminant analysis of morphometric data and in

the pattern of variation in color (Figs. 14, 16, 18) and pattern of markings on the

elytra (Figs. 20, 22, 24, 26), there are a few geographically isolated populations

which vary noticeably from the form of the typical C. /. laurentii of central

Colorado. Specimens of the Black Hills of South Dakota (SDl) exhibit somewhat

reduced maculation similar to that of specimens of the boreal regions (Figs. 20, 22,

24, 26), but they exhibit variable coloration (Figs. 14, 16, 18) and occur in an

isolated area of coniferous forest and dominant soil type similar to that of the Rocky

Mountains of central Colorado, and parts of Wyoming and Montana. The

populations from Sandoval County (NMl) and Bernalillo County (NM2), New

Mexico are similarly isolated populations which consist of specimens with reduced

maculation (Figs. 20, 22, 24, 26), variable coloration (Figs. 14, 16, 18) and are

morphometrically different (Spanton, 1983) from those of neighbouring populations.

Samples from Iron County, Utah (UT3), Kaibab Plateau, Arizona (AZl) and

Apache County, Arizona (AZ2) represent geographically isolated populations which

The Cicindela sylvatica Group

129

are visibly differrent from each other. UT3 is distinct in exhibiting the most

consistently heavily maculated specimens of any population in the species complex

(Figs. 20, 22, 24, 26), AZl specimens exhibit a pattern of maculation which is

reduced from that of surrounding populations and AZ2 specimens differ only in

being slightly more variable in coloration of the proepistemum and dorsal elytral

surface than those of most of the neighbouring populations. Rumpp (unpublished)

has proposed subspecific names for these latter three populations. I believe that

although these three populations (AZl, AZ2, UT3) are zoogeographically

interesting, the designation of a formal trinomen for each would not improve our

understanding of variation in this species.

Based on the preceding discussion, the morphometric differences among the

populations of the Pacific region. Rocky Mountain region and Boreal region

presented in the discriminant analysis section, and zoogeographical factors, and

despite the fact that the application of subspecific names to morphologically

heterogeneous groupings of populations is somewhat controversial, I recognize two

subspecies in addition to the nominate form C. longilabris longilabris discussed

previously.

The subspecies C. longilabris perviridis Schaupp here includes the populations

of the Sierra Nevada and Cascade Mountains from east-central California to

southwestern British Columbia (Fig. 39), typified by an increased frequency of

green elytral color, and increased elytral maculation relative to the nominal form, a

smaller average body size than C. longilabris or C. /. laurentii and a proportionately

longer, narrower pronotum than the nominate form.

The subspecies C. longilabris laurentii Schaupp, includes the populations of the

Rocky Mountain region from Arizona and New Mexico to Montana, specimens of

which are typified by brown to bronze or olive green dorsum (Fig. 18), bronze

proepistemum (Fig. 14), and elytral maculation (in most specimens) increased over

that of the nominate subspecies. The arid area of the Great Basin provides a

geographic separation between C. /. laurentii and C. /. perviridis (Fig. 39).

An area of hybridization occurs where the ranges of the three subspecies

converge geographically in southwestern Alberta, northwestern Montana, northern

Idaho, western Washington and southeastern British Columbia (Fig. 39). The

population sample AB6 from the Fincher Creek area of Alberta is an example of a

hybrid population. The series of specimens was collected the same day at the same

local site. Specimens in the series exhibit each of the various color states of the

proepistemum (Fig. 14), all but one of the color states of the abdomen (Fig. 16), and

all of the color states of the elytra (Fig. 18). The complete range of pattern is

represented, from completely immaculate specimens to specimens with 20 per cent

of their elytral surface covered with white markings. More intensive field collections

of large series of specimens from this area are required for a detailed study of the

pattern of hybridization between the forms.

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Spanton

Distribution. — The geographic range of C. longilabris is shown in Figure 39. In

eastern North America the southern limit approximates the southern limit of Podzols

as the dominant soil type, and the southern limit of the ranges of Jack Pine (Pinus

banksiana Lamb.), White Spruce (Picea glauca (Moench) Voss), Balsam Fir (Abies

balsamea (L.) Mill) and Balsam Poplar (Populus balsamifera Linnaeus) (Little,

1971). Across the prairie provinces the southern limit of the distribution of C.

longilabris corresponds with the southern limit of boreal forest and boreal

forest-grassland transition, and in the western United States the range of this species

approximates the range of montane and subalpine forests, or the Canadian life zone

of Merriam (1898). In the north the range of C. longilabris appears to be limited by

the northern limit of wooded country and the southern limit of continuous

permafrost.

A complete list of locality data is in the archives of the University of Alberta,

and is available on request to the Department of Entomology. Included here is a list

of those localities which I consider to be in the hybrid zone mentioned above. The

number of specimens and the collection(s) where they are housed is indicated, or

alternately, the reference is cited.

C. /. longilabris X C. 1. laurentii X C. 1. perviridis Intergrades

CANADA.

Alberta: Beaver Mines, 20 km W Fincher Creek (35, LU), Banff (14, CNC; 6, CU; 1, UMAA; 4, AMNH;

102, CAS), 4.5 mi N Banff (43, AMNH), Frank (1, CNC), Ft. McLeod “Brit. Amer” (1, AMNH), Loggan

(20, AMNH; 1, CAS), Waterton Lakes (3, CNC). British Columbia: Ainsworth (1, CU), Creston (6, CNC; 4,

CU; 2, UMAA; 40, UBC; 42, AMNH; 16, CAS), Femie (5, CAS), Golden (2, AMNH), Hosmer (Leffler,

1979), Kaslo (3, CU; 7, UMAA; 19, AMNH; 2, CAS), Loiggan (1, CU), Little Vermillion River (Leffler,

1979), Sanca (2, UBC), Wynndel (12, CNC; 1, UBC; 7, AMNH; 8, CAS; 1, UAE), Yoho Valley (Leffler,

1979).

UNITED STATES.

Idaho: Benewah Co.: Emerald Creek (Leffler, 1979), N. S. Ski Bowl (1, UIM), Potlach 21.5 mi NE (Leffler,

1979). Bonner Co.: Clark Fork 8 mi E (3, UIM), Preist Lake (1, UIM), Preist River Exp. For. (1, UIM),

Trout Creek 12 mi SE Sandpoint (6, UIM). Boundary Co.: Brush Lake (2, UIM), Caribou Cr. 17 mi W

Naples (2, UIM), Ruby Pass 13 mi NW Naples (1, UIM). Clearwater Co.: Elk River 3 mi N (1, UIM).

Elmore Co.: Atlanta (Leffler, 1979). Idaho Co.: Kooskia 15 mi E (2, REA), Lolo Pass (1, UIM), Moose

Creek (15, USU), Moose Creek R. S. Orangeville (5, REA). Kootenai Co.: Coer D’Alene (3, CAS; 1, UIM).

Latah Co.: Bovill (1, WSU), Flat Creek (1, REA), Harvard (1, UIM), Harvard 7 mi SE Sand Creek (1,

WSU), Little Bear Cr. Helmer (1, UIM), Moscow Mtn. (7, UIM; 1, SMEK; 2, WSU), Moscow, 6 mi NE (1,

UIM; 3, WSU), Troy (4, UIM). Lemhi Co.: Gibbonsville (1, UIM), Meadow Lake 6 mi N Gilmore (7, UIM).

Shoshone Co.: Pine Creek (3, CMP), Wallace (2, UIM). Valley Co.: Egger’s Creek (Leffler, 1979), McCall

(9, UIM), Yellow Pine (1, UIM). Localities of unknown counties: Spelling (1, WSU), Lk. Waha (1, UIM; 1,

CAS). Montana: Deer Lodge Co.: Lost Creek Pass (1, CNC). Rathead Co.: Kila, 8 mi S (Leffler, 1979).

Glacier Co.: St. Mary’s (1, UMAA). Jefferson Co.: Homestake Pass (Leffler, 1979). Lewis and Clark Co.:

Helena (2, CAS), Roger’s Pass Summit 20 km W Lincoln (1, LU). Missoula Co.: (2, USU), Blue Mountain

(Leffler, 1979), Greenhough (Leffler, 1979), Kitchen Creek (Leffler, 1979), Missoula (1, MSU; 1, USNM),

Pattee Canyon (Leffler, 1979). Ravalli Co.: Blodgett Mtn. (1, MSU), Blue Nose Peak (2, AMNH; 2, WSU),

Camp Creek (1, MSU), Come Lake (2, AMNH), Darby (1, MSU), Darby 19 mi SW (Leffler, 1979), East

Fork (1, AMNH), Girds Creek (5, MSU; 1, CAS), Hamilton (1, CNC; 2, MSU; 1, AMNH), Hamilton 6 mi

NW (Leffler, 1979). Sanders Co.: Kaniksu Nat. For. Bull R. Campground (1, SMEK), Thomson Falls (2,

CAS), Weeksville (MCZ), White Pine (Leffler, 1979). Silver Bow Co.: Butte (1, MSU). Localities of

The Cicindela sylvatica Group

131

unknown counties: Camp Pleasant (1, CAS), Flathead Nat. For. Big Creek (1, CMP), Glacier Park (1, CU),

Glacier Park, Indian Ridge (2, AMNH), Glacier Park, McGee Meadow (1, AMNH). Oregon: Umatilla Co.:

Athena Wild Horse Mtn. (1, AMNH), Bone Springs Blue Mtn. (12, AMNH), Meacham (2, CAS), Tollgate

(11, AMNH; 2, CAS). Union Co.: Oregon Trail Camp (1, SMEK). Localities of unknown counties: Moffat

Head, Blue Mtns. (1, CAS), Wallowa Mtns.: Chimney Lake (2, USU), Minam L. Area (1, WSU), Morcastle

Lake (1, WSU). Additional Oregon records from Leffler (1979): Baker Co.: Anthony Lake; Baker; Pine

Creek nr. Baker; Durkee. Grant Co.: Summit Dixie Pass. Umatilla Co.: Deadman’s Pass. Union Co.: Elgin;

Phillips Canyon 6.8 km NE Elgin; 8 mi E LaGrande. Wallowa Co.: Hat Point; French Forest Camp; Lost

Line River; Wallowa Lake. Washington: Columbia Co.: Blue Mtns. Tollgate Road (1, WSU). Pend Oreille

Co.: 7 mi W. Locke (3, PSU). Stevens Co.: Blackwelder (1, UWM). Walla Walla Co.: Walla Walla (1,

WSU). Additional Washington records from Leffler (1979): Columbia Co.: Blue Mtns., Goodman Springs:

Lewis Peak. Whitman Co.: Pullman.

Cicindela nebraskana Casey

Cicindela nebraskana Casey, 1909:268. Type locality: Nebraska. Casey, 1914:18. Leffler, 1979:484. Boyd

and Associates, 1982:6.

Cicindela montana LeConte, 1861:338 (not Charpentier, 1825). Type area: Valleys of the Bitter Root

Mountains of eastern Idaho and western Montana (here restricted: see ‘Notes about synonymy and

taxonomic history’). Casey, 1914:17.

Cicindela longilabris montana LeConte, 1875:157. Schaupp, 1884:87. Leng, 1902:122. Harris, 1911:20.

Cicindela longilabris nebraskana-, Harris, 191 1:20.

Cicindela montana canadensis Casey, 1913:17. Type locality: Calgary Alberta.

Cicindela spissitarsis Casey, 1913:18. Type locality: Aweme, Manitoba. Casey, 1914:17.

Cicindela canadensis Casey, 1914:17.

Cicindela calgaryana Casey, 1914:17. Type locality: Lethbridge, Alberta.

Cicindela montana uteana Casey, 1924:12. Type locality: Provo, Utah.

Cicindela longilabris chamberlainv, Knaus, 1925:182. Type locality: “Stein Mountains”, Steens

Mountains, Harney Co., Oregon.

Cicindela montana montana-, Wallis, 1961:49.

Cicindela montana spissitarsis-, Wallis, 1961:50.

C. montana chamberlainv, Wallis, 1961:50.

C. montana montana-, Leffler and Pearson, 1976:33.

Cicindela nebraskana chamberlaini Knaus; Leffler, 1979:486. Boyd and Associates, 1982:6. New

Synonymy.

Recognition. — Adults of C. nebraskana are distinguished from C. longilabris

adults by the relatively smooth elytral surface with punctations occurring in discrete

fashion, with smooth fields between, normally black abdomen varying to metallic

purple or blue or blue and green in a small percentage of specimens, absence of

humeral and apical lunules, and, in most specimens, absence of middle band, black

dorsal coloration in the majority of specimens, and a labrum which is light tan in

color in 44.8%, intermediate colored or mottled in 30.6% and black in 24.6% of

females, and light in 82.6%, intermediate or mottled in 15.6%, and black in 1.8% of

male specimens. Descriptions of C. nebraskana are found in LeConte (1861) under

the name C. montana, and in Leng (1902), Casey (1909) and more recently, Leffler

(1979).

Notes about synonymy and taxonomic history. — LeConte’s (1861) name C.

montana is a junior homonym of C. montana of Charpentier, (1825), itself a junior

synonym of C. hybrida riparia Dejean (Huber, 1969). Cicindela nebraskana Casey

1909 is the oldest available junior synonym, and therefore becomes the correct name

of the taxon.

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Spanton

LeConte (1861) indicated the type area for his Cicindela montana as “Valleys of

the Rocky Mountains”. Two specimens of his type series of three (housed at the

Museum of Comparative Zoology, Harvard University) bear the locality data “Bitter

Root” on their labels. These specimens were very probably collected in the Bitter

Root Mountains of eastern Idaho and western Montana, and I have restricted the

type area, accordingly.

Casey’s names C. spissitarsis, C. canadensis and C. calgaryana are based on

individual variants. Casey’s (1924) name C. montana uteana was based on one

specimen from Provo, Utah which exhibited a slightly cupreous brown dorsal

coloration and more metallic luster than is typical of the species. A few specimens

of C. nebraskana in that area exhibit some metallic green and blue abdominal color

and a slightly bronze brown dorsal coloration.

Knaus’s C. /. chamberlaini (1925) is based on a series of specimens in which a

number have a slender middle band which occurs in other specimens of this species.

Geographic variation. — The frequency of occurrence of females with a dark

colored labrum varies greatly between localities, but a geographic pattern is not

evident (Fig. 13). The color of the proepistemum is black in most specimens of most

populations sampled, (Fig. 15) but a significant number of specimens in Manitoba,

Idaho, Wyoming and Utah population samples are metallic green, blue, or bronze.

The color of the abdomen is also variable in some populations (Fig. 17). Populations

AB2 and MTl include many specimens with a brown abdomen and MBl, WYl,

WY2, and UTl have many specimens exhibiting metallic greens and blues.

Elytral color varies little in C. nebraskana. In most populations a majority of

specimens have a black dorsum (Fig. 19). UTl is a variable population with a larger

number of specimens exhibiting some brown or dark olive green elytra.

Most specimens in all populations of C. nebraskana are immaculate or very

nearly so (Figs. 21, 23, 25, 27). In those populations (IDl, WY2, Fig. 21) where the

percentage of the elytral surface covered with maculations is slightly increased, it is

because of the presence of a thin or incomplete middle band in some of the

specimens (Fig. 25). In all of the sampled populations the humeral lunule and apical

lunule (Figs. 23, 27) are absent in the majority of specimens.

Leffler (1979), in studying tiger beetles of the northwestern United States

recognized two subspecies, C. n. nebraskana east of the continental divide, and C. n.

chamberlaini west of the divide, based on a statistical difference in head width, the

eastern form having broader heads, on average. Head width was not selected in

discriminant tests 5 and 6 as a variable contributing significant variance to the

discriminant functions (Table 10). In the discriminant analysis section I reported

longer average femur lengths and elytral lengths in populations west of the divide,

which suggests that western populations of this species are larger in body size, on

average than eastern populations. These findings are not significant justification for

subspecific recognition. I treat C. nebraskana as a monobasic species.

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133

Fig. 40. Known distribution of Cicindela nebraskana Casey.

Distribution. — The geographic distribution of C. nebraskana is illustrated in

Figure 40. The northern limit of the distribution of this species approximates the

northern limit of Chemozemic soils in grassland and grassland-forest transition

zones across the prairie provinces of Canada and in British Columbia. In the west its

range follows that of the Ponderosa pine-bunchgrass vegetation zone. The range of

C. nebraskana is limited to the south by arid lands of the Great Basin. Conditions

affecting the eastern limits of the distribution of this species are unclear, but

probably involve edaphic factors which are not evident at the order and great group

level of soil classification.

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A complete list of locality data is in the archives of the University of Alberta and

is available on request to the Department of Entomology.

GENITALIA AND EVOLUTIONARY ASPECTS OF THE CICINDELA

SYLVATICA SPECIES GROUP

Genitalia of the C. sylvatica group

Rivalier (1950, 1954), in classifying the Palearctic and Nearctic Cicindela

concluded that C. longilabris (which presumably included C. nebraskana) and the

Palearctic species C. soluta Dejean, C. lacteola Pallas, C. japana Motschulsky, C.

gemmata Faldermann, C. sylvatica Linneaus and C. granulata Gebler form a group

of closely related species based on similarities of the sclerotized structures of the

internal sac of the male aedeagus.

Previously the male genitalia of C. soluta and its geographic variation were

studied by Mandl (1936), and Papp (1952) figured the male genitalia of C.

longilabris. Mandl (1970) examined the male genitalia of C. sylvatica and described

a new subspecies, C. sylvatica reiseri, based partly on slight differences in the

sclerites of the internal sac.

Drawings of the dorsal view, in its fully extruded state, and left lateral view of

the male aedeagus and the sclerites of the internal sac are here presented for the

species of the sylvatica group of Rivalier (1950, 1954) (Figs. 41^9). Figures 50 to

57 illustrate the female genitalia of the species of the same group. The nomenclature

applied here is that of Freitag et al. (1985).

Figures 41 and 42 illustrate male genitalia of C. longilabris. The median lobe of

the aedeagus is of moderate length, curved in lateral view, with lateral flanges at the

apical 0.25 of its length, which converge towards the apex producing a ventrally

curved point. The left lateral apical flange is usually slightly more pronounced than

that on the right. The parameres (Fig. 42) are usually 0.66 to 0.75 of the length of

the median lobe and taper to thin pointed apices. In all of the approximately 90

examined male specimens of the C. sylvatica group the right paramere is shorter

than the left. In 23 measured specimens the right paramere averaged 85% of the

length of the left, with a range of 78% to 99%. Figure 41 shows the arrangement of

sclerites of the internal sac of the aedeagus of C. longilabris. The numbered sclerites

are probably homologous with the correspondingly numbered structures on the

internal sac in the C. maritima group as described by Freitag (1965). The largest is

number 6, the median tooth, “la grande dent” of Rivalier (1950). The flagellum, “le

flagelle” of Rivalier (sclerite 4, Figs. 41, 42), is a long, tapering sigmoid-shaped

structure, thickened at its base, and pointed apically. A small sclerite 3, “le clou” of

Rivalier (1950) is ventral to the base of the flagellum. The large curved sclerite 2,

“la piece arciforme” of Rivalier (1950) and a smaller twisted sclerite 1, “la baguette”

of Rivalier (1950) are situated dorsally in the internal sac (Fig. 41).

The lateral flanges (Fig. 42) are more pronounced in some individuals, but no

taxonomically significant variation was found in development of lateral flanges.

The Cicindela sylvatica Group

135

Figs. 41^3. Aedeagus, left lateral aspect (extruded in mating position) and left lateral aspect of internal sac

showing internal sclerites of C. longilabris Say from Stanley Hill Cemetery, Thunder Bay District, Ontario.

(Fig. 41). Aedeagus, dorsal and left lateral aspects (extruded in mating position) and sclerites of the internal

sac of C. longilabris Say from Gander, Nfld. (Fig. 42). Aedeagus, dorsal and left lateral aspects and sclerites

of the internal sac of C. nebraskana Casey, from 10 mi. east of Patricia, Alberta. (Fig. 43). Legend: If, lateral

flange; ml, median lobe; p, paramere; 1, sclerite 1, probably homologous with sclerite 1 of Freitag (1965),

“la baguette” of Rivalier (1950); 2, sclerite 2, probably homologous with sclerite 2 of Freitag (1965) and “la

piece arciforme” of Rivalier (1950); 3, sclerite 3, probably homologous with sclerite 3 of Freitag (1965) and

“le clou” of Rivalier (1950); 4, flagellum, homologous with sclerite 4 of Freitag (1965), “le flagelle” of

Rivalier (1950); 5, sclerite 5, probably homologous with sclerite 5 of Freitag (1965); 6, median tooth, “la

grande dente” of Rivalier (1950), possibly homologous with sclerite 6 of Freitag ( 1965).

Quaest. Ent., 1988, 24 (1)

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Spanton

Figs. 44—46. Aedeagus, ventral and left lateral aspects, and sclerites of the internal sac of C. sylvatica

Linnaeus from Baikal, Siberia, USSR (Fig. 44); C. granulata Gebler from Turkestan, USSR (Fig. 45), and

C. japana Motschulsky from Kanazawa, Japan (Fig. 46). Sclerites of the internal sac are numbered as in

Figures 28 and 29, except where homologies are uncertain; a, possibly a fusion of sclerites 4 and 5; b,

sclerite of uncertain homology.

5

6

Figs. 47^9. Aedeagus, ventral and left lateral aspects, and sclerites of the internal sac of C. gemmata

Faldermann from Korea (Fig. 47), C. soluta Dejean from Hungary (Fig. 48), and C. lacteola Pallas labelled

“Turk” (Fig. 49). Legend; a, b, c, sclerites of unknown homologies; sc, sclerite cluster.

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Spanton

shape of aedeagus, or sclerites of the internal sac among the sampled populations of

C. longilabris.

Male genitalia of C. nebraskana (Fig. 43) exhibit inter-population variation no

greater than intrapopulation variation. There may be slightly more pronounced

flanges of the median lobe in C. nebraskana than in C. longilabris but larger series

of specimens would have to be examined to adequately test that hypothesis. Norman

Rumpp (unpublished data) has found a statistical difference in the size of the

genitalia between C. nebraskana and C. longilabris where the species occur

sympatrically in east central California. The former has a longer aedeagus and

longer median tooth of the internal sac, consistent with the larger overall body size

of C. nebraskana relative to the smaller form of C. longilabris which occurs in the

area (Rumpp, unpublished data).

Figure 44 shows the male genitalia of C. sylvatica. The median lobe of the

aedeagus is different from that of C. longilabris or C. nebraskana in its long narrow

apex. The flagellum (sclerite 4) of C. sylvatica is short and stout and the median

tooth (sclerite 6) lacks the expanded base evident in the two Nearctic species.

Sclerites 1, 2, and 5 (Fig. 44) are relatively small in size.

The male genitalia of C. granulata (Fig. 45) are distinguished by the thick

median lobe with an acutely pointed apex and the stout flagellum which is not

greatly thickened at its base, relative to that of C. longilabris (Fig. 42), C.

nebraskana (Fig. 43) or C. sylvatica (Fig. 44).

The median lobe of C. japana (Fig. 46) is intermediate in shape between that of

C. longilabris or C. nebraskana (Figs. 41, 42) and the narrower, elongated shape of

C. sylvatica (Fig. 44). The sclerites of the internal sac distinguish C. japana .

Sclerite a (Fig. 46) could be the result of a fusion of the flagellum (sclerite 4) and

sclerite 5, but this is uncertain. Sclerites 2 and 3 (Fig. 46) are small in size and

sclerite 6 is probably a reduced median tooth. The homology of sclerite b in Figure

46 is uncertain.

The male genitalia of C. gemmata are shown in Figure 47. The median lobe of

the aedeagus is large, consistent with the large size of the beetle and is markedly

curved in lateral view. The most notable character is the sclerite cluster of the

internal sac (Fig. 47).

The male genitalia of C. soluta (Fig. 48) are smaller in size, consistent with the

small body size of the beetle, and are unique among the species of the sylvatica

group in the shape of the projection at the apex of the aedeagus. The sclerite cluster

of the internal sac (Fig. 48) is similar to the sclerite cluster of C. gemmata (Fig. 47).

Structure c in Figures 47 and 48 could possibly be the median tooth (sclerite 6)

reduced in size and lacking the enlarged base present in C. longilabris (Figs. 41, 42).

The homologies of other sclerites of the internal sac of C. gemmata and C. soluta

are unclear.

The male genitalia of C. lacteola (Fig. 49) are distinct in the size and shape of

the median lobe of the aedeagus and the semi-circular ring of sclerites in the internal

The Cicindela sylvatica Group

139

sac which probably represent an enlargement and breaking up of sclerite 2. Sclerite

6 (Fig. 49), the median tooth of the internal sac in C. lacteola is narrower at its base,

and slightly more curved than that of other species in the species group.

Figure 50 shows the female genitalia of C. longilabris. Individuals vary slightly

in the size and shape of the oviduct sclerite, the degree of development of the ventral

notches of the second gonacoxae, and may vary markedly in the shape of the apex of

the second gonapophyses. The second gonapophyses may, in some older individuals

showing signs of much abrasion, wear down to 0.50 the length depicted in Figure

50. No taxonomically significant differences in female genitalic structures were

observed, among populations of C. longilabris.

The female genitalia of C. nebraskana (Fig. 51) and C. longilabris are not clearly

separable.

The female genitalia of the Palearctic species C. sylvatica (Fig. 52) show no

characters that are distinctly different from C. longilabris and C. nebraskana,

although syntergum 9 and 10 of the former species is slightly more rounded.

The female genitalia of the Palearctic species C. granulata (Fig. 53) are distinct

in having a relatively short, wide oviduct sclerite, two small sclerotized structures

visible in the right side of the bursa copulatrix and a slightly narrower syntergum 9

and 10 with lateral margins straighter than those of C. sylvatica.

The female genitalia of C. japana (Fig. 54) are distinguished by the combination

of a large oviduct sclerite, a small, quadrate sclerite in the membrane between the

second gonacoxae, and rounded apices of sternum 8. The rounded apices of sternum

8 of this specimen (Fig. 54) may be largely a result of wear, as there are no setae on

the apices and the second gonapophyses showed signs of much abrasion. The few

female specimens of C. japana I was able to examine were all old and worn.

The female genitalia of C. gemmata (Fig. 55) are distinct in having a wide,

rounded, slightly triangular shaped oviduct sclerite and the notched shape of the

second gonapophyses.

The female genitalia of C. soluta are distinguishable by several characters (Fig.

56). The shape of the oviduct sclerite is different from other species in the group,

there is a pronounced point on the medial edge of the second gonacoxae

immediately basal to the notch, the second gonapophyses have a notched shape to

their lateral edge, sternum 8 is uniquely shaped at its apex with one distinct seta at a

notch at the apex on each side, and the lateral portions of syntergum 9 and 10 are

more rounded in shape than other species of this group. The notched lateral edge of

the second gonapophyses of C. soluta (Fig. 56) suggests a close relationship with C.

gemmata (Fig. 55).

The female genitalia of C. lacteola (Fig. 57) are distinct in the truncated setose

apices of sternum 8, and the three small, dark, heavily sclerotized structures inside

the bursa copulatrix.

The number of setae on the ventral surface of the second gonapophyses is not

useful in distinguishing species of this group, because of its variability. In C.

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Spanton

Figs. 50-52. Female genitalic structures of C. longilabris Say, from Harmon Field, Nfld.(Fig. 50), C.

nehraskana Casey from 10 mi E of Patricia, Alberta (Fig. 51), and of C. sylvatica Linnaeus, from Baikal,

Siberia, USSR (Fig. 52). Legend: bcx, bursa copulatrix; co, common oviduct; mr, median ridge of ventral

sclerite of bursa copulatrix; n, ventral notch of second gonacoxa; os, oviduct sclerite; s, spermatheca; sd,

spermathecal duct; sgp, second gonapophysis; sgx, second gonacoxa; s8, sternum eight, ventral aspect;

t9&10, syntergum nine and ten, dorsal aspect.

Figs. 53-55. Female genitalic structures of C. granulata Gebler, from Turkestan, USSR (Fig. 53); C.japana

Motschulsky from Kanazawa, Japan (Fig. 54); and C. gemmata Faldermann, from Kanazawa, Japan (Fig.

55). Structures are labelled in Figure 50.

Quaest. Ent., 1988, 24 (1)

Figs. 56 and 57. Female genitalic structures of C. soluta Dejean from Hungary (Fig. 56) and of C. lacteola

Pallas from Aulic Ata (Fig. 57). Structures are labelled in Figure 50.

Fig. 58. Reconstructed phylogeny of the Cicindela sylvatica species group. Dark boxes indicate apotypies,

light boxes indicate plesiotypies, crosses indicate reversals, and parallel lines indicate homoplasies.

Characters (primary numbers) and character states (superscripts) are as indicated in Table 13.

Quaest. Ent., 1988, 24 (1)

144

Spanton

longilahris this varies from zero in some individuals to four in others and differs

between the right and left sides of many individuals.

Reconstructed Phytogeny

W. Horn (1926) made one of the earliest attempts to arrange some of the species

of the genus Cicindela into species groups, based on external structure. Rivalier

(1950, 1954, 1957, 1961, 1963) carried out studies of the male genitalic structures of

species of Cicindela, and based thereon arranged the species in genera, subgenera

and species groups. As Freitag (1974) indicated, Rivalier’ s classification has been

largely accepted in North America, although Rivalier’ s genera have been treated as

subgenera, and his subgenera as synonyms, as reflected in a recent checklist of the

North American tiger beetles (Boyd and Associates, 1982).

Rivalier (1950, 1954) grouped the North American species C. longilahris with

the Palearctic species C. soluta Dejean, C. lacteola Pallas, C. japana Motschulsky,

C. gemmata Faldermann, C. sylvatica Linneaus, and C. granulata Gebler. He

associated this sylvatica group with the C. hybrida, C. transhaicalica and C.

maritima groups (groups I, II, and IV, respectively, of Rivalier, 1950, 1954), in what

is now considered the subgenus Cicindela. Freitag (1974) considered the maritima

group, among the North American fauna, to be the most closely related to C.

longilahris and C. nebraskana. Both authors (Freitag, 1974; Rivalier, 1950, 1954)

considered these species groups to be among the most primitive in the genus

Cicindela . For these reasons, and because published information (Freitag, 1965,

1974), and specimens are more available to me than for either of the Palearctic C.

hybrida and C. transhaicalica groups, I have used the C. maritima group as the

outgroup for the C. sylvatica group, for purposes of character polarization for a

phylogenetic analysis.

The methods used in phylogenetic reconstruction have been discussed by Hennig

(1966), Ross (1974), Wiley (1981), and Watrous and Wheeler (1981), among many

others. Basically, such an analysis involves the identification of the sister group of

the taxon to be analysed, shared character states are assumed to have been inherited

from a common ancestor (in this study an ancestor common to both the sylvatica

and maritima species groups), and are designated as plesiotypic (primitive or

ancestral). Apotypic (derived) character states are then used as evidence of

phylogenetic affinity, or relative recency of common ancestry, between the species,

or taxa sharing such character states.

Phylogenetic analysis of character states. — Characters are listed numerically, in

the sequence in which they appear in Table 13 and Figure 58, and plesiotypic and

apotypic states are specified for each. The notation used by Ball (1985) is here used,

in which apotypic states believed to be part of multi-state transformation series are

numbered consecutively, the lowest numbered state being most like the plesiotypic

state. Apotypic states believed to have evolved independently from the plesiotypic

condition are indicated by different capital letters.

The Cicindela sylvatica Group

145

01. Female genitalia, 8th sternum. Four states: plesiotypic, with angular, pointed

apices (Figs. 50-52); apotypic state A, with rounded apices (Figs. 53, 54); apotypic

state B, each apex with a notch bearing a stout seta (Fig. 56); apotypic state C,

apices truncate, with 5 or 6 setae at apical margin (Fig. 57).

02. Male genitalia, apex of aedeagus. Three states: plesiotypic, apex of aedeagus

narrowed to a rounded point (Figs. 41^3); apotypic state A, apex of aedeagus

prolonged (Figs. 44, 45); apotypic state B, apex of aedeagus with a laterally

flattened projection (Fig. 48).

03. Male genitalia, sclerites of the internal sac. Four states: plesiotypic, six distinct

sclerites present (Figs. 41-45); apotypic state A, sclerites 4 and 5 fused (Fig. 46a);

apotypic state B, sclerite cluster ‘a’ present (Figs. 47, 48); apotypic state C, sclerite

2 divided into three parts (Fig. 49).

04. Humeral lunule configuration (Fig. 6). Four states: plesiotypic, complete lunule

(Fig. 8(4)); apotypic state Al, lunule reduced to two dots (Fig. 8(3)); apotypic state

A2, absent from most specimens; apotypic state B, lunule expanded and continuous

with markings.

05. Middle band configuration (Fig. 6). Three states: plesiotypic, lunule present;

apotypic state A, lunule absent; apotypic state B, lunule broadened and continuous

with other markings.

06. Apical lunule configuration (Fig. 6). Four states: plesiotypic, lunule complete;

apotypic state Al, reduced to one spot; apotypic state A2, lunule absent completely;

apotypic state B, lunule expanded and continuous with other markings.

07. Elytra, color of dorsal surface. Three character states: plesiotypic, grey-brown

to black; apotypic state A, brown and metallic green and/or blue; apotypic state B,

metallic red to bronze.

08. Frons, presence or absence of setae (other than supro-orbital setae). Two states:

plesiotypic, present; apotypic, absent.

09. Elytra, surface texture. Three character states: plesiotypic, roughly granulate,

appearing dull to the unaided eye; apotypic state A, finely granulate with slight

depressions surrounded by smooth fields, appearing shiny to the unaided eye;

apotypic state B, finely granulate, appearing smooth to the unaided eye.

10. Labrum, shape, lengthlwidth. Two states: plesiotypic, short; apotypic, long.

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Spanton

11. Lahrum, shape of median longitudinal ridge. Three states: plesiotypic, absent

(labrum flat, or nearly so); apotypic state Al, broad, rounded median ridge; apotypic

state A2, median ridge in the form of a sharp carina.

12. Body size. Two character states: plesiotypic, medium sized Cicindela\ apotypic,

large sized Cicindela.

13. Labrum, color. Two states: plesiotypic, light tan color; apotypic, dark brown to

black color, at least in some individuals.

14. Antennal scape, ''other setae” {¥'\g. 3). Three states: plesiotypic, zero to four in

number; apotypic state A, four to 10 in number; apotypic state B, more than 20. The

20+ number of setae found at this locus on the scape in specimens of C. soluta was

interpreted as having been independently derived from the plesiotypic state because

C. soluta shares two good synapotypies with C. gemmata, and because this character

is sufficiently labile among species of Cicindela that I could not assume that the

condition was derived from the four to 10 state exhibited by C. sylvatica and C.

granulata (Table 13, Fig. 58).

S

15. Female genitalia, second gonapophyses. Two states: plesiotypic, with smoothly

curved lateral margins (Figs. 50-54); apotypic, with notched lateral margins (Figs.

55, 56).

16. Female genitalia, oviduct sclerite. Three states: plesiotypic, shield shaped (Figs.

50-53); apotypic state A, larger in size (Fig. 54); apotypic state B, triangular shaped

(Fig. 55).

17. Female genitalia, quadrate sclerite in membrane between 2nd gonacoxae. Two

states: plesiotypic, quadrate sclerite absent; apotypic, quadrate sclerite present (Fig.

54).

For characters which vary within a taxon, that state exhibited by the majority of

specimens over most of the geographic range of the species was coded for cladistic

analysis. For example, character 7, color of the elytral surface, is quite variable in C.

longilabris including bronze to dull green specimens for C. 1. laurentii in the Rocky

Mountain region, and some green specimens of C. /. longilabris in Newfoundland,

and C. /. perviridis in California and Oregon, to predominantly black or dark brown

over most of the range of the species. Cicindela longilabris was coded as “0”

because the plesiotypic dark brown or black color exhibited by specimens in most of

the geographic range of the species. A similar coding strategy was applied for C.

longilabris for the characters of the elytral markings (characters 4,5,6), all of which

show some variation in this taxon.

The Cicindela sylvatica Group

147

Reconstructed Phytogeny. — The reconstructed phylogeny appears in Figure 58.

The first branching point on the cladogram is a divergence of C. lacteola from the

common ancestor of the other seven species in the group. Cicindela lacteola

exhibits a number of autapomorphies, including distinctly truncate, setose apices of

sternum eight (character 1), sclerite two of the internal sac divided into three parts

(character 3), humeral lunule, middle band and apical lunule expanded and

confluent (characters 4, 5, 6), and a metallic red -bronze elytral color (character 7).

The remaining seven species in the group share an apotypically reduced humeral

lunule (character 4), an apical lunule reduced to a single spot (character 6), labrum

with a broad rounded median ridge (character 1 1), and a larger body size (character

12). Cicindela japana, C. gemmata and C. soluta share the apotypic color of the

elytra (character 7). Cicindela gemmata and C. soluta share an apotypic sclerite

cluster ‘a’ of the internal sac of the male aedeagus (character 3), and the notched

second gonapophyses of the female genitalia (character 15). The ancestor of C.

nebraskana, C. longilabris, C. sylvatica, and C. granulata is assumed to have had an

apotypically dark colored labrum (character 13), a condition which is assumed to

have reversed to its plesiotypic state in C. granulata. Cicindela sylvatica and C.

granulata share a prolonged apex of the aedeagus (character 2), and an increased

number of ‘other setae’ on the antennal scape (character 14). Cicindela nebraskana

and C. longilabris both exhibit setae on the frons (character 8), which are lacking in

C. sylvatica and C. granulata.

Homoplasies were invoked to explain the distribution of characters 1, 8, 9, 10,

and 14, and reversals were required in characters 4, 6, and 12. I placed emphasis on

(informally weighted) characters of the male and female genitalia (characters 1, 2, 3,

15, 16, 17), the labrum (characters 10, 11, 13), and presence of setae on the frons

(character 8), and less emphasis on characters such as elytral maculations

(characters 4, 5, 6) and elytral color (character 7). I have shown that these latter

characters are highly variable across the range of C. longilabris and the recent work

of Schultz (1986) with C.formosa, and Acorn (in prep.) with C. scutellaris, show

that in other tiger beetle species as well, characters of color and color pattern are

extremely labile and subject to local selection. An obvious weakness of this

reconstruction is that elytral color is the only synapotypy uniting C. japana, C.

gemmata and C. soluta.

The lack of a synapomorphy at the basal node suggests that this group is

polyphyletic: perhaps C. lacteola, which exhibits a number of autapomorphies,

should not be classified in this species group. This lack of a basal synapomorphy for

the group is based, in part, on my assumption that the apotypic long labrum

(character 10) arose independently in C. lacteola, and in the lineage which gave rise

to the species C. nebraskana, C. longilabris, C. sylvatica, and C. granulata. The

alternate hypothesis, that a long labrum arose once in a common ancestor of the

group, and subsequently underwent reversal to the short plesiotypic state in the

clade which includes C. japana, C. gemmata and C. soluta would support the

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Spanton

monophyly of the species group.

Evolution of characters. — Some trends in the evolution of characters in this

group are worth noting. Three characters of the labrum were considered here

(characters 10, 11, 13). Species in the maritima group exhibit relatively short, and

flat labra. In the sylvatica group this tends toward a much longer labrum

(apotypically) in proportion to its width, especially in the clade which includes C.

nebraskana, C. longilabris, C. sylvatica and C. granulata It is interesting that the

species with the longest labra also exhibit more pronounced median ridges,

especially in C. sylvatica, in which the ridge is intensified into a sharp keel. The

longitudinal ridge in these species may function largely to strengthen a longer

labrum. It is also in this clade that a trend toward a dark colored labrum occurs. This

is most pronounced in C. sylvatica and C. nebraskana, although, as discussed

previously, some individuals of C. longilabris also possess a dark colored labrum.

The adaptive significance of a longer, dark colored labrum is unknown. As

mentioned above, C. lacteola also possesses a somewhat elongate labrum, which in

this reconstruction (Fig. 58) is assumed to have arisen independently. This is

assumed because the labrum of C. lacteola is not as elongate, nor is it associated

with a trend toward increase of a median ridge and dark color, as it is in the clade

which includes C. nebraskana, C. longilabris, C. sylvatica and C. granulata.

The most apotypic characters of the genitalia are found in species which are

plesiotypic with respect to labral characters. Most notable are the sister species C.

gemmata and C. soluta which, as mentioned above, share the apotypically notched

second gonapophyses (character 15) and the sclerite cluster (character 3, B) of the

internal sac of the male aedeagus. Pronounced autapotypies occur in the shape of the

apex of the eighth sternum (character 1 ) in females of C. soluta and C. lacteola, the

triangular-shaped oviduct sclerite (character 16) of females of C. gemmata and C.

japana, and in the shape of the apex of the aedeagus (character 2) in males of C.

lacteola. The functional significance of such genitalic characters has yet to be

suggested.

Independent and opposite states have evolved in the pattern of markings on the

elytral surface (characters 4, 5, 6) from the ancestral state in which each of the

markings is present and complete. In C. lacteola the markings have become

broadened and confluent. In several species of the group there is a trend toward

reduction in markings, with the humeral and apical lunules incomplete, present as

one or two dots. This is carried to an extreme in C. nebraskana in which the elytral

markings are absent, in most specimens. The pattern of markings on the elytral

surface is very labile. As noted previously, such characters are highly variable

within the species C. longilabris and may be subject to reversals and homoplasy

among species, thus great emphasis should not be placed upon such characters in

reconstructing a phylogeny.

The Cicindela sylvatica Group

149

59

Figs. 59-60. Approximate geographic distributions of the Palearctic species of the Cicindela sylvatica group.

59. C. sylvatica Linnaeus, C. granulata Gebler, and C. japana Motschulsky. 60. C. soluta Dejean, C.

lacteola Pallas, and C. gemmata Faldermann.

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Geographical History

Introduction. — I offer a few remarks concerning the possible history of

differentiation of the species C. nebraskana and C. longilabris and their immediate

relatives, according to the hypothesized phylogeny (Figure 58), C. sylvatica and C.

granulata. A detailed account of the evolutionary history of all members of the

group is not possible at present, as there is a paucity of information concerning the

distribution and habitat affinities of the Palearctic species.

The pattern. — The geographic ranges of C. nebraskana and C. longilabris are

presented in Figures 38 and 40, respectively. As mentioned previously, C.

longilabris is primarily a species of boreal forest and mixed coniferous/deciduous

forests, from Newfoundland to Alaska, north as far as tree line and south to the

southern limit of spruce {Picea spp.) and Jack Pine {Pinus banksiana) in the east,

and is also found in montane and subalpine forests in the mountains of western

North America as far south as Arizona and California. Cicindela nebraskana is

primarily a species of grasslands and forest/grassland transition zones in the western

prairies and in the Pacific Northwest. The approximate ranges of C. sylvatica and C.

granulata are presented in Figure 59. Cicindela sylvatica occurs from western

Europe (United Kingdom, France, and Scandinavia) east across coniferous forest

and mixed forest regions of the northern part of the Palearctic region to eastern

Siberia. Cicindela granulata occurs in a narrow band stretching in an east to west

direction, approximately coincident with the grassland regions of central Asia. The

approximate distribution of C. japana is also shown in Figure 59. Figure 60 shows

the approximate distributions of C. lacteola, C. gemmata, and C. soluta. Little is

known of the habitat affinities of these Palearctic species. Their collective ranges are

restricted to temperate regions.

Age of the C. sylvatica group. — The age of the C. sylvatica group is, in the

absence of a fossil record, largely speculative. Willis (1967) stated that the

cicindelids arose during the Permian, and that the genera of tiger beetles arose

during the Mesozoic. Freitag (1965) thought that the ancestral stock of the C.

maritima group was in existence in the early Tertiary. The warmest Cenozoic

climatic period for the northern hemisphere was during the mid Eocene, at which

time paratropical forests were thought to occur as far north as present day arctic

regions (Matthews, 1979a). Subsequent to the Eocene maximum, a gradual cooling

and increase in latitudinal climatic gradient are thought to have occurred,

culminating in the onset of glacial conditions in the northern Hemisphere during the

late Pliocene. As most species of the C. sylvatica group occur in the Palearctic

Region, it seems likely that this predominantly northern lineage had its beginnings

in the expanding temperate forests of the Palearctic region sometime subsequent to

the Eocene climatic maximum, perhaps during the Oligocene.

Differentiation of the species. — If we assume, as suggested above, an origin of

the group in Oligocene temperate forests of the Palearctic, then a common ancestor

of C. nebraskana, C. longilabris, C. sylvatica and C. granulata must have been in

The Cicindela sylvatica Group

151

existence in the late Oligocene or Miocene, Geological evidence suggests that an

Asian-Alaskan land connection has existed from the late Cretaceous to the Pliocene,

and it is thought that a boreal realm taxonomically more diverse than today’s Boreal

forest was Holarctic in extent in the Miocene (Matthews, 1979a), Such an ancestral

species could have had a Holarctic or Palearctic distribution in the Miocene, and the

divergence of the Nearctic lineage (C. nebraskana and C. longilabhs) and the

Palearctic lineage (C. sylvatica and C. granulata) could have been a result of either

a vicariant or dispersal event, probably also during the Miocene, but could have

occurred anytime prior to the the marine transgression of Beringia approximately 3

Mya.

The C. nebraskanalC. longilabhs and C. granulatalC. sylvatica species pairs

may represent parallel situations in which C. granulata in the Palearctic and C.

nebraskana in the Nearctic independently invaded grassland habitats, from a

coniferous forest inhabiting ancestor. This could have happened, in both instances

during late Miocene or early Pliocene, as it is thought that steppe habitats were in

existence in the interior of North America by the late Miocene (Matthews, 1979a,

Webb, 1977).

An alternate scenario would involve a single invasion of grasslands in either the

Palearctic or Nearctic Region, by a lineage which dispersed across Beringia during

the Quaternary, during glacial periods when steppe floral and faunal elements from

both Palearctic and Nearctic regions were present in Beringia (Matthews, 1982).

This would have to have been followed by vicariant speciation during an interglacial

period of the Quaternary when there was a marine barrier between Alaska and

Siberia. This latter scenario is less probable than the former, for two reasons. It

would imply that C. nebraskana is most closely related to C. granulata which is

inconsistent with the relationships indicated by structural characters of the species

(Figure 58), and also requires a speciation event during a Quaternary interglacial.

This seems unlikely in light of studies of fossil beetles which show that almost no

speciation has occurred through the course of the Quaternary period (Ashworth,

1979; Coope, 1977, 1979; Matthews, 1979a and b, 1980; Morgan and Morgan,

1980), and that only in Miocene and Pliocene deposits does one find fossils of

beetles which are specifically distinct from extant forms (Matthews, 1976, 1979b,

1980, 1982).

Similarly, it is unlikely that the ancestor of C. sylvatica, C. granulata, C.

longilabris, and C. nebraskana was a savannah/grassland inhabitant, as the

Beringian connection between Asia and the New World in the Late Tertiary was

predominantly a coniferous forest biome. Savannah and grasslands were found

farther south, in the interior of Asia and North America. As mentioned above, there

was an exchange of steppe biota via the Beringian steppe-tundra habitats of the

Pleistocene, but a steppe inhabiting ancestor would have had to expand across

Beringia in the Pleistocene, followed by a subsequent speciation event separating

the ancestor of C. longilabris! C . nebraskana from the ancestor of C. sylvaticaiC.

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granulata. Then two more, independent speciation events would have had to occur

to produce the four species we see today, presumably also in the Pleistocene. This

much speciation during the Pleistocene seems unlikely in light of fossil beetle

evidence which shows a remarkable constancy of form and lack of speciation in the

last two or more million years (Coope, 1979; Matthews, 1976, 1979a and b, 1980,

among others).

Differentiation within the Nearctic lineage. — Cicindela nebraskana probably

diverged from a common ancestor with C. longilabris sometime during the

Miocene, associated with the increase of grassland habitats in the interior of North

America. Subsequently, C. nebraskana has remained monobasic, showing relatively

little variation, compared with C. longilabris which is a polymorphic species. This

could be a result of C. nebraskana having passed the Quaternary glacial periods in

semi-arid regions of western North America, south of the ice fronts, in a contiguous,

if somewhat restricted geographic range possibly in the northern part of the Great

Basin region. Cicindela longilabris, in contrast, may have developed intraspecific

variation, and subsequent differentiation of subspecies as a result of fragmentation

of its range during glacial periods of the late Pliocene and Pleistocene. It is possible

that over the past two to three million years, during glacial periods, the range of C.

longilabris was compressed southward, and fragmented, with what is now the

nominate form C. /. longilabris having been isolated in spruce forests, which are

thought to have occurred in a belt across the eastern two-thirds of the continent,

south of the ice fronts (Watts, 1983; Ruhe, 1983). At the same time, populations

ancestral to C. /. laurentii might have been isolated in montane forests in the Rocky

Mountain region, and populations ancestral to C. /. perviridis were isolated in the

Sierra Nevada/Cascade Mountains in the Pacific region. Climatic conditions during

the relatively long glacial periods may have thus reduced genetic mixing among

what we now see as subspecific entities, while acting to maintain the genetic

continuity within the subspecies. For example, in the southern Rocky Mountains of

Arizona and New Mexico, Spaulding et al. (1983) have shown that some subalpine

tree species occurred during glacial episodes at elevations more than 1000 meters

below their present lower altitudinal limit. This would mean that some populations

of C. /. laurentii which are today isolated in forested areas of the Colorado Plateau,

the Wasatch and Uintah Mountains of Utah, the Abajo Mountains of southeastern

Utah, and the White Mountains of eastern Arizona may have had some genetic

contact during the periods of coldest climate via increased areas of coniferous forest

habitat. Thus the present pattern of geographic variation among and within the

subspecies of C. longilabris may have developed during Pleistocene climatic cycles

involving long glacial periods during which the subspecies of C. longilabris were

isolated, and populations within subspecies were genetically mixed, alternating with

relatively short interglacial periods during which the three subspecies mixed in a

hybrid zone, as seen today (Figure 39). Some temporary isolation of populations

occurred within subspecies (especially in the southern Rocky Mountain populations

The Cicindela sylvatica Group

153

of C. /. laurentii as seen today), and the range of the nominate subspecies C. /.

longilabris expanded northward with coniferous forest habitats.

CONCLUDING REMARKS

Questions requiring additional research have been presented in places throughout

the text. At the risk of some repetition I suggest the following.

Additional insight into the relationships between populations of C. longilabris,

especially in the western part of North America, relative to the interspecific

differences between C. longilabris and C. nebraskana might be gained by

electrophoretic studies. Additional field studies in the grassland-forest transition

regions of the western prairies, and in the Pacific Northwest might better elucidate

the nature of the ecological separation between the two species in question, and I

believe edaphic factors should figure prominently in such investigations. More

information concerning distribution, patterns of geographic variation, and habitat

affinities of the Palearctic species of the C. sylvatica group would aid greatly in

determining phylogenetic relationships in the group. Laboratory mating of adults

and rearing of larvae of the Palearctic species would add larval characters to the

information available for cladistic analysis of the group. Perhaps more importantly,

a rigorous phylogenetic analysis of the genus Cicindela is needed to test the

monophyly of the present supraspecific groupings. After such a cladistic framework

is established, phylogenetic analysis at the species group level will be on a much

more secure basis than at present.

ACKNOWLEDGEMENTS

I thank R. Freitag, sponsor of my M.Sc. program, for his advice and

encouragement during the course of this study, and the members of my advisory

committee, A. D. Macdonald, R. J. Ross, and J. P. Ryder, for their help and advice. I

thank my external examiner, G. E. Ball, and my colleagues J. H. Acorn and B. V.

Brown for constructive criticisms of the text.

I greatly appreciate the help and patience of the curators and private collectors

listed under “Materials”, who kindly loaned specimens. I thank A. Newton, T. L.

Erwin, and G. W. Byers for the courtesies extended during visits to their respective

establishments. Thanks are also extended to M. Palmer and to C. B. Knisely for their

courtesy and hospitality during visits, and for helpful discussions of rearing

techniques and field methods. I extend my thanks to W. Johnson and to D. Brzoska,

for kindly inviting me into their homes to examine collections of tiger beetles, and

to N. L. Rumpp, for sharing with me his unpublished metric data for male genitalia

of several populations of C. longilabris and C. nebraskana. I also appreciate the

many useful discussions of tiger beetle biology, evolution and taxonomy with M. M.

Kaulbars.

Quaest. Ent., 1988, 24 (1)

154

Spanton

Special thanks are due to Y. Spanton for technical assistance, and for much

patience throughout.

Financial support is gratefully acknowledged from the following sources: during

most of the duration of this study, from National Sciences and Engineering Research

Council Grant A4888 held by R. Freitag, and during the final stages of preparation

of this manuscript, from NSERC Grant A 1399 held by G. E. Ball.

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Issued May 1988

QUAESTIONES ENTOMOLOGICAE

ISSN 0033-5037

A periodical record of entomological investigation published at the Department

of Entomology, University of Alberta, Edmonton, Alberta.

Volume 24 Number 2 1988

CONTENTS

Smetana-Revision of the tribes Quediini and Atanyganthini. Part II. The

Himalayan region (Coleoptera: Staphylinidae) 163

Book Review-Jamieson, B. G. M. 1987. The Ultrastructure and

Phytogeny of Insect Spermatozoa 465

REVISION OF THE TRIBES QUEDIINI AND ATANYGNATHINI. PART II.

THE HIMALAYAN REGION (COLEOPTERA: STAPHYLINIDAE),^'

Ales Smetana

Biosystematics Research Institute

Agriculture Canada

Ottawa ON KIA 0C6 Quaestiones Entomologicae

CANADA 24:163-464 1988

ABSTRACT

A revision of the staphylinid tribes Quediini and Atanygnathini (Coleoptera) of

the Himalayan region is presented including a historical review, a discussion of

diagnostic characters and teclmicpies, and some remarks, mostly on the

zoogeography of the group.

Nine genera (8 in Quediini and 1 in Atanygnathini) and 104 species (96 in

Quediini and 8 in Atanygnathini) are recognized. Forty one species (36 in Quediini

and 5 in Atanygnathini) are described as new, new combinations are proposed for 5

species and new status for 1 species.

Quedius leptocephalus Coiffait 1982 is placed in synonymy with Q. inquietus

Champion 1925; Quedius flavoterminatus Cameron 1932 and Q. sherpai Coiffait

1982 with Q. apicicornis Eppelsheim 1895; Quedius mimeticus Cameron 1932, Q.

notabilis Cameron 1932 and Q. peraffinis Cameron 1932 with Q. beesoni Cameron

1932; Quedius analoka Smetana 1975 and Q. queinneci Coiffait 1983 with Q.

stevensi Cameron 1932; Quedius sodalis Cameron 1932 with Q. ripicola Cameron

1926; Quedius trisulensis Coiffait 1982 with Q. franzi Smetana 1975; Quedius

mandra Smetana 1975 and Q. deuvei Coiffait 1983 with Q. kashmirensis Cameron

1944; Quedius kasmiricus Cameron 1932, Q. nichinaiensis Coiffait 1982 and Q.

cachemiricus Coiffait 1982 with Q. fluviatilis Cameron 1932; Quedius decipiens

Cameron 1944 with Q. aureiventris Bernhauer 1915; Quedius dohertyi Cameron

1932 and Q. heterogaster Cameron 1944 with Q. muscicola Cameron 1932;

Quedius lama Coiffait 1982 and Q. dhaulagirensis Coiffait 1982 with Q. kanyasa

Smetana 1975; Quedius mussooriensis Cameron 1932 with Q. rugosus Cameron

1921 ; Quedius coeruleus Coiffait 1977 with Q. gardneri Cameron 1932; Quedius

akalita Smetana 1977 and Algon cyanipennis Coiffait 1982 with Indoquedius

sikkimensis (Cameron 1932); Acylophorus ventralis Coiffait 1983 with A. beesoni

Cameron 1926; Acylophorus ruficollis Kraatz 1859 with A. ruficollis Motschulsky

1858; Atanygnathus ruficollis Kraatz 1859 with A. pictus Motschulsky 1858 and

( ] 33rd contribution to the knowledge of Staphylinidae).

^Part I is the revision of the North American Quediini (Smetana 1971).

164

Smetana

Atanygnathus fuscus Kraatz 1859 with A. piceus Motschulsky 1858.

Heterothops flavicollis Motschulsky 1858 is transferred to the genus Gabronthus

Tottenham 1955 (comb.novj and the name Philonthus pulchellus Kraatz 1859

becomes a junior synonym of it fsyn.novj. Heterothops magniceps Bernhauer 1920

is transferred to the genus Ctenandropus (comb.novj; Quedius juno Sharp 1874

and Q. praeditus Sharp 1889 are transferred to the genus Indoquedius (comb.nov.J

Cyrtothorax vulneratus Fauvel 1878 is transferred to the genus Bolitogyrus

(comb.nov.j, and Acylophorus nepalicus Coiffait 1981 is transferred to the genus

Erichsonius Fauvel 1874 (comb.nov.) and placed tentatively in synonymy with E.

basalis Motschulsky 1858. Heterothops indicus Cameron 1926 is removed from

synonymy with H. oculatus Fauvel 1895 and reinstated as a separate species

(stat.nov.).

Lectotypes are designated for Quedius beesoni Cam., Q. mimeticus Cam., Q.

notabilis Cam., Q. adjacens Cam., Q. placidus Cam., Q. ripicola Cam., Q. sodalis

Cam., Q. chatterjeei Cam., Q. fluviatilis Cam., Q. muscicola Cam., Q. heterogaster

Cam., Q. anomalus Cam., Q. assamensis Cam., Q. aureipilis Cam., Q. rugosus Cam.,

Q. mussooriensis Cam., Q. intricatus Fauv., Q. gardneri Cam., Q. lineipennis Cam.,

Q. bipunctatus Epp., Q. oculatus Fauv., Heterothops oculatus Fauv., H. indicus

Cam., H. hindustanus Cam., H. persimilis Cam., Ctenandropus nigriceps Cam.,

Paratolmerus pilosiventris Cam., Acylophorus beesoni Cam., A. microcephalus

Cam., A. furcatus Motsch., A. flavipes Motsch., A. tibialis Cam., Anchocerus

monticola Cam., Atanygnathus ruficollis Kr., A. pictus (Motsch.), A. brevicollis

Fauv., A. piceus (Motsch.) and A. fuscus (Kr.).

A key to the two tribes, and keys to all genera and species are provided. The

tribes and genera are described; each species is described and illustrated, all

available bionomic and distributional data are presented and distributional records

are mapped.

RESUME

Une rhnsion des especes des trihus Quediini et Atany gnat hint (Coleoptera, Staphylinidae) de la region

Himalayenne, incluant une revue historique, une discussion des caracteres importants et des techniques,

ainsi que certaines remarques sur la hiogeographie du groupe, est presentee.

Neuf genres (8 parmi les Quediini et 1 parmi les Atanygnatliini) et 104 especes (96 parmi les Quediini et

8 parmi les Atanygnathini) sont reconnues. Quarante et une nouvelles especes (36 parmi les Quediini et 5

parmi les Atanygnathini) sont decrites, de nouvelles comhinaisons sont proposees et le statut d'une espece

est redefinit.

Quedius leptocephalus Coiffait 1982 est placee en synonymie avec Q. inquietus Champion 1925);

Quedius flavoterminatus Cameron 1932 et Q. sherpai Coiffait 1982 avec Q. apicicornis Eppelsheim 1895;

Quedius mimeticus Cameron 1932, Q. notabilis Cameron 1932 et Q. peraffinis Cameron 1932 avec Q.

beesoni Cameron 1932; Quedius analoka Smetana 1975 et Q. queinneci Coiffait 1983 avec Q. stevensi

Cameron 1932; Quedius sodalis Cameron 1932 avec Q. ripicola Cameron 1926; Quedius trisulensis Coiffait

1982 avec Q. franzi Smetana 1975; Quedius mandra Smetana 1975 et Q. deuvei Coiffait 1983 avec Q.

kashmirensis Cameron 1944; Quedius kasmiricus Cameron 1932, Q. nichinaiensis Coiffait 1982 et Q.

cachemiricus Coiffait 1982 avec Q. fluviatilis Cameron 1932; Quedius decipiens Cameron 1944 avec Q.

aureiventris Bernhauer 1915; Quedius dohertyi Cameron 1932 et Q. heterogaster Cameron 1944 avec Q.

Revision of the Tribes Quediini and Atanygnathini

165

muscicola Cameron 1932; Quedius lama Coijfait 1982 et Q. dhaulagirensis Coiffait 1982 avec Q. kanyasa

Smetana 1975; Quedius mussooriensis Cameron 1932 avec Q. rugosus Cameron 1921 ; Quedius coeruleus

Coiffait 1977 avec Q. gardneri Cameron 1932; Quedius akalita Smetana 1977 et Algon cyanipennis Coiffait

1982 avec Indoquedius sikkimensis (Cameron 1932); Acylophorus ventralis Coiffait 1983 avec A. beesoni

Cameron 1926; Acylophorus ruficollis Kraatz 1859 avec A. ruficollis Motschulsky 1858; Atanygnathus

ruficollis Kraatz 1859 avec A. pictus Motschulsky 1858 et Atanygnathus fuscus Kraatz 1859 avec A. piceus

Motschulsky 1858.

L’espece Heterothops flavicollis Motschulsky 1858 est transferee au genre Gabronthus Tottenham 1955

(nouv. comh.) et le nom Philonthus pulchellus Kraatz 1859 devient un synonyme plus recent de Gabronthus

flavicollis (Motschulsky) (nouv. syn.). Heterothops magniceps Bernhauer 1920 est transferee au genre

Ctenandropus (nouv.comh.), Quedius juno Sharp 1874 et Q. praeditus Sharp 1889 au genre Indoquedius

(nouv. comh.), Cyrtothorax vulneratus Fauvel 1878 au genre Bolitogyrus (nouv.comh.) et Acylophorus

nepalicus Coiffait 1981 au genre Erichsonius Fauvel 1874; cette derniere espece est tentativement placee en

synonvmie avec E. basalis Motschulsky 1858. Heterothops indicus Cameron 1926, auparavant considere

comme un synonyme plus recent de H. oculatus Fauvel 1895, est traite comme une espece valide (nouv.

stat.).

Un lectotype est designe pour chacune des especes suivantes: Quedius beesoni Cam., Q. mimeticus

Cam., Q. notabilis Cam., Q. adjacens Cam., Q. placidus Cam., Q. ripicola Cam., Q. sodalis Cam., Q.

chatterjeei Cam., Q. fluviatilis Cam., Q. muscicola Cam., Q. heterogaster Cam., Q. anomalus Cam., Q.

assamensis Cam., Q. aureipilis Cam., Q. rugosus Cam., Q. mussooriensis Cam., Q. intricatus Fauv., Q.

gardneri Cam., Q. lineipennis Cam., Q. bipunctatus Epp., Q. oculatus Fauv., Heterothops oculatus Fauv., H.

indicus Cam., H. hindustanus Cam., H. persimilis Cam., Ctenandropus nigriceps Cam., Paratolmerus

pilosiventris Cam., Acylophorus beesoni Cam., A. microcephalus Cam., A. furcatus Motsch., A. flavipes

Motsch., A. tibialis Cam., Anchocerus monticola Cam., Atanygnathus ruficollis Kr., A. pictus (Motsch.), A.

brevicollis Fauv., A. piceus (Motsch.) et A. fuscus (Kr.).

Ce travail inclut des tahleau.x de determination pour les trihus, genres et especes. Les tribes et genres

sont decrits; chaque espece est decrite et illustree, les donnees sur I'ecologie et la repartition geographique

sont indiquees et des cartes de distribution sont fournies.

TABLE OF CONTENTS

Introduction 166

Historical Review 167

Dignostic Characters 169

Techniques 170

Systematics 178

1. Tribe Quediini 179

1. Genus Quedius Stephens 181

2. Genus Indoquedius Cameron 300

3. Genus Bolitogyrus Chevrolat 314

4. Genus Heterothops Stephens 318

5. Genus Ctenandropus Cameron 334

6. Genus Paratolmerus Cameron 335

7. Genus Acylophorus Nordman 337

8. Genus Anchocerus Fauvel 361

2. Tribe Atanygnathini 368

1. Genus Atanygnathus Jacobson 368

Conclusions 380

Acknowledgements 383

Quaest. Ent., 1988, 24 (2)

166

Smetana

References 385

Figures 391

Index 459

INTRODUCTION

My interest in the tribe Quediini of the Himalayan region started some 12 years

ago while working on the quediine material collected in Nepal by Dr. H. Franz.

Although the material was rather limited, it nevertheless clearly indicated that 1) the

Himalaya harbors a rich fauna of this tribe, and 2) the species from higher elevations

(above 3000 m) are of temperate origin and show obvious relationships to the

east-palaearctic fauna, particularly to species described from higher mountains in

Sichuan in China.

Some time later, during my first joint collecting trip to the Nepal Himalaya with

my friend Ivan Ldbl, Geneve, Switzerland, I realized that the Quediini fauna and, as

a matter of fact, the entire fauna of the family Staphylinidae of the middle and

higher elevations throughout the Himalaya is even less known that I suspected. It

was during the long strenuous treks of the first trip that I decided to go ahead with

the revision of the tribe.

This revision was originally intended to include only the species occurring in the

Nepal Himalaya. But, as work on the revision progressed, it became obvious that the

only way to do the project right was to include the fauna of the entire Himalaya,

since the original material of all species described from throughout the Himalaya

had to be examined anyway. In addition, the material collected in Nepal, particularly

during my five collecting trips (three together with Ivan Lobl and one together with

my wife), formed by far the most numerous and diverse material available from the

area, so that the expansion to include the entire Himalaya did not actually add

enough additional material to substantially increase the work required to complete

the project.

The only previous treatment of the tribes from the Himalaya was that by

Cameron (1932) within his treatment of the family Staphylinidae in the series “The

Fauna of British India ....”. When using Cameron’s keys and descriptions in this

book, I suiprisingly often ran into problems in interpreting the species. Later, after

revising the types, I realized, that, oddly enough, many of Cameron’s comparisons

of the Himalayan species with well known European species were misleading and

some quite wrong. His species and generic descriptions omitted important characters

and many of the species were assigned to the wrong genus or subgenus. These

problems significantly contributed to my difficulty in using his works and to the fact

that many of Cameron’s species were later redescribed by those authors who did not

study the older types (see also Historical Review). However, in spite of these

criticisms, Cameron’s contribution to the knowledge of the family Staphylinidae,

particularly from the Oriental region, is enormous. The reason for the criticism is

Revision of the Tribes Quediini and Atanygnathini

167

purely practical: to emphasize that the study of the types is ""conditio sine qua non\

a condition without which there is no solid base for taxonomic work. This statement

cannot be overemphasized.

The present study incorporates all available information about the systematics,

geographical distribution, and bionomics of adults. The larvae of the Himalayan

species are virtually unknown.

In the Himalayan region (the term is used here freely and has nothing to do with

the zoogeographical faunal regions - see the discussion under Techniques) the tribe

Quediini consists of 8 genera and 96 species, and the tribe Atanygnathini of 1 genus

and 8 species. The Himalaya has recently become a focus of interest and a lot of

collecting is being done there. This is fortuitous, because, considering the rate of

destruction of the forests in Nepal, quite a few species are likely to disappear before

they can be discovered.

I hope that this revision will stimulate interest in this group and that additional

collecting in the future will provide the additional material needed to fill in many of

the gaps that still exist in our knowledge of these two tribes in the Himalayan

region.

HISTORICAL REVIEW

A few papers, dealing to various extents with the Quediini and Atanygnathini of

the Himalayan region have been published in the past.

The two widely distributed Palaearctic species, Quedius fulgidus and Q.

ochripennis do not really belong to the Himalayan fauna and are not considered in

this review. I also do not consider the species Quedius chlorophanus and Q.

spectahilis described from India by Erichson (1840) and Kraatz (1859) respectively,

and Acylophorus microcerus Eauvel 1895 from Burma (see more in the discussion

following the description of Quedius).

In 1858 Motschulsky (1858 a,b) described several species of the genera

Acylophorus and Atanygnathus from “Indes orientates”. Three of them, Acylophorus

furcatus, ruficollis and Atanygnathus pictus occur in the Himalayan region and

are the first quediines and atanygnathines to be described from this area. The species

Acylophorus flavipes and Atanygnathus piceus probably do not occur in the

Himalaya; their actual distributional ranges are not known.

In 1859 Kraatz described Acylophorus ruficollis, Atanygnathus ruficollis and A.

fuscus from “Ceylan” and/or “India orientalis”. Two of them, Acylophorus ruficollis

and Atanygnathus ruficollis, occur in the Himalayan region; all three names are

junior synonyms of species described by Motschulsky in 1858.

In 1878 Eauvel treated the genus Cyrtothorax (synonym of Bolitogyrus); one of

the species he described from “Cochinchine”, B. vulneratus, occurs in the

Himalayan region.

Quaest. Ent., 1988, 24 (2)

168

Smetana

In 1895 Eppelsheim (1895 a,b) described Quedius filicornis, Q. hipunctatus and

Q. apicicornis, all from the Himalayan region. The first two species belong to the

genus Indoqiiedius.

In 1895 Fauvel, in his series on new species of Staphylinidae from India and

Malaysia, described the species Quedius oculatus, Q. inthcatus, Heterothops

oculatus, Acylophorus puncticeps, and Atanygnathus hrevicollis. All species are

valid; three of them occur in the Himalayan region and two (Q. inthcatus and A.

puncticeps) are still known only from the original specimens collected in Burma.

Quedius oculatus belongs to the genus Indoquedius.

In 1905 Fauvel erected the genus Anchocerus for one new species, A. hirmanus,

from Burma.

In 1915 Bernhauer described the species Quedius himalayicus and Q.

aureiventris from the Himalaya (Kumaon).

In 1922 and 1925 respectively Champion described the species Quedius conicus

and Velleius inquietus, both from Kumaon. The former species belongs to the genus

Indoquedius and is a junior synonym of Q. filicornis, the latter species belongs to

the subgenus Microsaurus of Quedius.

In 1926 Cameron, in his series on new staphylinids from India, erected the genus

Ctenandropus for the single new species C. nigriceps and described the species

Heterothops indicus, Quedius sodalis, Q. adjacens, Q. ripicola, Q. fluviatilis, Q.

anomalus, Q. chatterjeei, Acylophorus heesoni and Anchocerus monticola all of

them from the Himalayan region. Heterothops indicus was combined with H.

oculatus (Cameron 1932), but is in fact a distinct species. The species Q. sodalis and

Q. ripicola proved to be identical.

In 1932 Cameron treated both tribes (as one tribe Quediini) in the series “The

Fauna of British India...”. His treatment included the genera Heterothops,

Ctenandropus, Algon Sharp 1874, Securipalpus Schubert 1908, Baijpalpus

Cameron 1932, Cyrtothorax (synonym of Bolitogyrus), Velleius Mannerheim 1831,

Quedius, Acylophorus, Anchocerus and Atanygnathus', many new species were

described, particularly in the genus Quedius. The genera Algon and Securipalpus

(congeneric with Rientis Sharp 1874) were subsequently excluded from Quediini

(Smetana 1977 b) as is now the genus Barypalpus (see the discussion following the

Quediini). The genus Velleius does not occur in the Himalayan region, the only

species included by Cameron in fact belongs to the subgenus Microsaurus of

Quedius (see above).

In 1944 Cameron described additional species from the Himalayan region:

Quedius kashmirensis, Q. flavocaudatus, Q. deceptor, Q. decipiens and Q.

heterogaster. Quedius decipiens proved to be identical with Q. aureiventris and Q.

heterogaster with Q. muscicola.

In the following 30 years nothing was published on the subject until Smetana

(1975) gave results of the study of the Quediini collected in Nepal by H. Franz in

1970 and 1971. Only the genera Quedius and Acylophorus were represented in this

Revision of the Tribes Quediini and Atanygnathini

169

material. Ten new species of the genus Quedius were described in this paper; two of

them were found to be identical with species previously described by Cameron:

Quedius analoka with Q. stevensi and Q. mandra with Q. kashmirensis.

In 1976 Scheerpeltz (1976b) described the species Q. cyanopterus from Nepal. It

proved to be a junior synonym of Q. apiciconus.

In 1977 Smetana (1977a) presented the results of the study of the Quediini of the

Bhutan Expedition 1972 of the Naturhistorisches Museum in Basel, Switzerland. All

species in this paper were recorded from Bhutan for the first time. Quedius

(Indoquedius) akalita and Anchoeenis punctatissimus were described as new; the

former species was found to be identical with Indoquedius sikkimensis.

In 1977 Coiffait presented the results of the study of the subfamilies

Xantholininae and Staphylininae of the same expedition to Bhutan. He described

one new species of Quedius: Q. coeruleus. It proved to be identical with Q. gardneri

previously described by Cameron.

In the early eighties Coiffait (1981, 1982 a,b, 1983 a,b) described within a series

of papers on the Staphylinidae of the Himalayan region 15 new species in the genera

Quedius (10), Algon (1) (the species is identical with Indoquedius sikkimensis),

Heterothops (2) and Acylophorus (2). Except for the two species of Heterothops and

one species of Quedius (Q. daksumensis) all other species were found to be identical

with species described by previous authors. The larger number of synonyms created

was to some extent caused by the fact that Coiffait relied too much on the keys and

descriptions of Cameron, which, after the study of Cameron’s material and types,

proved to be very inaccurate and partly misleading (see also Introduction).

DIAGNOSTIC CHARACTERS

The diagnostic characters of the Quediini were discussed in detail in my revision

of the tribe Quediini of America north of Mexico (Smetana 1971:3-6). I refer the

user to the above paper for all the information he may need.

In my 1971 revision I did not discuss the sclerites of the male and female genital

segments, since they were not studied at that time. Subsequently (Smetana

1976:176-178) the possible importance of the tergite 10 and sternite 9 of the male

genital segment was demonstrated (wrongly referred to as tergite 10 and sternite 7).

Since both the male and female genital segments are referred to in this revision, I

briefly discuss them below.

The male genital segment consists of tergite 9, which is divided into two

sclerites, each narrowed distally into a stylus, a fairly small tergite 10 located

between the two sclerites of tergite 9, and of fairly large sternite 9 closing the genital

segment ventrally (Figs. 42, 171, 297, 308, 324, 351). Both tergite 10 and sternite 9

are thickly sclerotized (Figs. 219, 237, 238, 271, 272), except they are

semimembranous in Ctenandropus and obsolete in Atanygnathus (Figs. 306, 357).

In some genera the styli of tergite 9 each bear one strong or, more or less strong.

Quaest. Ent., 1988, 24 (2)

170

Smetana

apical spines (Figs. 308, 324) or spiniform setae (Fig. 351 ) and their setation may be

modified by being either very strong or long, or both (Figs. 171, 297, 306). The

shape of tergite 10 and sternite 9, particularly the condition of their apical margins

(e.g., rounded, truncate or emarginate - Figs. 219, 237, 238) seems to provide useful

characters.

The female genital segment consists of tergite 9, which is divided into two

sclerites, each narrowed distally into a stylus; a fairly large tergite 10 located

between the two sclerites of tergite 9 (both conditions similar to those of the male

genital segment), and the ventral gonocoxites 1 and 2, with each of the second

gonocoxites having a minute papilla-like stylus bearing a strong and long apical

seta; each stylus of tergite 9 usually bears two strong and very long apical setae

(Figs. 384-387, 392-399). The condition of the apical portion of tergite 10 (Figs.

385, 393) seems to be of importance.

Unlike in my 1971 revision (see above), the abdominal segments are numbered

morphologically, therefore the 6th visible segment is segment 8 and is the segment

immediately preceding the genital segment (see above).

The measurements of body parts were taken in the same way as described in my

1971 revision.

TECHNIQUES

Material

I attempted to accumulate for this revision as many specimens from all known

sources, as was reasonably possible. Undoubtedly there are specimens housed

somewhere that I did not see; however, I am sure that the material at hand is the

largest and most complete collection ever assembled from the Himalaya. In addition

to the fairly large collection from the British Museum (Natural History), London,

which was mostly collected by G.C. Champion and M. Cameron in northern India

prior to Second World War, the vast majority of my material comes from the

expeditions to northern India of Drs. C. Besuchet and I. Lobl from the Museum

d’Histoire Naturelle, Geneve, Switzerland, and from my expeditions to the Nepal

Himalaya, most of them undertaken together with my friend I. Lobl from the above

institution. At least two thirds of the available material came from these two

sources. The value of this material is accentuated by the fact that it is accompanied

by detailed field notes on habitats, et cetera.

The material at hand totalled about 3600 specirnens.

Revision of the Tribes Quediini and Atanygnathini

171

Quaest. Ent., 1988, 24 (2)

Map 1. Countries and Indian States of the area treated in this revision.

172

Smetana

Map 2. 1981 expedition to Nepal, collection localities. 1, above Sundarijal, 2000 m. 2, Chaubas, 2500 m,

2600 m. 3, Burlang Bhanjyang, 2600 m. 4, Chipling, 2300 m. 5, Gul Bhanjyang, 2600 m. 6. Kutumsang,

2200-2700 m. 7, near Mere Dara, 3000 m. 8, Mere Dara, 3100-3300 m. 9, ridge between Mere Dara and

Thare Pati, 3500 m. 10, below Thare Pati, 3300-3500 m. 11, Malemchi, 2750-2900 m. 12, below Tarke

Ghyang, 2000 m. 13, Tarke Ghyang, 2750 m. 14, above Tarke Ghyang, 3000-3400 m. 15, Yangri-top, 4150

m. 16, Yangri Ridge, saddle Phedi, 3800 m. 17, Yangri Ridge, 4200-4800 m. 18, above Shermathang, 2900

m. 19, Kakani, 2200 m. 20, Tarang Marang, 1000 m. 21, 4 km S Tarang Marang, 900 m. 22, 3 km N

Bahunepati, 900 m. 23, Barahbise, 1600 m. 24, NE Barahbise, 2500 m. 25, Dobate Ridge NE Barahbise,

2700-3000 m. 26, below Jangtang Ridge NE Barahbise, 3150 m. 27, Jangtang Ridge NE Barahbise,

3250-3300 m.

Area

The paper treats the fauna of the following area: 1) The great mountain system of

the Himalaya, extending some 2500 km between the rivers Indus in the west and

Brahmaputra in the east and covering an area more than 590,000 square kilometers.

Included are from north to south: a) Great Himalaya, or the main range, with a

crest-line of about 6000 m. b) Lesser Himalaya with altitudes of about 2100-4600

m, running parallel and sometimes merging with the main range and enclosing some

large valleys, like Vale of Kashmir in the west, or the Nepal Valley in the central

portion; the range Mahabharat Lekh, with the highest mountain Phulcoki (2762 m),

closing the Kathmandu Valley in the south, belongs to this system, c) Outer

Himalaya with altitudes of about 700-1500 m and consisting mainly of the Siwalik

Range, which in Nepal is known as the Churia Range. 2) Also included are the hill

ranges in northeastern India, such as the Khasi Hills and the Garo Hills in

Revision of the Tribes Quediini and Atanygnathini

173

Map 3. 1982 expedition to Nepal, collection localities. 1, Dunge Dara N of Tumlingtar, 1100 m. 2,

Khandbari, 1700 m. 3, Pangma, 1900 m. 4, Forest above Ahale, 2250-2400 m. 5, Kuwapani, 2100 m. 6,

Forest NE Kuwapani, 2450-2550 m. 7, Num, 2000 m. 8, Arun River at Num, 1500-1600 m. 9, below

Sheduwa, 2100-2550 m. 10, Sheduwa, 2800 m. 11, above Sheduwa, 3000-3050 m. 12, “Bakan” west of

Tashigaon, 3200-3250 m. 13, above Tashigaon, 3100-3600 m.

Meghalaya, or Naga Hills in Nagaland, as long as material for study was available

from them. 3) For practical reasons also species from Burma, treated by Cameron

(1932) in the series The Fauna of British India..., are included. However, not

included are species described from Burma after Cameron’s publication, particularly

those described by Scheerpeltz (1965). The mountains in Burma are not part of the

Himalaya proper, although those in northern Burma are in every respect an eastern

extension of the Himalaya. There is also a practical reason why Scheerpeltz’s

species are not treated. Not only would their inclusion further extend the extent of

this revision, but also, there are difficulties in interpreting these species most of

which were described from single specimens (many of them females), or without

considering the characters on the aedoeagus (the slides with the aedoeagi and the

drawings were apparently lost during the confused times around the end of the

Second World War, see Scheerpeltz 1965:93-95). The interpretation of these

species should be attempted within the framework of a revision of the Oriental

Quediini and after the accumulation of recent collections from Burma.

Quaest. Ent., 1988, 24 (2)

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Smetana

Map 4. 1983 expedition to Nepal, collection localities. 1, Turture Dara, 800 m. 2, Tarku, 850 m. 3,

Besisahar, 900 m. 4, Bahudanda, 1300 m. 5, below Tal, 1500 m. 6, Tal, 1600 m. 7, Forest W Bagarchhap,

2200-2250 m. 8, Latha Manang W Bagarchhap, 2350-2450 m. 9, 4 km SE Pisang, 3050 m. 10, 2 km SE

Pisang, 3050 m. 11, Manang airstrip, 3200 m. 12, Manang to Thorong Pass, 2600-4000 m. 13, below

Thorong Pass, east side, 4400 m. 14, Thorong Pass, west slope, 4300-4550 m. 15, Jomsom, 2750 m. 16, 3

km SE Marpha, 2650 m. 17, 2 km N Kalopani, 2550 m. 18, Lete, 2550 m. 19, Ghoropani Pass, N slope,

2700-2750 m. 20, Ridge E Ghoropani Pass, 3100-3150 m. 21, Pun Hill at Ghoropani Pass, 3050-3100 m.

22, Ghoropani Pass, 2850 m.

A few species (Acylophorus flavipes, A. hipunctatus and Atanygnathus piceus)

were included here for practical reasons (to treat all the species included in Cameron

1932); they either do not {Acylophorus hipunctatus), or very likely do not occur in

the Himalayan region.

Distributional records and maps

All records and the general distribution given for each species are based, with

very few exceptions, on specimens that I have seen.

The geographic distribution of each species is also shown on maps, but again

only records of specimens that I have examined are included.

Specimens under “Type material” do not appear again under the “Distribution”,

except where such a specimen represents the only record from a particular province

or state.

Revision of the Tribes Quediini and Atanygnathini

175

Map 5. 1984 expedition to Nepal, collection localities. 1, Pangnia, 1700 m. 2, Forest above Ahale, 2200 m.

3, Sangrati, 2 km SW Kuwapani, 2250 m. 4, Forest NE Kuwapani, 2250-2400 m. 5, Pass NE Mangmaya,

2300 m. 6, Ridge NE Mangmaya, 2800 m. 7, Ridge S Mansingma, 2600-2800 m. 8, Goru Dzure Dara, W

slope, 3600 m. 9, Goru Dzure Dara, E Slope, 3350 m. 10, Forest S Mansingma, 2200-2300 m. 1 1, Induwa

Khola Valley, 1700-2150 m. 12, Induwa Khola Valley, 2600-2850 m. 13, 2 km E Mansingma, 2900 m. 14,

Arun Valley at Num main bridge, 1000-1 150 m. 15, above Num, 1900-2200 m.

Map 1 shows all countries and states of India in the area treated in this revision,

to allow quick general orientation.

There were some difficulties in listing some records because of a) different ways

of transcription from the native languages, b) because different collectors related

some collecting sites to different settlements and c) because of difficulties in

obtaining detailed maps of many areas, names provided by native people were used,

but were subsequently found unusable since they were not to be found on any maps

or in any available lists. This is particularly true for some of the recent collecting

sites in Nepal.

The most important cases are listed below:

1) All Nepalese localities from the 1982 and 1984 trips, listed as in Khandbari

District, are correctly in Sindhupalchok District. I used Khandbari because it is a

well known center on the main trail from the Tumlingtar airstrip to the Tibetan

border. I continue to use the term Khandbari District, because it is on all locality

Quaest. Ent., 1988, 24 (2)

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Smetana

Map 6. 1985 expedition to Nepal, collection localities. 1, 1.5 km N Trisuli Bazar, 550 m. 2, above

Bokhajhundo, 1950 m. 3, 1.5 km NE Bhargu, 2000 m. 4, Langtang Khola Valley, 2.5 - 3.5 km E Syabru,

1720-1750 m. 5, Langtang Khola Valley, Forest Camp, 1900-1950 m. 6, Shin Gumba above Dhunche,

3300-3400 m. 7, North slope above Syabru, 3600-3900 m. 8, Gosaikund area, Larabina Pass, 4000-4100 m.

9, Gosaikund Lakes, 4200-4450 m. 10, between Gosaikund and Ghopte, 3600 m. 11, between Ghopte and

Thare Pati, 3100-3250 m. 12, Thare Pati - Mere Dara, 3200-3600 m. 13, Siwapuri Dara, 2300-2550 m.

labels attached to the beetles.

2) The locality “Forest above Ahale” in Khandbari District (1982 and 1984 trips)

is identical with the locality “Chichila s. Ahale” of other collectors;.

3) The locality “Dobate Ridge NE Barahbise” (1981 trip) should correctly read

“Pokhare NE Barahbise” (not to be confused with the city of Bokhara). The term

“Dobate Ridge” was provided by native people, but it is otherwise unknown.

4) The locality “Jangtang Ridge NE Barahbise” (1981 trip) should correctly read

“Yardang Ridge NE Barahbise”.

5) The localities “Arun Valley at Num main bridge” and “Val. Arun ss/Num”

(1984 trip) are identical. They both refer to the area around the large bridge across

the river Arun just north of Num, and are not to be confused with the narrow

hanging bridge across Arun west of Num on the trail to Sheduwa and further on to

the Makalu base camp.

Revision of the Tribes Quediini and Atanygnathini

177

Maps 2-6 give all collection localities of the 1981-1985 expeditions to allow

easy location of any of them.

Illustrations

All illustrations were drawn to scale in transmitted light by using a

Prado-Universal microprojector (Leitz, Wetzlar). Some details shown in the

drawings, such as minute setae on the parameres or on the median lobe, or the

details of the internal sac of the aedoeagus of some taxa cannot be seen clearly

through a stereomicroscope. A compound microscope is recommended for the study

of these details.

All photographs of body parts were taken at the facilities of the Electron

Microscope Centre, Chemistry and Biology Research Institute, Ottawa, with the aid

of an AMR 1000 scanning electron microscope at an accelerating voltage of 10 KV.

The parts to be photographed were routinely coated with gold.

Type material

I was able to study the original material of almost all previously described taxa

included in this revision. The exceptions are Quedius spectahilis Kraatz 1859,

Quedius chlorophanus Erichson 1840, and Acylophorus microcerus Eauvel 1895. I

was unable to locate the type material of these species and it is possible that these

specimens have been destroyed or lost. I did not attempt to study the type material

of the two well known Palaearctic species Q. fulgidus and Q. ochhpennis.

The detailed information on the type material of all the respective species is

included under “Type material” following the species description.

The type material of the species described as new in this revision is deposited in

the collections of the Institutions from which it was originally sent out for study.

The type material of the species I collected in Nepal is deposited in the section of

my own collection in Ottawa I still own (subfamilies Staphylininae and

Xantholininae). It is readily available for study; this portion of my collection will

eventually be deposited in the Museum d’Histoire Naturelle, Geneve, Switzerland.

Miscellaneous

This revision is based exclusively on adults. Therefore, whenever a “species” is

compared with another “species” or a “genus” with another “genus”, it is understood

that the adults of the two particular species, or the adults of the species of the two

particular genera, are being compared. Similarly, if under Bionomics A-us h-us is

said to occur in such and such habitats, it is the adults of A-us h-us that prefer these

habitats.

In the references for each genus and species (except for original references) only

papers pertaining to the area included in this revision, and only subgenera and/or

synonyms based on specimens from the area are included. However, at least one

paper giving the complete references and synonymies is mentioned at appropriate

Quaest. Ent., 1988, 24 (2)

178

Smetana

place in the revision.

The manuscript of this revision was concluded in May 1986. Any information

published after this date is not included.

The Institutions which loaned material for this revision are listed below together

with the abbreviations used in the text when referring to the material studied. The

assistance of the curators responsible for the loan of the specimens from their

respective institutions is gratefully acknowledged.

BMNH British Museum (Natural History), London, Mr. M.E. Bachus, P.M.

Hammond, Mrs. S.L. Shute

CNCC Canadian National Collection of Insects, Ottawa

FMNH Field Museum of Natural History, Chicago, Dr. L.F. Watrous

MHNG Museum d’Histoire Naturelle, Geneve, Dr. I. Lobl

NHMB Naturhistorisches Museum, Basel, Dr. M. Brancucci

SMFM Senckenberg Museum, Frankfurt a.M.. Dr. R. zu Strassen

USNM United States National Museum, (now known as National Museum of

Natural History), Smithsonian Institution, Washington, D.C., Dr. P.J.

Spangler

The following individuals also generously loaned or donated material from their

private collections:

HCCC H. Coiffait, collection in Museum National d’Histoire Naturelle, Paris,

France

HFCC Dr. H. Franz, Modling, Austria

SMFM Dr. J. Martens, Mainz, Federal Republic of Germany (material in SMFM)

GDRC Mr. G.M. de Rougemont, London, England

SYSTEMATICS

The two tribes Quediini and Atanygnathini were treated by most modem authors

as one tribe, the Quediini. However, most recently, I have pointed out (Smetana,

1984:279) that the genus Atanygnathus actually does not exhibit any of the

characters of the Quediini and that a separate tribe should be restored for it (see

Smetana, /.c., and page 368 for more details). The two tribes are easily separated

using the following key:

1 Tarsal formula 5,4,4. Maxillary and labial palpus conspicuously

elongate Atanygnathini, p. 368

1' Tarsal formula 5,5,5. Maxillary and labial palpus not

conspicuously elongate Quediini, p. 179

Revision of the Tribes Quediini and Atanygnathini

179

1. Tribe Quediini

Ganglhauer 1895:378; Cameron 1932:264 (r-.v Smetana 19776:178-180: 1984:279.

The tribe is characterized by; front angles of pronotum distinctly produced

beyond anterior margin of prosternum; inflexed lateral portions of front pronotal

angles markedly inflexed, meeting prosternum at sharp, almost 90° angle in some

taxa; anterior margin of prostemum clearly below level of lower surface of pronotal

front angles, both sclerites separated by suture; prothoracic hypomera (epipleura)

markedly inflexed and therefore not visible in lateral view; superior marginal line of

prothoracic hypomera not curved ventrally in apical portion of prothorax and

therefore situated entirely on dorsal surface; inferior line of prothoracic hypomera, if

present, usually disappearing before reaching front margin of procoxal cavities and

therefore not meeting superior marginal line; head with infraorbital ridge ventrally,

at least on basal portion of head (ridge rarely missing).

The above characters of Quediini were discussed in detail (Smetana

1977b; 178-180) and I refer the reader to this paper for more information and

illustrations.

In the genera Quedius, Indoquediiis and BoUtogynis the aedoeagus rests in the

abdomen rotated 90°, resting on its right side, that is with the proximal opening and

the paramere on left side (the orientation of the aedoeagus was unfortunately not

recorded for Quedius gardneri and Q. lineipennis). On the other hand, in the genera

Heterothops, Ctenandropus, Pciratolmerus and Acylophorus the aedoeagus rests in

the abdomen with the ventral side (where the proximal opening and the paramere

are) facing ventrally.

Unlike in my previous treatments of this tribe (Smetana 1958, 1971), the

Quediini here do not include the genus Atanygnathus which is believed to form a

separate tribe (see Smetana 1984;281 and page 368 in this revision).

The tribe is represented by many genera in all zoogeographical regions, eight

genera occur in the Himalayan region. Several genera, assigned to Quediini by

Cameron (1932), in fact do not belong there. The genera Rientis Sharp 1874

iSecuripalpus Schubert 1908) and Algon Sharp 1874 were already assigned to the

tribe Staphylinini, Subtribe Xanthopygi (Smetana 1977b; 180) and the same is true

for the genus Barypalpus Cameron 1932. The single species, B. ruficornis Cameron

1932, most likely belongs to the genus Rientis.

Key to genera of Quediini

1 Antenna not geniculate (Figs. 380, 388, 401 ) 2

1' Antenna geniculate (Fig. 409) 6

2 (F) Last segment of maxillary palpus very small, thin and sharp,

subulate; at base only about half as wide as penultimate

segment at apical margin and much shorter than penultimate

Quaest. Ent., 1988,24 (2)

180

Smetana

2'

3 (2)

3'

4 (2')

4'

5 (4')

5’

6 (!’)

6’

segment 3

Last segment of maxillary palpus not subulate, at base more

than half as wide as penultimate segment at apical margin and

about as long or longer than penultimate segment 4

Head constricted before neck; infraorbital ridge present. Middle

tibia spinose on lateral face

4. Genus Heterothops Stephens, p. 318

Head not constricted before neck; infraorbital ridge absent.

Middle tibia not spinose on lateral face

5. Genus Ctenandropus Cameron, p. 334

Middle tibia not spinose on lateral face. First five segments of

antenna lacking fine and dense pubescence, each bearing only

strong and long setae. Frons behind antennal insertions with

V-shaped impression. Posterolateral and basal margins of

pronotum markedly and abruptly explanate

3. Genus BoUtogyrus Chevrolat, p. 314

Middle tibia markedly spinose on lateral face. First three

segments of antenna lacking fine and dense pubescence, each

bearing only strong and long setae (Figs. 380, 388, 401). Frons

behind antennal insertions without V-shaped impression (Fig.

379). Posterolateral and basal margins of prontoum no more

than slightly and gradually explanate 5

Two or three setiferous punctures between anterior and

posterior frontal punctures situated directly at median margin of

eye (Fig. 400). Dorsal surface of head and pronotum without

microsculpture (microsculpture rarely present in lateral

pronotal groove). Dorsal rows of pronotum each with two

punctures 2. Genus Indoquedius Cameron, p. 300

No setiferous punctures between anterior and posterior frontal

punctures (Fig. 379); if, rarely, punctures present, then

separated from median margin of eye by distance at least equal

to diameter of puncture. Dorsal surface of head and pronotum

with distinct microsculpture and dorsal rows of pronotum each

with at least three punctures; if, rarely, dorsal surface of head

and pronotum without microsculpture, then dorsal rows of

punctures on pronotum absent

1. Genus Quedius Stephens, p. 181

Middle and hind tarsus with empodial setae short, hardly

visible between claws and considerably shorter than claws 7

Middle and hind tarsus with empodial setae long, distinctly

protruding between claws and at least as long as claws, but

usually longer (Fig. 413)

Revision of the Tribes Quediini and Atanygnathini

181

7. Genus Acylophorus Nordmann, p. 337

7 (6) Middle tibia with numerous strong and long spines on lateral

face. First antennal segment lacking fine and dense

pubescence 8. Genus Anchocerus Fauvel, p. 361

7' Middle tibia with one or two fine and short spines on lateral

face. First antennal segment with fine and dense pubescence

6. Genus Paratolmerus Cameron, p. 335

1 . Genus Quedius Stephens ^

QuecU US Stephens 1829:22; Cameron 1932:280.

Subg. Microsaurus Dejean 1 833:61 ; Stephens 1835:435; Cameron 1932;281.

£r//t//<»5 Mulsant et Rey — 1875:616; Cameron 1932:281.

Subg. Distichalius Casey 1915:398, 404.

Subg. /?<7p/;//7/5 Stephens 1829:23; Cameron 1932:281.

Sauridus Mulsant et Rey 1875:700; Cameron 1932:281.

Type species: Staphyliniis tristis Gravenhorst 1802, designated by Curtis 1837,

pi. 638.

Descriptive notes. — A conventional formal description of this genus is presented

by Smetana (1971:40) and Coiffiat (1987:9). I discuss some unconventional

characters, particularly those used to distinguish higher taxa of the region covered in

this revision.

The head has on dorsal side a set of setiferous punctures characteristically

located (chaetotaxy); they were described and named in Smetana 1971 and I refer

the reader to this paper for details. In general, there are no additional setiferous

punctures present at medial margin of the eye between the anterior and posterior

frontal punctures. They are present only in one species (Q. taruni), but unlike in

species of Indocpiedius, they are separated from medial margin of the eye by a

distance at least equal to the diameter of the puncture. In a few taxa, there are also

other additional punctures present around the posterior frontal puncture {e.g., Q.

franzi), but those are far away from the margin of the eye. Additional setiferous

punctures may be present between the two anterior frontal punctures {e.g., Q.

kashmirensis). The frons behind the antennal insertions is even and lacks the

V-shaped impression characteristic of species of the genus Bolitogyrus. The eyes

vary considerably in size, ranging from minute and flat and considerably shorter

than tempora to very large and convex, occupying most of the lateral portions of the

head. The dorsal surface of the head bears fine microsculpture; it is missing only in

Q. lineipennis, which differs also in other characters from all other species (see

below and page 300). The three basal antennal segments lack fine and dense

pubescence; the large setae on antennal segments, particularly on those past segment

3, are relatively short and moderately strong (Fig. 380, 388). The penultimate

Only subgenera and references pertaining to the area treated in this revision are given. See Smetana (1971:

38-40) for a complete synonymy with all references up to 1970.

Quaest. Ent., 1988, 24 (2)

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Smetana

segments of both the maxillary and labial palpi bear only a few fine setae.

The pronotum is usually narrowed anteriorly and evenly transversely convex; its

posterolateral and basal margins are occasionally slightly and gradually explanate.

The dorsal rows on pronotum are each composed of three punctures, rarely of four

punctures {Q. tanmi), or are entirely missing {Q. Uneipennis). The scutellum is

impunctate, variably punctate, or impunctate but bearing irregular transverse rugae

on the basal portion. Elytra vary considerably in length, in correlation with the

development of the wings. They are usually variably, simply punctate, occasionally

each elytron bears three inconspicuous longitudinal rows of three or four coarser

punctures.

On the legs the front tarsi are sexually dimorphic, with four first segments

slightly to considerably (Fig. 381) more dilated in the males than in the females.

Middle and hind tibiae are spinose on lateral face. The empodial setae of all tarsi are

short, considerably shorter than claws (Fig. 389).

On the abdomen the tergites are simple, rarely the first three visible tergites bear

a median basal keel {Q. gardneri). The tergite 7 bears a whitish apical seam of

palisade setae in winged species; brachypterous and apterous species lack this seam

(Figs. 382, 383, 391). Female tergite 8 is simple, without a median notch. Male

sternite 8, and occasionally also sternites 7 and 6, bear secondary sexual characters

consisting of an emargination in the middle of the apical margin (Fig. 18) and a

small flattened and smooth triangular area before it.

Male genital segment with tergite 10 and sternite 9 not emarginate apically,

except emarginate in Q. Uneipennis; styli of tergite 9 blunt apically, without strong

apical spine (Fig. 171), rarely bearing dense and strong, conspicuously long setae

(Fig. 42). Female genital segment with tergite 10 obtusely rounded or arcuate

apically (Figs. 384, 392).

Aedoeagus with median lobe symmetrical or occasionally asymmetrical with

parameres present. The median lobe never bears microsetae apically. The parameres

are fused in one solid sclerite, rarely divided anteriorly in two branches (Fig. 233).

The paramere usually bears sensory peg setae on the surface adjacent to the median

lobe; these setae vary considerably in number and arrangement. They are missing

from some assemblages of species {e.g., Q. beesoni, Q. antennalis, Q. apicieornis,

or Q. hpicola and Q. milansaar, and Q. kailo, Q. tanderi and Q. angnimai). These

assemblages of species, based on other characters, do not seem to be closely related;

the loss of the sensory peg setae on the paramere therefore apparently occured

independently several times during the evolution. The paramere bears variably

developed setae on the apical portion. There are typically two pairs of apical setae

and a pair of subapical setae on each lateral margin below the apex (Figs. 5, 9, 31);

subapical setae are sometimes missing {Q. placidus - Fig. 38), or they are sometimes

shifted apically to form with the apical setae one apical group {e.g., Q. beesoni or Q.

tanderi - Figs. 13, 68). In addition to the normal setae, ususlly there are microsetae

at lateral margins (Figs. 94, 102), occasionally also on the surface adjacent to the

Revision of the Tribes Quediini and Atanygnathini

183

median lobe {e.g., Q. inquietus or Q. heesoni - Figs. 5, 13). The internal sac of

aedoeagus is in general simple and lacks large heavily sclerotized structures (Figs.

47,57).

The genus Quedius has more than 600 species occurring in all zoogeographical

regions. Fifty-seven species are at present known from the Himalayan region.

Classification and comparisons. — I use here the three “classical” subgenera of

Quedius: Microsauriis, Distichalius and Raphirus as in Smetana (1971). However, it

should be mentioned that the concepts of these (and also other) subgenera of

Quedius, which were established based on limited faunas, need to be revised; other

sets of characters will have to be found to meaningfully define those that really

deserve the subgeneric status. The currently used characters, such as the size of the

eyes, the character of the elytral punctation, etc, clearly are not sufficient. The

difficulties to characterize and delimit these subgenera grow with the size of the

geographical area included in the study. New characters perhaps can be found in the

development of the male and female genital segments and in the re-examination and

re-evaluation of the chaetotaxy of the head and pronotum, particularly that of the

head. For example, all Palaearctic, Nearctic and also Himalayan species belonging

to the subgenus Microsaurus always have two setiferous punctures posteromediad

of the posterior frontal puncture (Fig. 379), whereas all species belonging to the

subgenus Raphirus (in my sense, which includes Sauridus) have only one puncture

posteromediad of posterior frontal puncture (Fig. 421 ). The species belonging to the

subgenus Distichalius (characterized mainly by the presence of the two additional

setiferous punctures between the anterior frontal punctures) also have, just like those

of the subgenus Microsaurus, two punctures posteromediad of the posterior frontal

puncture. Also, the presence and absence of other setiferous punctures on the head,

such as additional punctures between the anterior frontal punctures, or between the

anterior and posterior frontal puncture on each side of the head may be more

significant than considered at present (see also Bordoni 1973:42, etc.).

These are just a few simplified thoughts that need more exploration and

correlation with other possible characters. Should these ideas prove correct, they

would lead to the reassignment of many species to different subgenera.

Within the fauna treated in this paper, two species are quite isolated and their

assignment to existing subgenera poses serious problems. I believe that their

subgeneric or perhaps generic assignment can be properly assessed only within a

study involving possibly the entire Oriental fauna. These species are 1 ) Q. gardneri,

assigned here tentatively to the subgenus Raphirus, and 2) Q. lineipennis which is

not assigned to any subgenus (see also the discussions following the descriptions of

either of the species).

Two species of Quedius: Q. spectahilis Kraatz 1859 and Q. chlorophanus

Erichson 1840, are not treated in this revision, because their types are apparently

lost and their correct interpretation is therefore not possible. Quedius spectahilis was

described from “India borealis”, apparently from one single female. It is probable

Quaest. Ent., 1988, 24 (2)

184

Smetana

that one of the species related to Q. apicicornis, possibly Q. heesoni (of which some

specimens have antennae and abdomen uniformly black), is in fact synonymous

with it. Quediiis chlorophanus was described from Bengal (“Habitat in Bengalia”);

it very likely does not occur in the Himalayan region; I also have some doubts about

the assignment of this species to Quedius, although Erichson (l.c.) compares its

general habitus to that of Q. fulgidus.

Key to subgenera of Quedius

1 Pronotum with dorsal rows of punctures, each row with three or

rarely more punctures; or, rarely, dorsal rows irregular and

formed by very coarse pit-like punctures. Clypeus without

extremely fine punctures 2

r Pronotum without dorsal rows of punctures. Clypeus with

extremely fine punctures

Q. lineipennis Cameron (subgenus ?), p. 297

2 (1) First three abdominal tergites with elevated median keel

3. subg. Raphirus Stephens (pars) (Q. gardneh), p. 295

2' None of abdominal tergites with elevated median keel 3

3 (2’) Head, pronotum and elytra bright metallic green or dark green,

bluish-green or purplish-green. Head with deep and coarse

punctures forming more or less distinct rugae on posterior

portion, or at least with numerous coarse punctures

posteromediad of each eye

3. subg. Raphirus Stephens (pars), p. 231

3' Head, pronotum and elytra at most with metallic reflections.

Head with scattered setiferous punctures 4

4 (3') No additional setiferous punctures between anterior frontal

punctures 5

4' Two additional setiferous punctures between anterior frontal

punctures 2 subg. Distichalius Casey, p. 227

5 (4) Two or three additional punctures on each side along medial

margin of eye between anterior and posterior frontal punctures

3 subg. Raphirus Stephens (pars) {Q. taruni), p. 271

5' No additional punctures between anterior and posterior frontal

punctures 6

6 (5') Two or rarely three setiferous punctures between posterior

frontal puncture and posterior margin of head (Fig. 379)

(occasionally one puncture missing unilaterally). Eyes small to

moderately large, only occasionally distinctly larger than

tempora 1. subg. Microsaurus Dejean, p. 185

6' Only one puncture between posterior frontal puncture and

Revision of the Tribes Quediini and Atanygnathini

185

posterior margin of head (fig. 421). Eyes large, distinctly to

considerably longer than tempora

3. subg. Raphirus Stephens (pars), p. 231

1. Suhgenus Microsaurus Dejean

Type species: Staphylinus lateralis Gravenhorst 1802, fixed by Westwood

(1838:16).

The subgenus is used here in general in the same sense as in Smetana ( 1971:49).

However, see the discussion following Quedius (.s', lat.) for the ever increasing

difficulties to satisfactorily characterize and delimit the conventional “classical”

subgenera of Quedius.

Taxonomic notes. — I do not recognize species-groups in this subgenus, for

similar reasons as discussed under Acylopliorus. As there, in Microsaurus there are

assemblages of species linked together by some characters, but 1 found it impossible

to meaningfully characterize them as definite species-groups. I had a distinct feeling

that there are many gaps and/or missing links which possibly could be found in

purely tropical faunas of other areas of the Oriental region.

Quedius inquietus is fairly isolated by the markedly incrassate antennae with

outer segments subserrate; this was the main reason why the species was originally

placed in the genus Velleius by Champion ( 1925: 107).

The species Q. apicicornis, Q. heesoni (most specimens), Q. flavocaudatus, Q.

antennalis and Q. hirmanus share the character state of the pale tips of antenna and

the pale tip of abdomen, and, except for Q. apicicornis, the character state of

scutellum bearing irregular transverse rugae. Most species in this assemblage share

the character state of the absence of the sensory peg setae on the underside of the

paramere {Q. apicicornis, Q. heesoni, Q. antennalis and possibly Q. hirmanus)',

however, the sensory peg setae are present on the paramere of Q. flavocaudatus. The

two species Q. apicicornis and Q. heesoni are very similar externally and also share

some characters on the aedoeagus, but Q. apicicornis lacks the transverse rugae on

the scutellum, characteristic of all other species of this assemblage.

The species Q. martensi, Q. dui, Q. adjacens, Q. ochripennis and Q. fulgidus are

an obviously artificial assemblage which is difficult to characterize; they do not fit

in any other assemblage. They are mostly of the habitus of a “classical

Microsaurus"' with more or less small eyes, except for Q. adjacens with moderately

large eyes.

Quedius placidus and Q. lesagei are unique by the presence of irregular

longitudinal rows of coarser punctures on the elytra.

The species Q. stevensi, Q. ripicola and Q. milansaar are characterized by the

very deep and coarse punctation of the elytra. They are quite similar to each other in

all external characters; however, despite this similarity, Q. stevensi differs from both

other species by the drastically different, unique shape of the aedeoagus and by the

Quaest. Ent., 1988, 24 (2)

186

Smetana

styli of tergite 9 of the male genital segment bearing dense, strong and

conspicuously long setae (Fig. 42).

The remaining five species Q. franzi, Q. goropanus, Q. taiideri, Q. kailo and Q.

angnimai are brachypterous with more or less short elytra and with abdominal

tergite 7 lacking the whitish apical seam of palisade setae. They are endemic

Himalayan species with more or less restricted ranges, occurring at elevations above

3000 m.

Key to species of Microsaurus

1 Antenna markedly incrassate, segment 4 transverse, outer

segments subserrate. Aedoeagus as in Figs. 2-5. Length

10.5-13.0 mm 1. Q. inquietus (Champion), p. 189

1' Antenna not markedly incrassate, segment 4 about as long as

wide, outer segments not subsen'rate. Aedoeagi different 2

2 (F) Scutellum with sculpture consisting of irregular transverse

rugae in middle of basal portion 3

2' Scutellum smooth, without sculpture of irregular transverse

rugae on basal portion 6

3 (2) Dorsal rows on pronotum each with four punctures. Elytra red.

Aedoeagus as in Figs. 15, 16. Length 12.0-13.3 mm

4. Q. flavocaudatus Cameron, p. 200

3' Dorsal rows on pronotum each with three punctures. Elytra

never red (occasionally reddish-piceous in teneral specimens).

Aedoeagi different 4

4 (3') Abdominal tergite 3 (first visible) with smooth impunctate area

in middle. Sublateral rows on pronotum each with two to four

punctures, last puncture situated behind level of large lateral

puncture. If only two punctures present (usually only

unilaterally), posterior puncture may be situated at level of

large lateral puncture. Aedoeagus as in Figs. 11-13. Length

8.0-12.0 mm 3. Q. heesoni Cameron, p. 196

4' Abdominal tergite 3 (first visible) evenly punctate, without

smooth impunctate area in middle. Sublateral rows on

pronotum each with one or two punctures, posterior puncture

(if present) situated no further back than about level of large

lateral puncture 5

5 (4') Large lateral puncture on pronotum situated not far from lateral

margin, separated from it by distance about equal to width of

apex of antennal segment 2. Aedoeagus as in Figs. 17, 19, 20.

In general smaller and less robust species. Length 1 1.0 mm

5. Q. antennalis Cameron, p. 201

Revision of the Tribes Quediini and Atanygnathini 187

5' Large lateral puncture on pronotum situated very far from

lateral margin, separated from it by distance almost twice as

large as width of apex of antennal segment 2. Male unknown.

In general larger and more robust species. Length 12.0 mm

6. Q. hinnanus Cameron, p. 202

6 (2') Abdominal tergite 7 (fifth visible) with whitish apical seam of

palisade setae. Elytra relatively long, at suture about as long as

and at sides variably longer than pronotum at midline. Fully

winged species 7

6' Abdominal tergite 7 (fifth visible) without whitish apical seam

of palisade setae. Elytra short, both at suture and at sides

variably shorter than pronotum at midline. Brachypterous species 17

7 (6) Posterior frontal puncture situated much closer to posterior

margin of head than to posteromedian margin of eye.

Aedoeagus as in Figs. 25-27. Length 8.7 mm

8. Q. did spec, now, p. 204

7' Posterior frontal puncture situated at least slightly closer to

posteromedian margin of eye than to posterior margin of head.

sometimes quite close to margin of eye 8

8 (7') Punctation of elytra not uniform, each elytron with three

inconspicuous longitudinal rows of three or four coarser punctures 9

8' Punctation of elytra uniform, without longitudinal rows of

coarser punctures 10

9 (8) Eyes small, tempora much longer than length of eyes seen from

above (ratio 1.66). Aedoeagus as in Figs. 36-38. Length

7. 8-8.0 mm 12. Q. placidus Cameron, p. 210

9' Eyes large, tempora considerably shorter than length of eyes

seen from above (ratio 0.23). Aedoeagus as in Figs. 40, 41.

Length 5. 7-6. 5 mm

13. Q. lesagei spec, now, p. 21 1

10 (8') Sublateral rows on pronotum each with no more than two

punctures, posterior puncture situated before level of large

lateral puncture 1 1

10' Sublateral rows on pronotum each with at least three punctures,

posterior puncture situated behind level of large lateral

puncture (rarely last puncture missing unilaterally) 12

11 (10) Posterior frontal puncture situated close to posteromedian

margin of eye, separated from it by distance somewhat larger

than diameter of puncture. Eyes large and convex, about as

long as tempora seen from above. Elytra at least slightly dark

bluish or greenish-blue. Aedoeagus as in Figs. 7-9. Length

7.1-12.0 mm 2. Q. apicicornis Eppelsheim, p. 192

Quaest. Ent., 1988, 24 (2)

188

Smetana

1 1 ' Posterior frontal puncture situated far from posteromedian

margin of eye, just slightly closer to posteromedian margin of

eye than to posterior margin of head. Eyes small and rather flat,

considerably shorter than tempora seen from above. Elytra

piceous-black. Aedoeagus as in Eigs. 22, 23. Length 10.0 mm ..

7. Q. martensi Smetana, p. 203

12 (10') Posterior frontal puncture situated away from posteromedian

margin of eye, separated from it by distance at least twice as

large as diameter of puncture 13

12' Posterior frontal puncture situated quite close to posteromedian

margin of eye, separated from it by distance not longer than

diameter of puncture or actually touching it; no additional

setiferous puncture between it and posteromedian margin of eye 15

13 (12) One additional setiferous puncture between posterior frontal

puncture and posteromedian margin of eye 14

13' No additional setiferous puncture between posterior frontal

puncture and posteromedian margin of eye. Elytra red. Length

7.0-1 1.5 mm 1 1. Q. fiilgidiis (Eabricius), p. 209

14 (13) Elytra red. Aedoeagus as in Eigs. 33, 34. Length 8.0-10.2

mm 10. Q. ochripemiis Menetries, p. 207

14' Elytra piceous-black. Aedoeagus as in Eigs. 29-31. Length

9.5-1 1.2 mm 9. Q. adjacens Cameron, p. 205

15 (12') Aedoeagus conspicuously large and voluminous, paramere very

wide basally, markedly narrowed anteriorly (Eig. 44). Length

6. 9-8. 6 mm 14. Q. stevensi Cameron, p. 212

15' Aedoeagus narrow and elongate, paramere of different shape

(Eigs. 46^9, 51-54) 16

16 (15') Apical portion of median lobe of aedoeagus forming short

rounded hook in lateral view (Eig. 48). Length 6. 8-8. 3 mm

15. ripicola Cameron, p. 215

16' Apical portion of median lobe of aedoeagus forming large,

elongate hook in lateral view (Eig. 53). Length 7. 5-8. 3 mm

16. Q. mdansaar spec, now, p. 217

17 (6') Sublateral rows on pronotum each with three or four punctures,

posterior puncture situated distinctly behind level of large

lateral puncture 1 8

17' Sublateral rows on pronotum each with only two punctures,

posterior puncture situated before level of large lateral

puncture. Head with one additional setiferous puncture

anteromediad and occasionally another additional setiferous

puncture posteromediad of posterior frontal puncture.

Aedoeagus as in Eigs. 74-77. Length 7. 6-9. 5 mm

Revision of the Tribes Quediini and Atanygnathini

189

2\. Q. angnimai spec, nov., p. 223

18 (17) Head with two or three additional setiferous punctures

posteromediad and usually one anteriad of posterior frontal

puncture; posteromedian additional punctures form with two

usual punctures near posterior margin of head an irregular

group of four or five punctures. Aedoeagus as in Figs. 56-59.

Length 5.8-7. 1 mm 17. Q. franzi Smetana, p. 217

18' Head without additional setiferous punctures near posterior

frontal puncture

19 (18') Median lobe of aedoeagus, except for small lateral lobe on each

side just below middle, entirely covered by paramere; paramere

with sensory peg setae forming two lateral groups situated well

posteriad of apex of paramere (Figs. 61-64). Male sternite 8

with three strong setae on each side (Fig. 60). Length 6. 9-7. 4

mm 18. Q. goropcmus Smetana, p. 220

19' Median lobe of aedoeagus without small lateral lobe on each

side, entirely covered by paramere except for apex; paramere

without sensory peg setae (Figs. 66-68, 70-72). Male sternite 8

with four strong setae on each side (Fig. 65)

20 (19') Apical narrowed portion of median lobe short; paramere with

four apical setae at each side of median emargination (Figs. 67,

68). Length 6. 9-7.4 mm 19. Q. tanderi spec, now, p. 222

20' Apical narrowed portion of median lobe long; paramere with

three apical setae at each side of median emargination (Figs.

71, 72). Length 7. 1-7.3 mm 20. Q. kailo spec, nor., p. 223

19

20

1. Quediiis (Microsaurus) inquietus (Champion)

Figs. 1-5; Map 1

Velleius inqiiietiis Champion 1925: 107; Cameron 1932:279.

Quedius leptocephaliis Coiffait 1982b:276 (syn.nov.)

Description. — Deep black, head no more than feebly iridescent, pronotum and abdomen iridescent;

appendages black, tarsi usually slightly paler, rather piceous- black to piceous. Head in male naiTower than

pronotum, of rounded quadrangular shape, wider than long (ratio 1.2), parallel-sided behind eyes, with

obtuse posterior angles. Head in female small and relatively narrow, much narrower than pronotum, as long

as wide to feebly longer than wide (ratio 1.08), parallel-sided to slightly naiTowed behind eyes, posterior

angles quite obtuse and hardly noticeable; eyes small or moderately large (female) and only slightly

protruding from lateral outlines of head, tempora distinctly (male, ratio 1.6) or slightly (female, ratio 1.3)

longer than length of eyes seen from above; no additional punctures between anterior frontal punctures;

posterior frontal puncture situated closer to posteromedian margin of eye than to posterior margin of head,

two additional setiferous punctures between it and posterior margin of head; temporal puncture situated

closer to posterior margin of head than to posterior margin of eye; deflexed portion of tempora with

numerous fine punctures bearing rather short but stiff setae; surface of head with very dense and fine

microsculpture of transverse waves. Antenna short and stout, incrassate anteriorly, segments 2 and 3 short,

segment 2 only feebly to slightly longer than wide, segment 3 slightly stronger and about as long as to

slightly longer than segment 2, segment 4 slightly transverse, following segments becoming increasingly

Qiiaest. Ent., 1988, 24 (2)

190

Smetana

transverse and subsenate, last segment shorter than 2 preceding segments combined. Pronotum wider than

long (ratio 1.2), broadly rounded basally and markedly narrowed anteriorly, with lateral portions somewhat

explanate, disc markedly transversely convex; dorsal rows each with three fine punctures; sublateral rows

each with two punctures, posterior puncture situated before level of large lateral puncture; microsculpture of

pronotum similar to that of head, but still finer and denser. Scutellum large, usually impunctate, but in some

specimens with two or three fine punctures with setae comparable in length to those on elytra. Elytra rather

long, at base hardly narrower than pronotum at widest point, at suture slightly longer (ratio 1.2), at sides

distinctly longer (ratio 1.4) than pronotum at midline; punctation and pubescence fine and dense, pubescence

deep black, surface between punctures without microsculpture. Wings fully developed. Abdomen with

tergite 7 bearing distinct whitish apical seam of palisade setae, punctation and pubescence of abdominal

tergites similar to that on elytra, but slightly sparser, pubescence deep black. Front tarsus dilated in both

sexes.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with three

long setae on each side; apical margin with rather shallow, obtusely triangular emargination and numerous,

dark and long setae (Fig. 1 ), small triangular area before emargination flattened and smooth. Aedoeagus

(Figs. 2-5) small, both median lobe and paramere asymmetrical apically; median lobe obtuse apically; apical

part, when paramere removed, with two subapical hooks markedly projecting ventral ly in lateral view.

Paramere not quite reaching apex of median lobe, with four apical setae, median pair distinctly longer than

lateral pair, and with one pair of setae at each lateral margin rather far below apex; underside of paramere

distinctly excavate, with only two sensory peg setae at each lateral margin below apex and with numerous

extremely fine setae.

Length 10.5-13.0 mm.

Type material. — Quedius inqidetus. Champion (1925:107) described the species

from a single specimen from Kumaon. The male holotype in the British Museum

(Natural History), London, is labelled as follows: “3109” (label upside down)/“Type

H.T.” (round label with red margin)/ “Sunderhunga V.W. Almora Divn.

8000-12000 feet June ’19 H.G.C.”/ “G.C. Champion. Brit. Mus. 1925^2.” (label

upside down)/ “E.M.M. 1925. det. G.C.C.” (label upside down)/ “Velleius inquietus.

Champ.”/ “Velleius himalayanus type Ch.”. The specimen was dissected, and the

aedoeagus and sternite 8 were mounted into Canada Balsam; tergite 8 and the

genital segment were glued to plate with beetle.

Quedius ieptocephaliis. Coiffait (1982b:276) described the species from a single

female specimen from Nepal. It is deposited in the Naturhistorisches Museum,

Basel, Switzerland, and is labelled as follows: “Manigow lO.VI. 1200-1 900 m”/

“Nepal 1978 Bhakta B. Ch.”/ “TYPE”/ “Quedius (Microsaurus) leptocephalus H.

Coiffait 1981”. This female agrees in all essential characters with the male holotype

of Q. iuqiiietus\ the name Q. leptocephalus is a junior synonym of Q. inquietus. The

specimen was accordingly labelled.

Geographical distribution. — Quedius inquietus is at present known from Uttar

Pradesh, West Bengal and from Nepal (Map 7).

Material studied. — 4 specimens.

INDIA. Uttar Pradesh. Kumaon (see Type material of Q. inqidetus). West Bengal. Gonglu-Garibas,

3050 m, 7. VI.75, W. Wittmer (NHMB ) 1 .

NEPAL. Chandam Bari, 3350 m, 22. VI. 78, B. Bhakta (NHMB) 1.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons and variations. — Quedius inquietus is very distinctive due to the

very short, incrassate antenna with outer segments subserrate, in combination with

the deep black colouration of the body, and the small head in the female sex. It

Revision of the Tribes Quediini and Atanygnathini

191

Quaest. Ent., 1988, 24 (2)

Map 7. Distribution records for: Quedius inquietus ( A. )\Q. adjacens ( ^)\Q. martensi ( • ); and Q. dm ( ■ ).

192

Smetana

cannot be confused with any other species of Quedius occurring in the Himalayan

region. The antenna of Q. inquietus resembles to some extent that of the species of

the genus Velleius. However, the antennal segments 5-10 are not pectinate as in the

species of Velleius, the scutellum is impunctate, the general habitus and the type of

the aedoeagus is that of a typical Microsaunis. There seems to be little doubt that

the species belongs to the subgenus Microsaurus of Quedius. Champion (1925:108)

himself assigned it to Velleius with some doubts.

The holotype of Q. inquietus is a male, not a female as suspected by Champion

(1925:108).

In the female of Q. inquietus, the head is distinctly narrower and more narrowed

posteriorly behind the eyes, and the eyes are larger than in the male. However, the

female head looks narrower than it actually is (see the description).

2. Quedius (Microsaurus) apicicornis Eppelsheim

Figs. 6-9, 379-387; Map 8

Quedius upicicovnis 1 895b:39 1 ; Cameron 1932:286; Smetana 1975:325; Coiffait 1982a:32

Quedius flavoterminatus Cameron 1932:290 (syn.uov.)

Quedius cyanopterus Scheerpeltz 1976b: 150; Coiffait 1982a:32

Quedius sherpai Co'\tEd\i 1982a:77 {syn.uow)

Descnption. — Black, head and pronotum usually feebly to distinctly, abdomen markedly iridescent,

elytra usually more or less dark bluish, apical third of abdominal segment 7 (fifth visible) and apex of

abdomen yellowish-red, occasionally segment 7 only vaguely paler apically and also base of segment 8

darkened, rarely almost entire abdomen dark; palpi brownish to piceous; antennae piceous-black, usually

with segments 8-10 pale yellowish to whitish-yellow with last segment variably darkened, either partially or

entirely; occasionally also segment 7 partially or entirely pale, and/or last segment entirely pale, or only

segments 9-10 pale; legs piceous-black to black with more or less paler tarsi. Head (Fig. 379) in general of

rounded quadrangular shape, about as long as wide, moderately to distinctly narrowed posteriorly behind

eyes, posterior angles rounded and indistinct; eyes convex and prominent, tempora about as long as length of

eyes seen from above; no additional punctures between anterior frontal punctures; posterior frontal puncture

situated close to posteromedian margin of eye, but separated from it by distance distinctly larger than

diameter of puncture; two additional setiferous punctures between it and posterior margin of head; temporal

puncture situated closer to posterior margin of eye than to posterior margin of head; surface of head with

very fine and dense microsculpture of transverse waves becoming gradually irregular anteriorly on clypeus.

Last segment of maxillary palpus much longer than penultimate segment, slightly attenuate basally. Antenna

moderately long and only slightly thickened toward apex, segment 3 much longer than segment 2 (Fig. 380),

segments 4-7 longer than wide, gradually becoming shorter, outer segments about as long as wide,

occasionally slightly longer than wide, last segment shorter than two preceding segments combined.

Pronotum about as long as wide, broadly rounded basally, slightly to distinctly narrowed anteriorly, rather

markedly transversely convex, lateral portions hardly to slightly explanate laterally, lateral margins evenly

arcuate, occasionally somewhat flattened basally; dorsal rows each with three fine punctures, sublateral rows

each with only two punctures, posterior puncture situated before level of large lateral puncture;

microsculpture on pronotum similar to that on head, but still finer and denser. Scutellum large, impunctate,

smooth. Elytra moderately long, at base narrower than pronotum at widest point, at suture shorter (ratio

0.80), at sides about as long as pronotum at midline; punctation and pubescence moderately dense, fine to

moderately coarse, surface between punctures without microsculpture. Wings fully developed. Abdomen

with tergite 7 bearing whitish apical seam of palisade setae (Figs. 382, 383); punctation and pubescence of

abdominal tergites finer than that of elytra and about equally dense, gradually becoming slightly sparser

toward apex; middle portion of tergite 3 (first visible) and occasionally also that of following tergite, to less

extent, impunctate. Front tarsus dilated in both sexes.

Revision of the Tribes Quediini and Atanygnathini

193

Male. First four segments of front tarsus slightly more dilated than in female (Fig. 381 ). Apical margin

of stemite 8 with moderately wide and not deep emargination (Fig. 6), triangular area before emargination

flattened and smooth. Aedoeagus (Figs. 7-9) elongate, median lobe parallel-sided in middle part, then

dilated toward apical part which is markedly narrowed anteriad, deeply emarginate apically and broadly

grooved at midline before emargination to accommodate apical part of paramere; both portions of apical part

of median lobe just laterad to median emargination distinctly hooked on surface adjacent to paramere.

Paramere variably exceeding apex of median lobe, very nanow and elongate, variably dilated, rounded,

obtusely truncate to feebly emarginate apically; four short apical setae and one pair of long lateral setae on

each side close to apex; underside of paramere without sensory peg setae, often variably swollen; internal

sac with typically shaped, markedly sclerotized structure with short and stout subacute tubercles.

Length 7. 1-1 2.0 mm.

Type material. — Quedius apiciconiis. Eppelsheim (1895b:391) described the

species from a single female from Sikkim. The holotype is deposited in the

Eppelsheim collection in the Naturhistorisches Museum in Wien, Austria. It is

labelled as follows: pink square label/ “Sikkim”/ “apicicornis Sikkim Epp.

Himalaya, leg. Waagen”/ “Je ne I’ai pas”/ “c. Epplsh. Stend d.”/ “Typus” (white

label)/ “Typus” (red label). It is very well preserved and intact (see also Smetana

1975:325).

Quedius cyanopterus. Scheerpeltz (1976b: 150) described the species from six

specimens from Nepal. They are deposited in the Zoologische Staatssammlung des

Bayerischen Staates, Miinchen, Eederal Republic of Germany, and are labelled as

follows: Spec. No. 1: ‘V ”/ “Nepal Kathmandu 10.-15. IV.62 1400 m leg. G. Ebert”/

“Quedius cyanopterus Scheerp.”/ “Holotypus”/ “Typus Quedius cyanopterus

Scheerpeltz”/ “Quedius cyanopterus nov. spec. det. Scheerpeltz. 1966”. Spec. no. 2:

“?”/ “Nepal Ting-Sang-La 3800 m 6/7. V. 62, Leg. G. Ebert”/ “Quedius cyanopterus

Scheerp.”/ “Allotypus”/ “Quedius cyanopterus O. Scheerpeltz”. Spec. No. 3: first

two labels as in allotype, plus label “Cotypus Quedius cyanopterus O. Scheerpeltz”.

Spec. No. 4 and 5: same three labels as in Spec. No. 3. Spec. No. 6: “$”/ “Nepal

Prov. nr. 1 East Pultschuk”/ “30. V. 1964 1700 m Loffler leg. (6)”/ “Staatsslg.

Miinchen”/ “Cotypus Quedius cyanopterus O. Scheerpeltz”.

All six specimens are conspecific; the allotype is in fact a male as is the

holotype. They cannot be distinguished from the holotype of Q. apicicornis; the

name Q. cyanopterus is a junior synonym of Q. apicicornis, as already suggested by

Coiffait (1982:32). All specimens were accordingly labelled.

Quedius sherpai. Coiffait (1982a:77) described the species from three specimens

from Nepal. I was able to study two specimens deposited in the Senckenberg

Museum, Erankfurt a.M., Eederal Republic of Germany. They are labelled as

follows: Spec. No. 1: “Trisuli-Tal Gosaikund, 3200 m 23.-26.IV. 1973”/

“Nepal-Expedition Jochen Martens”/ “TYPE”/ “Holo-typus” SMFC 15106

(underside)/ “Quedius (Microsaurus) sherpa H. Coiffait 1979”. Spec. No. 2

(female): first two labels as in holotype, plus “PARATYPE”/ “Para-typus SMFC

15107” (underside). Both specimens cannot be specifically distinguished from the

holotype of Q. apicicornis (see the discussion for all details); the name Q. sherpai is

a junior synonym of apicicornis. Both specimens were accordingly labelled.

Quaest. Ent., 1988, 24 (2)

194

Smetana

Qiiedius flavoterminatus. Cameron (1932:290) described the species from a

single specimen from the Darjeeling district. The female holotype in the British

Museum (Natural History), London, is labelled as follows: “Type” (round label with

red margin)/ “Rangirum, 6000' Darjeling, Bengal. J.C.M. Gardner 8.IX.1929”/

“500”/ “Quedius flavoterminatus Cam. TYPE”/ “M. Cameron. Bequest. B.M.

1955-147”.

The specimen is in poor general shape and the base of the abdomen is damaged.

It cannot be distinguished from small females of Q. apiciconiis. The name Q.

flavoterminatus is a Junior synonym of Q. apicicornis. My determination label

“Quedius apicicornis Epp. Smetana det. 1983” has been attached to it.

Geographical distribution. — Quedius apicicornis is known from Nepal, West

Bengal, Sikkim and Bhutan (Map 8).

Material studied. — 149 specimens.

BHUTAN. Dechhi Paka, 3300 m, 19-20.VI.72 (NHMB) 1.

INDIA. “India” (BMNH) 1. West Bengal. Ghum Distr., Tiger Hill, 8300-10000’, V.VI.1931, Cameron

(BMNH) 1. Sikkim. See Type material of “Q. apicicornis”.

NEPAL. Kathmandu Distr. Siwapuri, 24. III. 82, de Rougemont (GDRC) 3; Siwapuri Dara, 2300- 2500

m, 29. IV., 30. IV., l.V. and 3.V.83, A. Smetana (ASCC, BMNH, CNCC) 15. Kliandhari Distr. above

Sheduwa, 3000-3050 m, 3 1 .II1.-2.IV.82, A. & Z. Smetana (ASCC, CNCC) 9; “Bakan”, W of Tashigaon,

3200 m, 3.-5.1V.82, A. & Z. Smetana (ASCC, CNCC, BMNH) 1 1; above Tashigaon, 3100 m, 7.-8. .IV. 82, A.

& Z. Smetana (ASCC, CNCC) 7; Forest above Ahale, 2400 m, 25.III.82, A. & Z. Smetana (ASCC) 1; Forest

NW Ktiwapani, 2250-2550 m, 1 1. -15. IV. 82, A. & A. Smetana; 24.IV. 84, Smetana & Lbbl (ASCC, MHNG)

7; above Num, 1900-2200 m, 23.IV.84, Smetana & Lobl (ASCC) 1; Induwa Khola Valley, 2000-2600 m,

15.-18.IV.84, Smetana & Lbbl (ASCC, CNCC, MHNG) 9; Forest S Mansingma, 2200-2600 m,

1 1.-13.IV.84, Lbbl & Smetana (MHNG) 2. Lalitpiir Distr. Phulcoki, Franz (CNCC) 2; Phulcoki, 2400-2650

m, 20.-22.1V. 82, A. & Z. Smetana; 14.-15.X.83, Smetana & Lbbl; 28.-30.IV.84, Lbbl & Smetana (ASCC,

BMNH, CNCC, MHNG) 30. Manang Distr. Forest W Bagarchhap, 2200 m, 2 FIX. 83, Smetana & Lbbl

(ASCC, MHNG) 4; Marsyandi above Bagarchhap, 2400 m, 13.- 14. IV. 80, Martens & Ausobsky (SMFM) 4;

same 2550 m, 14. 17. IV. 80, Martens & Ausobsky (SMFM)2. Mustang Distr. S Lethe, 2450-2600 m,

30.IV.-1.V.80, Martens & Ausobsky (SMFM) 1; Thaksang, 3150-3400 m, 26.-29.IV. 80, Martens &

Ausobsky (SMFM) 1. Nuwakot Distr. Malemchi, 2800 m, 17.IV.81, Lbbl & Smetana (ASCC) 3;

Kutumsang, 2200-2700 m, 6-7.IV.81, Lbbl & Smetana (ASCC, CNCC) 3; Gul Bhanjang, 2600 m, 6.IV.81,

Lbbl & Smetana (MHNG) 2. Parhat Distr. Ghoropani Pass, N Slope, 2700-2800 m, 5-6. X. 83, Smetana &

Lbbl (ASCC, CNCC) 4; ridge E Ghoropani Pass, 3100 m, 7.X.83, Smetana & Lbbl (MHMG) 1; forest below

Fulung, Franz (CNCC) 1. Dhaulagiri Thankur, N Dhorpatan, 3350 m, 24. -28. V. 73, J. Martens (SMFM) 1; S

Dhaulagiri Dhorpatan, 3000-3200 m, 18.-21.V.73, J. Martens (SMFM) 1; Gosaikund, Trisuli Valley, 3200

m, 23.-26.IV. 73, J. Martens (SMFM) 1; Chordung - Jin, 2900 m, III.73, J. Martens (SMFM) 1; Ghar Khola

below Deorali Pass, 2000 m, 25. III. 78 (MHNG) 2; Pha Kumpe, 2300 m, IV. 84, Morvan (GMRC) 5; Dobate

Ridge NE Barahbise, 2800 m, 3.V.81, Lbbl & Smetana (ASCC) 2.

Bionomics.-— In Nepal Quedius apicicornis was typically collected in

association with dead fallen trees. Most specimens were taken from under loose

bark or in moss on fallen trees. Specimens from Malemchi were taken from moss

and debris soaked with fermenting sap around pieces of a large tree felled during a

storm. Occasionally, specimens can also be found under rocks or in the forest floor

debris.

Comparisons and variations. — Quedius apicicornis varies considerably in most

external characters, particularly in the colouration of the antennae and of the tip of

the abdomen, in the shape of the head and pronotum, and also in the coarseness of

Revision of the Tribes Quediini and Atanygnathini

195

Quaest. Ent., 1988, 24 (2)

Map 8. Distribution records for: Quedius apicicornis.

196

Smetana

the punctation of the elytra. The apical portion of median lobe of aedoeagus is rather

constant in shape (Fig. 3); however, the apical portion of the paramere varies

considerably (see the description, Fig. 5 and Figs. 2 and 3 in Smetana 1975:324) and

so also does the distance by which it exceeds the apex of median lobe.

The aedoagus of Q. sherpai is supposed to differ from that of Q. apicicornis by

the absence of the hooks on the apical portion of median lobe (see the description).

However, the hooks are present on the median lobe of the holotype of Q. sherpai

and can be clearly seen especially in lateral view.

3. Quedius (Microsaurus) heesoni Cameron

Figs. 10-13; Map 9

Quedius heesoni Cameron 1932:285

Quedius mimeticus Cameron 1932:286 (syn.nov.)

Quedius notahilis Cameron 1932:286 {syn.nov.)

Quedius perajfinis Cameron 1932:286 (syn.nov.)

Description.— In all external characters very similar to Q. apicicornis and displaying similar

variability in colouration of antennae and tip of abdomen as described for Q. apicicornis. However, it differs

as follows: elytra black, without bluish metallic lustre, abdominal tergite 7 entirely dark, even in specimens

with apex of abdomen yellowish. Posterior frontal puncture situated closer to posteromedian margin of eye

and separated from it by distance not or only slightly larger than diameter of puncture. Antenna in general

stouter. Pronotum with lateral portions in general slightly more explanate, especially posteriorly; sublateral

rows of punctures with two to four (usually three) punctures, posterior puncture situated behind level of

large lateral puncture. Scutellum impunctate; however, triangular area at base transversely rugose.

Punctation of elytra in general coarser and deeper.

Male. First four segments of front tarsus slightly more dilated than in female. Apical margin of stemite 8

(Fig. 10) with emargination similar to that of Q. apicicornis (Fig. 6). Aedoeagus (Figs. 1 1-13) similar to that

of Q. apicicornis-, however, median lobe no more than slightly emarginate apically, apical portion shallowly

and broadly impressed medially to accomodate apical part of paramere and not hooked on surface adjacent

to paramere. Paramere not or Just about reaching apex of median lobe, usually slightly emarginate apically,

without sensory peg setae; four apical setae considerably longer than those of Q. apicicornis-, internal sac

different, without markedly sclerotized structure with short stout subacute tubercles.

Length 8.0-12.0 mm.

Type material. — Quedius heesoni. Ten syntypes in the British Museum (Natural

History), London, are labelled as follows: Spec. No. 1 (d): “Type” (round label with

red margin/ “Dung”/ “Deoban, 9331. Chakrata, U.P.”/ “Cameron 3.5.21”/ “Quedius

beesoni TYPE Cam.”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE”

(round label with blue margin). Spec. No. 2 ($): “Deoban, 9331. Chakrata, U.P.”/

“Dr. Cameron 3.5.21”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE”

(round label with blue margin). Spec. No. 3 (d): “Chakrata Dist. Sainj Khud 6500'”/

“Dr. Cameron 27.V.22”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE”

(round label with blue margin). Spec No. 4 ($): “Chakrata Dist. Chulli Khud 8000'”/

“Dr. Cameron 14.V.22”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE”

(round label with blue margin). Spec. No. 5 (c?): “Chakrata Dist. Patra Khud 8000'”/

“Dr. Cameron 15.V.22”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE”

(round label with blue margin). Spec. Nos. 6-8 (29?, Ic3): “Chakrata Dist. Kanasar

7050'”/ “Dr. Cameron 14-22. V. 22”/ “M. Cameron. Bequest. B.M. 1955-147.”/

Revision of the Tribes Quediini and Atanygnathini

197

“SYN-TYPE” (round label with blue margin). Spec. No. 9 (?): “Ghum district

V-VI-31 Dr. Cameron”/ “M. Cameron. Bequest. 1955-147.”/ “SYN-TYPE” (round

label with blue margin). Spec No. 10 (9): “Ghum dist. Rongdong Valley V-VI-31

Dr. Cameron”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE” (round

label with blue margin). All specimens are conspecific. All males were dissected.

The first male, bearing the label “TYPE” is hereby designated as the lectyotype of

Q. heesoni. The aedoeagus, genital segment and stemite 8 of this specimen were

glued to plate with beetle. The label “Lectotype Q. beesoni Cam. Smetana des.

1983” was attached to it.

Quedius mirneticus. Four syntypes in the British Museum (Natural History),

London, are labelled as follows: Spec. No. 1(9): “TYPE” (round label with red

margin)/ “Ghum Dist. Rongdong Valley V-VI-31 Dr. Cameron”/ “Q. mirneticus

Cam. TYPE”/ “M. Cameron. Bequest B.M. 1955-147.”/ “SYN-TYPE” (round label

with blue margin). Spec. Nos. 2-4 (299, Id): “Ghum dist. Rongdong Valley V-VI-31

Dr. Cameron”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE” (round

label with blue margin). All specimens are conspecific. The male specimen was

dissected and the aedoeagus, genital segment and sternite 8 were glued to plate with

beetle. The specimen is hereby designated as the lectotype of Q. mirneticus', the label

“Lectotype Q. mirneticus Cam. Smetana des. 1983” was attached to it.

The specimens of this original series cannot be specifically distinguished from

the lectotype of Q. heesoni. The name Q. mirneticus is a synonym of Q. heesoni.

Quedius notahilis. Five syntypes in the British Museum (Natural History),

London, are labelled as follows: Spec. No. 1 (d): “TYPE” (round label with red

margin)/ “Ghum distr. Rongdong Valley V-VI-31 Dr. Cameron”/ “Q. notabilis Cam.

TYPE”/ “Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE” (round label with

blue margin). Spec. Nos. 2-4 (299, Id): “Ghum dist. Rongdong Valley V-VI-31 Dr.

Cameron”/ “M. Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE” (round label

with blue margin). Spec. No. 5: (d): “Ghum dist. Mangpo V-31 Dr. Cameron”/ “M.

Cameron. Bequest. B.M. 1955-147.”/ “SYN-TYPE” (round label with blue margin).

Both males were dissected. The first male, bearing the label “TYPE” is hereby

designated as the lectotype of Q. notahilis. The aedoeagus, genital segment and

sternite 8 of this specimen were glued to plate with beetle. The label “Lectotype Q.

notabilis Cam. Smetana des. 1983” was attached to it.

The specimens of this original series cannot be specifically distinguished from

the lectotype of Q. heesoni. The name Q. notahilis is a synonym of Q. heesoni.

Quedius perajfinis. Two syntypes in the British Museum (Natural History),

London, are labelled as follows: Spec. No. 1 (d):'^ “Type” (round label with red

margin)/ “Ghum dist. Rongdong Valley V-VI-31 Dr. Cameron”/ “Q. peraffinis Cam.

TYPE”/ “M. Cameron. Bequest. B.M. 1955-147.”. Spec. No. 2: “Ghum dist,

Rongdong Valley V-VI-31 Dr. Cameron”/ “M. Cameron. Bequest. B.M.

4

It

is a male, although Cameron (1932:286) stated “c? unknown”.

Quaest. Ent., 1988,24 (2)

198

Smetana

1955-147.”/ “peraffinis Cam. syntype”/ “SYN-TYPE” (round label with blue

margin).

The first specimen (male) was dissected and the aedoeagus, genital segment and

stemite 8 were glued to plate with beetle. The tip of the abdomen, as well as both

front legs are missing in this specimen. The specimen is hereby designated as the

lectotype of Q. peraffinis', the label “Lectotype Q. peraffinis Cameron Smetana des.

1983” was attached to it.

Both specimens cannot be specifically distinguished from the lectotype of Q.

heesoni. The name Q. peraffinis is a synonym of Q. heesoni.

Geographical distribution. — Quedius heesoni is distributed from northern Uttar

Pradesh eastward through Nepal to the Darjeeling area (Map 9).

Material studied. — 74 specimens.

INDIA. Uttar Pradesh. Kumaon: Pindar Valley, 8000-1 1000', Champion (BMNH) 3; Sukhatal, 8000',

V.1920, Champion (BMNH) 2; Chakrata Distr.: Deoban, 9331', 3.V.21, Cameron (BMNH) 2; Kanasar,

7030', 14-22.V.22, Cameron (ASCC, BMNH) 5; Chutli Khud, 8000', 17.V.22, Cameron (ASCC, BMNH) 3;

Binal Gad, 7000-8000', 24.V.22, Cameron (BMNH) 1. West Bengal. Darjeeling Distr.: Tiger Hill,

8500-10000', V-VI-1931, Cameron (BMNH) 1; Rongdong Valley, V-VI-1931, Cameron (BMNH, CNCC)

7.

NEPAL. Kathmandu Distr. Siwapuri Dara, 2400-2450 m, 30.1V and 2.V.85, A Smetana (ASCC) 2.

Lalitpur Distr. Phulcoki, 2600-2750 m, 13-15.X.83, Smetana & Lobl (ASCC, MHNB) 8; same, 2600 m,

29. IV. 84, Smetana & Lobl (ASCC) 1. Manang Distr. Lorest W Bagarchhap, 2200 m, 21-24. IX. 83, Smetana

& Lobl (ASCC, CNCC, MHNG) 7. Parhat Distr. Ghoropani Pass N slope, 2700 m, 6.X83, Smetana & Lobl

(ASCC, MHNB) 5; Ghoropani vie., 2700-3100 m, 5.-9.X.83, Smetana & Lobl (MHNG) 2. Sindhupalchok

Distr. Malemchi, 2800 m, 17-18.IV.81, Lobl & Smetana (ASCC) 3; Dobate Ridge NE Barahbise, 2800 m,

2.V.8 1 , Lobl & Smetana (ASCC) 1 .

Bionomics. — Little is known about the habitat preferences of this species.

Cameron (1932:286) mentions that it occurs in Polyporus', one specimen was

collected in “dung” (see Type material of Q. heesoni). All specimens from Nepal

were taken by sifting a wide variety of mushrooms and moss and debris around

them on large fallen broadleaved trees.

Comparisons and variations. — Quedius heesoni can easily be distinguished

from Q. apicicornis by the characters mentioned in the description. The variability

in the colouration of the apical portion of the antennae and of the apex of abdomen

was the reason why Cameron (1932) described the species under four different

specific names, although he suspected that at least some of them might be only

colour varieties of one species. The name Q. heesoni was used for specimens with

both antennae and abdomen uniformly dark, Q. mimeticus for specimens with the

antennae uniformly dark and the apex of abdomen yellow, Q. notabilis for

specimens with antennal segments 8-10 and apex of abdomen yellow, and Q.

peraffinis for specimens with antennal segments 8-10 yellow and the abdomen

uniformly dark. There are some other slight differences between the specimens

belonging to the different colour varieties {e.g., those of Q. heesoni tend to be larger

and more robust than those of Q. mimeticus or Q. peraffinis and those of Q. notabilis

tend to have longer outer segments of the antenna). On the other hand, the males of

all colour varieties have identical aedoeagi; there is, therefore, hardly any doubt that

Revision of the Tribes Quediini and Atanygnathini

199

Quaest. Ent., 1988, 24 (2)

Map 9. Distribution records for: Quedius heesoni.

200

Smetana

they all represent differently coloured populations of the same variable species.

4. Quedius (Microsaurus) flavocaudatus Cameron

Figs. 14-16; Map 10

Quedius flavocaudatus Cameron 1 944: 1 3

Description. — Black, head and pronotum slightly, abdomen markedly iridescent, elytra red,

abdominal tergite 7 (fifth visible), corresponding sternite and rest of abdomen yellowish-red; labial palpi

testaceous, maxillary palpi piceous, becoming paler toward apex; antennae black, last four segments pale

yellowish; legs piceous-black to black with paler tarsi. Head of rounded quadrangular shape, wider than long

(ratio 1.36), markedly nan'owed posteriorly behind eyes, posterior angles rounded and indistinct; eyes

convex but only slightly protruding from lateral contours of head, tempora about as long as length of eyes

seen from above; no additional punctures between anterior frontal punctures; posterior frontal puncture

situated very close to posteromedian margin of eye and separated from it by distance about equal to diameter

of puncture, two (rarely three unilaterally) additional setiferous punctures between it and posterior margin of

head; temporal puncture situated closer to posterior margin of eye than to posterior margin of head; surface

of head with extremely fine and dense microsculpture consisting of transverse waves becoming gradually

irregular to almost meshed anteriorly on clypeus and anteromediad of each eye. Last segment of maxillary

palpus distinctly longer than penultimate segment, slightly attenuate basally. Antenna moderately long and

only slightly thickened toward apex; segment 3 distinctly longer than segment 2, segments 4 and 5 slightly

longer than wide, following segments about as long as wide, gradually becoming slightly shorter, last

segment shorter than two preceding segments combined. Pronotum wider than long (ratio 1.23), broadly

rounded basally and arcuately narrowed anteriorly, with lateral portions slightly explanate, disc moderately

transversely convex; dorsal rows each with four rather coarse punctures; sublateral rows each with three

(occasionally four unilaterally) punctures, posterior puncture situated behind level of large lateral puncture;

microsculpture of pronotum similar to that of head but still finer and denser. Scutellum large, impunctate,

but with small triangular area at base transversely rugose. Elytra moderately long, at base distinctly narrower

than pronotum at widest point, at suture feebly shorter (ratio 0.89), at sides about as long as pronotum at

midline; punctation and pubescence moderately dense and coarse, pubescence black, surface between

punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 bearing whitish apical

seam of palisade setae; punctation and pubescence of abdominal tergites distinctly finer and slightly denser

than that of elytra, not becoming appreciably sparser toward apex; pubescence black except rusty on pale

apical segments. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus slightly more dilated than in female. Apical margin of sternite 8

with moderately deep and wide, arcuate emargination (Fig. 14), triangular area before emargination flattened

and smooth. Aedoeagus (Figs. 15-17) rather small, narrow and elongate; median lobe slightly widened and

then suddenly narrowed into wedge-shaped, narrow apical portion with obtuse apex; subapical portion of

median lobe on opposite side to paramere markedly sclerotized and widely, arcuately emarginate at base of

wedge-shaped apical portion. Paramere elongate, parallel-sided in middle portion and then slightly dilated

apically; apical portion not quite reaching apex of median lobe, with deep V-shaped emargination; with two

moderately long apical setae at each side of apical emargination and one pair of lateral setae on each side

close to apex; underside of paramere with several small sensory peg setae at each side of apical

emargination; internal sac simple, without conspicuous sclerites.

Length 12.0-13.3 mm.

Type material. — Cameron (1944:13) described the species from a single male

specimen from Naini Tal, Uttar Pradesh. The holotype in the British Museum

(Natural History), London, is labelled as follows: “Type” (round label with red

margin)/ “6000-8000 Naini Tal U.P. Balwant Singh 4.V.1935”/ “In soil”/ “Q.

flavocaudatus Cam. TYPE”/ “M. Cameron. Bequest. B.M. 1955-147”. The

specimen was dissected and the aedoeagus and strenite 8 were mounted in Canada

Balsam.

Revision of the Tribes Quediini and Atanygnathini

201

Geographical distribution. — Quedius flavocaudatiis is distributed from northern

Uttar Pradesh eastward to central Nepal (Map 10).

Material studied. — 5 specimens.

INDIA. “India” (BMNH) I. Uttar Pradesh. See Type material.

NEPAL. Lalitpur Distr. Phulcoki, 2600 m, 20. IV. 82, A. & Z. Smetana (ASCC) 1. Nuwakot Distr.

Malemchi, 2900 m, 18.IV.81, Ldbl & Smetana (ASCC) 1. N. Dhorpatan, Parkapan, 1977, P. Morvan

(GDRC) 1.

Bionomics. — The specimen from near Malemchi was sifted from thick moss

from lower portions of the trunk of an old oak tree; the specimen from Phulcoki was

taken by sifting old mushrooms, moss and bark around them found on an old fallen

oak tree.

Comparisons. — Quedius flavocaudatus can easily be recognized by the large

size and the colouration, in combination with the presence of four punctures in each

dorsal row on the pronotum.

5. Quedius (Microsaurus) antennalis Cameron

Figs. 17-20; Map 10

Quedius antennalis Cameron 1932:285

Description . — Piceous-black, elytra, pronotum and head dark reddish-brown piceous, abdomen

slightly iridescent, posterior half of tergite 7 (fifth visible) and apex of corresponding stemite and apex of

abdomen reddish-yellow; labial and maxillary palpi rufo-testaceous; antennae with three basal segments

rufo-testaceous, following three segments slightly paler, however, very vaguely infuscate in middle portion,

last five segments very pale, whitish-yellow; legs piceous-black with paler tarsi. Head of rounded

quadrangular shape, wider than long (ratio 1.34), markedly narrowed posteriorly behind eyes, posterior

angles quite indistinct; eyes convex and slightly protruding from lateral contours of head, tempora slightly

shorter than length of eyes seen from above (ratio 0.77); no additional punctures between anterior frontal

punctures; posterior frontal puncture situated near posteromedian margin of eye and separated from it by

distance distinctly larger than diameter of puncture, two additional setiferous punctures between it and

posterior margin of head; temporal puncture situated closer to posterior margin of eye than to posterior

margin of head; surface of head with extremely fine and dense microsculpture of transverse waves, including

clypeus. Last segment of maxillary palpus distinctly longer than penultimate segment (ratio 1.5), slightly

attenuate basally. Antenna long and only slightly thickened toward apex; segment 3 distinctly longer than

segment 2, segments 4-6 elongate, much longer than wide, following segments gradually becoming shorter

and wider; however, even segments 9 and 10 appreciably longer than wide, last segment slightly shorter than

two preceding segments combined. Pronotum wider than long (ratio 1.25), broadly rounded basally and

continuously, slightly arcuately narrowed anteriorly, with lateral portions slightly explanate, disc rather

markedly transversely convex; dorsal rows each with three punctures; sublateral rows each with two

punctures, posterior puncture at about level of large lateral puncture; large lateral puncture situated not far

from lateral margin, separated from it by distance equal to about width of apex of second antennal segment;

surface of pronotum with microsculpture similar to that on head, but still finer and hardly visible. Scutellum

large, impunctate, but with extensive triangular area transversely rugose. Elytra moderately long, at ba.se

distinctly narrower than pronotum at widest point, at suture as long as, at sides feebly longer (ratio 1.11) than

pronotum at midline; punctation moderately coarse, slightly asperate and dense, pubescence black, surface

between punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 bearing whitish

apical seam of palisade setae; punctation of abdominal tergites fine, considerably finer than that on elytra,

evenly covering surface of tergites (including middle portion of first visible tergite), pubescence black, paler

on pale tip of abdomen.

Male. Eirst four .segments of front tarsus markedly dilated. Apical margin of stemite 8 with moderately

wide and rather shallow, arcuate emargination (Eig. 18), triangular area beofre emargination flattened and

smooth. Aedoeagus (Eigs. 17, 19, 20) of characteristic shape, large and elongate; median lobe slightly

Quaest. Ent., 1988, 24 (2)

202

Smetana

dilated and then slightly, evenly narrowed toward subtruncate and subemarginate apical margin, apical

portion of median lobe on side opposite to paramere markedly excavate, gutter-like. Paramere short,

parallel-sided with apical margin subtruncate and subemarginate medially, its apical margin separated from

apex of median lobe by distance equal to about median length of paramere; with eight unequally long setae

at apical margin; no sensory peg setae on underside of paramere.

Female. Unknown.

Length 1 1 .0 mm.

Type material. — Cameron (1932:285) described the species from a single male

specimen from the Darjeeling area. The holotype in the British Museum (Natural

History), London, is labelled as follows: “Type” (round label with red margin/

“Gopaldhara, B.W. Darjiling, 4720 ft. 23. X. -193 H. Stevens.”/ “H. Stevens. Brit.

Mus. 1922-397.”/ “Quedius antennalis Cam. TYPE”. The specimen was dissected

and the aedoeagus and sternite 8 were mounted in Canada Balsam.

Geographical distrihution. — Quedius antennalis is at present known only from

the type locality in the Darjeeling area (Map 10).

Material studied. — The holotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Quedius antennalis is very similar to Q. hirmanus and the latter

species actually may not be specifically different. The discussion following the

description of Q. hirmanus should be consulted for more details.

Quedius antennalis also resembles Q. heesoni; however, it differs, in addition to

the differently shaped aedoeagus, by the differently developed sublateral rows on

pronotum (see the key and respective descriptions), and by the absence, in Q.

antennalis, of the smooth area in the middle of the first visible abdominal tergite.

6. Quedius (Microsaurus) hirmanus Cameron

Map 10

Quedius hirmanus Cameron 1932;284

Description. — in all characters quite similar to Q. antennalis but different as follows; Form in

general larger and more robust. Black, head and pronotum feebly, abdomen slightly iridescent, posterior half

of abdominal tergite 7 (fifth visible), corresponding sternite and apex of abdomen reddish-yellow; labial

palpi testaceous, maxillary palpi piceous, becoming paler toward apex; antennae black, last four segments

yellowish; legs piceous with paler tarsi. Outer antennal segments slightly shorter. Left dorsal row on

pronotum with three, right row with only two punctures (first puncture missing); left sublateral row with

only one (front) puncture, right sublateral row with two punctures, posterior puncture at about level of large

lateral puncture; large lateral puncture situated far from lateral margin, separated from it by distance almost

twice as large as width of apex of second antennal segment. Elytra wider with punctation deeper and coarser.

Male. Unknown.

Length 12.0 mm.

Type material. — Cameron (1932:284) described the species from a single

female from Burma. The holotype in the British Museum (Natural History), London,

is labelled as follows: “64517”/ “Type” (round label with red margin)/ “Doherty”/

“Birmah Ruby Mes”/ “Quedius birmanus Cam. TYPE”.

Geographical distrihution. — Quedius hirmanus is at present known only from

the type locality in Burma (Map 10).

Revision of the Tribes Quediini and Atanygnathini

203

Material studied. — The holotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — It is possible that Q. birmanus does not actually specifically

differ from Q. antennalis. The differences in general form and colouration are not

significant (the holotype of Q. antennalis may be a slightly discoloured specimen). I

consider Q. birmanus as a separate species mainly because of the differently situated

large lateral puncture on the pronotum and the wider and more coarsely punctate

elytra. The status of Q. birmanus cannot be definitely assessed until more specimens

of both species, and particularly the males of Q. birmanus are available for study.

The first puncture in the right dorsal row on the pronotum is missing in the

holotype; however, there is an additional round impression (not setiferous) behind

the last puncture of the right row, imitating an additional puncture.

7. Quedius (Microsaurus) martensi Smetana

Figs. 21-23; Map 7

Quedius martensi Smetana 1975:329

Description. — Piceous-black, apical margins of abdominal tergites and apex of abdomen

indistinctly paler. Palpi rufo-brunneous, antennae brownish-piceous with three basal segments darker, legs

piceous-black, inner surface of front tibiae and all tarsi rufo-brunneous. Head of rounded quadrangular

shape, as long as wide, almost parallel-sided behind eyes, posterior angles obtuse, indistinct; eyes very small

and rather flat, tempora considerably longer than length of eyes seen from above (ratio 2.03); no additional

setiferous punctures between anterior frontal punctures; posterior frontal puncture situated slightly closer to

posteromedian margin of eye than to posterior margin of head; two additional setiferous punctures between

it and posterior margin of head; temporal puncture situated about half as far from posterior margin of head as

from posterior margin of eye; deflexed portion of tempora with some fine punctures posteriorly; surface of

head with fine and very dense microsculpture of irregular transverse lines with scattered longitudinal

connections. Antenna rather short, segment 3 slightly longer than segment 2, segments 4 and 5 slightly

longer than wide, following segments about as long as wide, becoming gradually wider and shorter, outer

segments slightly transverse, last segment distinctly shorter than two preceding segments combined.

Pronotum wider than long (ratio 1.14), basal margin continuously rounded with lateral margins, latter

distinctly narrowed anteriorly; lateral portions of pronotum explanate posteriorly; dorsal rows each with two

punctures (first puncture usually present at apical margin of pronotum seems to be missing); sublateral rows

each with two punctures, posterior puncture situated before level of large lateral puncture; surface of

pronotum with microsculpture identical to that of head, except perhaps feebly denser. Scutellum impunctate,

smooth. Elytra moderately long, at base narrower than pronotum at widest point, at suture about equally

long, at sides slightly longer (ratio 1.12) than pronotum at midline; punctation and pubescence superficial,

rather fine and sparse; interspaces between punctures much larger than diameters of punctures; surface

between punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible)

bearing whitish apical seam of palisade setae; punctation and pubescence of abdominal tergites distinctly

denser than that of elytra, small middle portion of first visible tergite impunctate.

Male. First four segments of front tarsus markedly dilated. Apical margin of sternite 8 with moderately

wide, shallow subangular emargination (Fig. 21), small triangular area before emargination flattened and

smooth. Aedoeagus (Figs. 22, 23) small, apical part of median lobe, when paramere removed, with two

small subapical hooks. Paramere just about reaching apex of median lobe, markedly laterally compressed in

middle part, dilated toward apex; with four rather short setae at apical margin and two slightly shorter setae

at each lateral margin below apex; underside of paramere markedly excavate, with only two sensory peg

setae at each lateral margin close to apex.

Length 10.0 mm (abdomen slightly extended).

Qiiaest. Ent., 1988, 24 (2)

204

Smetana

Type material. — The male holotype, deposited in the Franz collection, Modling,

Austria, is labelled as follows: “Khumbu, Zusammenfluss Imja, Phunki Drangka

3150-3300 m, 1.10.70, J. Martens”/ “HOLOTYPE Quedius martens! A. Smetana

1975”.

Geographical distribution. — Quedius martensi is known only from the type

locality in Khumbu Himalaya in Nepal (Map 7).

Material studied. — The holotype.

Bionomics. — No details are known about the habitat requirements of this

species. It was most probably taken in a subalpine birch forest (see Franz 1971:443).

The small eyes suggest some secretive way of life, possibly in nests of some

mammal.

Comparisons and variations. — Quedius martensi is well characterized, in

addition to the characters on the aedoeagus, by the colouration, the small eyes, the

chaetotaxy of the head and pronotum, and by the sparse punctation of the elytra. For

a comparison with Q. dui see the discussion there.

In the original description of this species, I described the surface of the elytra

between the punctures as “covered with extremely fine and rather dense

microsculpture consisting of intricate curved lines and irregularities”. However,

after closer examination in diffuse light, using a translucent filter, and after thorough

cleaning of the surface, the surface appears to be without any microsculpture.

Quedius martensi may have normally three punctures in each of the dorsal rows

on the pronotum. The usual first puncture, situated close to apical margin, may be

missing bilaterally in the holotype.

8. Quedius (Microsaurus) dui spec. nor.

Figs. 24-27; Map 7

Description. — Form rather stout; piceous, head black, elytra dark reddish-brown, apical margins of

abdominal tergites and apex of abdomen indistinctly paler. Palpi dark brown, antennae piceous, legs

piceous-brown with inner portions of middle and hind tibiae darkened, tarsi slightly paler, particularly their

last segment. Head of obtusely quadrangular shape, slightly transverse (ratio length: width=0.85),

parallel-sided behind eyes, posterior angles broadly arcuate; eyes small and flat, not protruding from lateral

contours of head, tempora twice as long as length of eyes seen from above; no additional setiferous

punctures between anterior fontal punctures; posterior frontal puncture and temporal puncture both situated

much closer to posterior margin of head than to posterior margin of eye; two small setiferous punctures

between posterior frontal puncture and posterior margin of head; tempora with some very fine punctures

posteriorly; surface of head with dense and rather coarse microsculpture of mostly transverse waves

becoming slightly confused on clypeus. Antenna short and rather stout, segment 3 distinctly longer than

segment 2, segments 4 and 5 as long as wide, segments 6-10 wider than long, gradually becoming shorter

and wider, segment 10 distinctly transverse, segment 1 1 about as long as two preceding segments combined.

Pronotum slightly wider than long (ratio 1.09), basal margin continuously rounded with lateral margins,

lateral margins distinctly, slightly arcuately narrowed anteriorly; dorsal rows each with two punctures;

sublateral rows each with two punctures, posterior puncture situated slightly before level of large lateral

puncture; surface of pronotum with microsculpture similar to that of head, however, waves of

microsculpture directed obliquely anteriad and becoming distinctly finer toward lateral margins. Scutellum

impunctate, surface with dense and very fine microsculpture of irregular transverse lines. Elytra at base

narrower than pronotum at widest point, hardly widened posteriorly, at suture as long as, at sides slightly

Revision of the Tribes Quediini and Atanygnathini

205

longer (ratio 1.12) than pronotum at midline; punctation fine and dense, interspaces between punctures no

more than slightly larger than diameters of punctures; pubescence dense, brownish-piceous; surface between

punctures without microsculpture. V/ings fully developed. Abdomen with tergite 7 (fifth visible) bearing

whitish apical seam of palisade setae; punctation and pubescence finer and denser than that of elytra, evenly

covering tergites, gradually becoming somewhat sparser toward apex of abdomen.

Male. First four segments of front tarsus markedly dilated. Stemite 8 with two strong setae on each side,

apical margin with wide and fairly deep, almost arcuate emargination (Fig. 24), small triangular area before

emargination flattened and smooth. Aedoeagus (Figs. 25-27) moderately large, median lobe slightly dilated

and then rather suddenly narrowed into subacute apex. Paramere not quite reaching apex of median lobe,

markedly constricted in middle portion, more apically slightly dilated and then narrowed toward truncate and

slightly emarginate apex; four long and about equally long setae at apical margin and one shorter seta at each

lateral margin below apex; sensory peg setae on underside of paramere forming isolated groups: one central

near apex of paramere and one irregular group near each lateral margin way below apex of paramere.

Length 8.7 mm.

Type material. — Holotype (male): “Punjab: Kulu Div. VI. 1932. H.G.

Champion”/ “H.G. Champion Coll. B.M. 1953-156.”/ “Quedius (Microsaurus)

spec. now Smetana det. 1984”. In the collection of the British Museum (Natural

History), London.

Geographical distribution. — Quedius did is at present known only from the type

locality in Himachal Pradesh (Map 7).

Bionomics. — Nothing is known about the habitat requirements of this species;

however, judging from its general habitus and the small and flat eyes, it may live in

burrows of some animal.

Comparisosn. — Quedius did is quite distinctive among the Himalayan species

due to the following combination of characters: body shape and colouration (see the

description), eyes small and flat, half as long as tempora, posterior frontal puncture

situated much closer to posterior margin of head than to posterior margin of eye;

dorsal rows on pronotum each with only two punctures, scutellum without rugae on

basal portion. It cannot be confused with any other Himalayan species.

Quedius martensi also has small eyes; however, it differs by the position of the

posterior frontal puncture (slightly closer to posteromedian margin of the eye than to

posterior margin of the head) and of the posterior puncture of the sublateral rows on

the pronotum (situated slightly before level of the large lateral puncture) and by the

sparse punctation of the elytra.

Etymology. — The specific epithet is the Nepali numeral dui (=two). It refers to

the number of punctures in each dorsal row on the pronotum.

9. Quedius (Microsaurus) adjacens Cameron

Figs. 28-31; Map 7

Quedius adjacens Cameron 1926:368; 1932:287

Description. — Piceous-black to black; pronotum feebly, abdomen distinctly iridescent. Palpi

piceous, antennae black, legs piceous-black with femora sometimes paler, rather piceous. ttead of rounded

quadrangular shape, slightly wider than long (ratio 1.18), with truncate apical margin of clypeus and almost

parallel-sided lateral margins behind eyes (holotype); or distinctly wider than long (ratio 1.35), with broadly,

arcuately emarginate apical margin of clypeus (emargination exposing articulation membrane between

clypeus and labrum) and lateral margins behind eyes noticeably narrowed posteriorly (two additional

Quaest. Ent., 1988, 24 (2)

206

Smetana

specimens from Cameron’s collection); eyes relatively large, moderately convex; tempora about as long as

to slightly shorter than length of eyes seen from above; no additional setiferous punctures between anterior

frontal punctures; posterior frontal puncture situated distinctly closer to posteromedian margin of eye than to

posterior margin of head, one additional setiferous puncture between it and posteromedian margin of eye and

two additional setiferous punctures between it and posterior margin of head (situated along posterior margin

of head); temporal puncture situated about half as far from posterior margin of head as from posterior margin

of eye; deflexed portion of tempora with a few fine punctures; surface of head with very fine and dense

microsculpture of transverse waves becoming irregular and forming some meshes on clypeus. Antenna

moderately long, segment 3 slightly longer than segment 2, segments 4-5 somewhat longer than wide,

following segments gradually becoming shorter, outer segments about as long as wide, last segments shorter

than the preceding segments combined. Pronotum hardly wider than long (ratio 1.07), basal margin

continuously rounded with lateral margins and tending to be feebly bisinuate, lateral margins distinctly,

arcuately narrowed anteriorly, posterolateral portions of pronotum slightly explanate; dorsal rows each with

three punctures; sublateral rows each with three or four punctures (with only two punctures unilaterally in

one specimen), last puncture situated behind level of large lateral puncture; surface of pronotum with

microsculpture similar to that on head. Scutellum impunctate, surface with dense and very fine

microsculpture of transverse lines. Elytra moderately long, at base feebly narrower than pronotum at widest

point, at suture equally long, at sides slightly longer (ratio 1.19) than pronotum at midline; punctation

superficial, pubescence rather fine and not dense, interspaces between punctures distinctly larger than

diameters of punctures; surface between punctures without microsculpture. Wings fully developed.

Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; punctation and

pubescence in general similar to that of elytra, except bases of tergites somewhat more densely punctate and

pubescent.

Male. First four segments of front tarsus markedly dilated. Apical margin of sternite 8 with moderately

wide and not deep, obtusely arcuate emargination (Fig. 28), small triangular area before emargination

flattened and smooth. Aedoeagus (Figs. 29-31) small, elongate and slightly asymmetrical; median lobe with

rounded apex and with one median and two lateral subapical denticles on face adjacent to paramere.

Paramere narrow, slightly asymmetrical, not reaching apex of median lobe; with four setae at apical margin

(median pair longer than lateral setae) and two similar setae at each lateral margin below apex; underside of

paramere flat, with two short and often irregular rows of three to four peg setae.

Fength 9.5-1 1.2 mm.

Type material. — There are three specimens in the British Museum (Natural

History), London, under the name Q. adjacens\ however, only the first specimen

(male) can be considered as belonging to the original series. It is labelled as follows:

“Type” (round label with red margin)/ “Deaban, 9331. Chakrata, U.P. Dr. Cameron.

3.5.21.”/ “Dung”/ “TYPE Quedius adjacens Dr. Cameron“/ “M. Cameron. Bequest.

B.M. 1955-147.”. The specimen was dissected and sternite 8 and the aedoeagus

were mounted in Canada Balsam; it is hereby designated as the lectoptype of Q.

adjacens; the label “LECTOTYPE. Quedius adjacens Cameron A. Smetana des.

1984” was attached to it.

Geographical distribution. — Quedius adjacens is distributed in the western

portion of the Himalaya; it is at present known from Kashmir, Himachal Pradesh

and Uttar Pradesh (Map 7).

Material studied. — 6 specimens.

INDIA. Kashmir. Gulmarg,VI-VII. 193 1, Cameron (BMNH) 1; Yusmarg, VI. 1981, de Rougemont

(GRDC) 1. Himachal Pradesh. Simla Hills, Narkanda, 9230’, 15.IX.21, Cameron (BMNH) 1. Uttar Pradesh.

See Type material.

Bionomics. — No details are known about the habitat preference of this species.

The lectotype was collected under dry dung.

Revision of the Tribes Quediini and Atanygnathini

207

Comparisons. — The difference in the configuration of the head, particularly of

the apical margin of the clypeus (see the description) between the lectotype and the

two additional specimens from the Cameron collection I have seen, is remarkable.

However, in all other characters, including the shape of the median lobe and the

paramere of the aedoeagus, the three specimens are identical and there is no doubt in

my mind that they are conspecific.

10. Quedius (Microsaurus) ochripennis (Menetries)

Figs. 32-34; Map 10

Staphyliiius ochripennis Menetries 1832:145

Quedius ochripennis'. Cameron 1932:289"’

Description. — Black, elytra pale red, abdomen slightly iridescent. Palpi dark brown, antennae dark

brown with three basal segments darker; legs piceous-brown with slightly paler tarsi, inner portions of

middle and posterior tibiae darkened. Head of obtusely quadrangular shape, wider than long (ratio 1.21),

slightly narrowed posteriorly, posterior angles broadly arcuate; eyes moderately large and convex, slightly

protruding from lateral contours of head; tempora as long as length of eyes seen from above; no additional

setiferous punctures between anterior frontal punctures; posterior frontal puncture situated closer to

posteromedian margin of eye than to posterior margin of head, one additional setiferous puncture between it

and posteromedian margin of eye and two setiferous punctures between it and posterior margin of head;

temporal puncture situated closer to posterior margin of head than to posterior margin of eye; tempora

punctate; surface of head with dense and fine microsculpture of transverse waves with intermixed

microscopic punctures. Antenna rather short, segment 3 distinctly longer than segment 2, segment 4 slightly

longer than wide, segment 5 as long as wide, segments 6-10 transverse, gradually becoming shorter and

wider, segment 1 1 about as long as two preceding segments combined. Pronotum slightly wider than long

(ratio 1.11), basal margin continuously rounded with lateral margins, lateral margins slighty, arcuately

naiTowed anteriorly; dorsal rows each with three punctures; sublateral rows each with three punctures,

posterior puncture situated behind level of large lateral puncture; surface of pronotum with microsculpture

similar to that on head, waves becoming gradually much finer toward lateral margins. Scutellum impunctate,

with very fine and dense microsculpture of transverse waves. Elytra at base hardly narrower than pronotum

at widest point and hardly widened posteriorly, at suture as long as, at sides somewhat longer (ratio 1.22)

than pronotum at midline; punctation fine and not dense, interspaces between punctures mostly distinctly

larger than diameters of punctures; pubescence not den.se, testaceo-brunneous; surface between punctures

without microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical

seam of palisade setae; punctation usually indistinctly finer than that of elytra, pubescence black.

Male. First four segments of front tarsus more dilated than in female. Sternites 6 and 7 with small more

densely punctate and dark pubescent area in middle of basal half; apical margin of sternite 8 with shallow

arcuate emargmation (Fig. 32), small triangular area before emargination flattened and smooth. Aedoeagus

(Figs. 33, 34) rather small; median lobe dilated and then slightly nan'owed toward truncate apical margin

with small median emargination, face adjacent to paramere with two short longitudinal carinae. Paramere

with lancetdike apical portion, apical margin subtruncate to truncate, not quite or Just about reaching apex of

median lobe; four apical setae and two short setae at each lateral margin below apex; underside of paramere

with several iiTCgularly placed sensory peg setae along each lateral margin.

Length 8.0-10.2 mm.

Type material. — Menetries (1832:145) described the species from specimens

collected under rocks at “Perimbal sur les montagnes de Talyche” (Talysh

Mountains on the border between Soviet Union and Iran). I did not study the

'Complete references and synonymy of this species can be found in several revisional papers {e.g.. Smetana

1958:345; Bordoni 1974:60).

Quaest. Ent., 1988, 24 (2)

208

Smetana

Map 10. Distribution records for: Quedius flavocaudatus ( % Q. antennalis ( aC); Q. birmanus ( ^ ); and Q. ochripemns ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

209

original series but there is little doubt that this species as it has been interpreted by

subsequent authors is the species which Menetries described.

Geographical distribution. — Qiiediiis ochripeimis is widely distributed

throughout Europe (except for the northern parts), the mediterranean area (including

North Africa) and farther east through Asia Minor, Iran, the Caucasus, the

middle-Asiatic republics of Soviet Union, Afghanistan and Pakistan to the western

portion of the Himalayan range (Map 10).

Material studied. — 1 specimen.

INDIA, Himachal Pradesh. Simla Hills, Narkanda, 9230', 14. IX. 3 1 , Cameron (BMNH) 1.

PAKISTAN. Khagan Valley. Naran, 2370-2750 m (Coiffait 19825:234, specimen not seen).

Bionomics. — Nothing is known about the habits of the Himalayan specimens.

Comparisons. — Qiiedius ochripeimis can easily be recognized among the

Himalayan species, in addition to the shape of the aedoeagus, by the presence of one

setiferoLis puncture between posterior frontal puncture and posteromedian margin of

eye, in combination with the red elytra.

1 1. Quedius (Microsaurus) fulgidus (Fabricius)

Sraphylinns fidgidus Fabricius 1787:220

Quedius fidgidus\ Cameron 1932:288

Complete references, synonymy and detailed description can be found in several

revisionary papers {e.g., Smetana 1958:349, 1971:66; Bordoni 1974:65); they will

not be repeated here.

Length 7.0-1 1.5 mm.

Geographical distribution. — Quedius fulgidus is widely distributed in the

Palaearctic region, particularly in Europe. Its mostly synanthropic occurrence

provides an excellent opportunity for introduction; it is therefore not suiprising that

the species was introduced into many parts of the world, so that it is now nearly

cosmopolitan in distribution.

Cameron (1932:289) records Q. fulgidus from “Northern India”. 1 have not seen

any specimens of this provenance in the Cameron collection in the British Museum.

I have also not seen any specimens from the Himalayan region in any of the

materials studied; however, the occurrence of this species in the Himalaya is

possible.

Classification and comparisons. — Quedius fulgidus can readily be distinguished

from all other species of the subgenus Microsaurus from the Himalayan region by

the following combination of characters: piceous-black to black, elytra red; posterior

frontal puncture on head situated distinctly closer to posteromedian margin of eye

than to posterior margin of head, no additional setiferous puncture between it and

posteromedian margin of eye; sublateral rows on pronotum each with 3-7 often

irregularly located punctures, last puncture located considerably behind level of

large lateral puncture; abdominal sternites 6 and 7 each with small, more densely

punctate and yellowish pubescent area in middle of basal half.

Quaest. Ent., 1988, 24 (2)

210

Smetana

Among the species occurring in the Himalayan region, only Q. ochripennis

resembles in general aspect Q. fulgidus\ however, Q. ochripennis can readily be

distinguished by the presence of an additional setiferous puncture between the

posterior frontal puncture on the head and the posteromedian margin of the eye.

12. Quedius (Microsaurus) placidus Cameron

Figs. 35-38; Map 1 1

Quedius placidus Cameron 1932:282

Description. — Piceous-black, apical margins of abdominal tergites feebly to slightly paler,

pronotum or elytra sometimes paler; abdomen slightly iridescent; palpi testaceo-brunneous, antennae

rufo-brunneous, legs brunneous, middle and hind tibiae and middle femora darkened, tarsi paler. Head of

rounded quadrangular shape, about as long as wide, posterior angles entirely rounded, indistinct; eyes small

and flat, hardly protruding from lateral contours of head, tempora much longer than length of eyes seen from

above (ratio 1.66); no additional setiferous punctures between anterior frontal punctures; posterior frontal

puncture situated much closer to posteromedian margin of eye than to posterior margin of head, one or two

setiferous punctures between it and posterior margin of head (close to the latter); temporal puncture situated

distinctly closer to posterior margin of head than to posterior margin of eye; surface of head with extremely

fine and dense microsculpture of transverse waves. Antenna moderately long and moderately thickened

toward apex, segment 3 distinctly longer than segment 2, segment 4 slightly longer than wide, segment 5 as

long as wide, following segments gradually becoming shorter, penultimate segment moderately transverse,

last segment shorter than two preceding segments combined. Pronotum barely wider than long (ratio 1.07),

broadly arcuate basally, markedly narrowed anteriorly; dorsal rows each with three punctures; sublateral

rows each with two punctures, posterior puncture situated slightly behind level of large lateral puncture;

microsculpture of pronotum similar to that on head. Scutellum impunctate. Elytra fairly long, at base

nan'ower than pronotum at widest point, at suture about as long as, at sides feebly longer than pronotum at

midline (ratio 1.11), finely and not densely punctate, each with sutural, discal and lateral, irregular and not

conspicuous longitudinal rows of coarser setiferous punctures; surface between punctures without

microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of

palisade setae; punctation of tergites about equal to that on elytra, slightly coarser and denser on basal

portions, first visible tergite with middle portion impunctate; pubescence distinct, dark. Front tarsus dilated

in both sexes.

Male. First four segments of front tarsus slightly more dilated than in female. Apical margin of stemite 8

with shallow and not wide arcuate emargination (Fig. 35), small triangular area before emargination

flattened and smooth. Aedoeagus (Figs. 36-38) long and rather narrow; median lobe, except for tip, entirely

covered by paramere, markedly naiTowed into long apical part with subacute apex; paramere very long,

apically narrowed into rounded apex; four rather long setae at apical margin; sensory peg setae on underside

of paramere numerous, forming two irregular longitudinal rows.

Length 7. 8-8.0 mm.

Type material. — Two syntypes in the British Museum (Natural History),

London, are labelled as follows: Spec. No. 1 (male): “Type” (round label with red

margin)/ “SYN-TYPE” (round label with blue margin)/ “Ghum dist. Rongdong

Valley V-VI-31 Dr. Cameron”/ “Quedius placidus Cam. TYPE”/ “M. Cameron.

Bequest. B.M. 1955-147.”. Spec. No. 2 (female): same labels as Spec. No. 1, except

for label “Type”.

The first (male) specimen was dissected and the sternite 8 and the aedoeagus

were mounted in Canada Balsam, the genital segment was glued to plate with beetle.

The specimen is hereby designated as the lectotype of Q. placidus-, the label

“Lectotype Quedius placidus Cameron Smetana des. 1984” has been attached to it.

Revision of the Tribes Quediini and Atanygnathini

211

Geographical distribution. — Quedius placidus is distributed in the eastern

portion of the Himalaya, from the Darjeeling area to Bhutan (Map 1 1 ).

Material studied. — 6 specimens.

BHUTAN. Tongsa, 2150 m, 24.VI.72 (CNCC, NHMB) 3; Tangu, 22 km N Thimphu, 30.VI.72

(NHMB) 1.

INDIA. West Bengal. See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Quedius placidus can easily be recognized by the small and flat

eyes in combination with the chaetotaxy of the head, and the presence of irregular

longitudinal rows of coarser punctures on the elytra.

For a comparison of Q. placidus and Q. lesagei see the discussion under the

latter species.

13. Quedius (Microsaurus) lesagei spec.nov.

Figs. 39^1; Map 13

Description . — Piceous-black with pronotum usually feebly paler, anterior portion of head narrowly

and indefinitely paler, sutural and apical margins of elytra narrowly paler in some specimens, basal portions

of tergites, pleurites and apex of abdomen paler, rather dark rufo-brunneous; abdomen slightly iridescent;

palpi testaceous, first three antennal segments rufo-testaceous, remaining segments piceous, legs

rufo-testaceous, medial face of middle and especially hind tibiae darkened. Head short, rounded and little

constricted behind eyes, wider than long (ratio 1.31 ); eyes large, moderately convex, feebly protruding from

lateral contours of head; tempora considerably shorter than length of eyes seen from above (ratio 0.24); no

additional setiferous punctures between anterior frontal punctures; posterior frontal puncture situated close

to posteromedial margin of eye, separated from it by distance no larger than diameter of puncture, two

punctures between it and posterior margin of head; temporal puncture separated from posterior margin of

eye by distance slightly smaller than diameter of puncture; surface of head with fine and dense

microsculpture of irregular transverse waves gradually changing into irregular meshes on anterior portion,

and with sparse microscopic punctures. Antenna short and moderately dilated toward apex, segment 3

slightly longer than segment 2, segment 4 slightly longer than wide, segments 5-8 about as long as wide,

gradually becoming slightly wider and shorter, segments 9 and 10 slightly transverse, last segment shorter

than two preceding segments combined. Pronotum feebly wider than long (ratio 1.09), broadly arcuate

basally, moderately narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with 1

or 2 punctures close to anterior margin; microsculpture of pronotum finer and denser than that on head.

Scutellum impunctate. Elytra fairly long, at base narrower than pronotum at widest point, at suture about as

long as, at sides feebly longer than pronotum at midline (ratio 1.09), moderately densely and finely punctate,

each with sutural, discal and lateral, irregular and inconspicuous longitudinal rows of coarser setiferous

punctures; surface between punctures without microsculpture. Wings fully developed. Abdomen with tergite

7 (fifth visible) bearing whitish apical seam of palisade setae; punctation of tergites slightly finer than that of

elytra, sparse, somewhat denser at tergal bases, first visible tergite with middle portion impunctate;

pubescence dark. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus slightly more dilated than in female. Apical margin of stemite 8

with shallow and not wide, obtusely triangular emargination (Fig. 39), small triangular area before

emargination flattened and smooth. Aedoeagus (Figs. 40, 41) rather small, narrow and elongate; median lobe

vaguely constricted in middle portion, slightly asymmetrical and with arcuate apex; paramere very narrow

and elongate, almost parallel-sided and with obtusely arcuate apex, by far not reaching apex of median lobe;

four rather small apical setae and two longer setae at each lateral margin below apex; sensory peg setae on

underside of paramere forming irregular group at apical margin, with a few larger peg setae posteriad of it.

Length 5. 7-6. 5 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Dis. Arun Valley at Num main bridge 1050 m. 22.IV. 1984 Smetana & Lobl”. In the

Qiiaest. Ent., 1988, 24 (2)

212

Smetana

collection A. Smetana, Ottawa, Canada.

Paratypes (4): Nepal: same data as holotype (ASCC, CNCC, MHNG) 4.

Geographical distribution. — Quedius lesagei is known from the Arun river

valley near Num in eastern Nepal (Map 13).

Bionomics. — The specimens of the original series were taken by sifting

mushrooms, moss, rotting wood and bark on an old fallen tree.

Comparisons. — Quedius lesagei resembles Q. placidus in general habitus and in

the presence of irregular and inconspicuous longitudinal rows of coarser setiferous

punctures on the elytra; however, Q. placidus can easily be distinguished, in

addition to the differences on the aedoeagus, by the considerably smaller eyes (see

the description).

Etymology. — Patronymic; the species was named in honour of my colleague and

friend. Dr. L. LeSage from the Coleoptera Unit, Biosystematics Research Centre,

Ottawa.

14. Quedius (Microsaurus) stevensi Cameron

Figs. 42-44, 396-399; Map 1 1

Quedius stevensi Cameron 1932:288

Quedius analoka SmeVdna 1975:331 {syn.nov.)

Quedius queinneci Coit'fad 1983a: 167 (syn.nov.)

Description . — Piceous to almost black, elytra and/or pronotum occasionally paler, or latter with

indistinctly paler margins, apical margins of abdominal tergites and apex of abdomen more or less, usually

indistinctly, paler; abdomen slightly iridescent; palpi rufopiceous, antennae piceous with three basal

segments usually darker, legs piceous with at least partially darkened tibiae, and paler, rufo-testaceous tarsi.

Head rounded, slightly wider than long (ratio 1.16); eyes large and convex, tempora distinctly shorter than

length of eyes seen from above (ratio 0.55); no additional setiferous punctures between anterior frontal

punctures; posterior frontal puncture situated close to posteromedian margin of eye, separated from it by

distance about equal to diameter of puncture, two additional punctures between it and posterior margin of

head; temporal puncture situated closer to posterior margin of eye than to posterior margin of head; tempora

with a few punctures; surface of head with very fine and dense microsculpture of transverse waves. Antenna

moderately long, segment 3 slightly longer than segment 2, following segments longer than wide, gradually

becoming shorter and slightly wider, segments 9 and 10 about as long as wide, last segment shorter than two

preceding segments combined. Pronotum slightly wider than long (ratio 1.15), widely arcuate basally and

rather markedly narrowed anteriorly, evenly transversely convex; dorsal rows each with three punctures;

sublateral rows each with three punctures (two first ones close to apical margin), posterior puncture situated

distinctly behind level of large lateral puncture; surface of pronotum with microsculpture similar to that on

head but slightly finer and denser. Scutellum impunctate. Elytra fairly long, at base only slightly narrower

than pronotum at widest point, at suture slightly (ratio 1.17), at sides distinctly longer (ratio 1.30) than

pronotum at midline; punctation coarse and deep, interspaces between punctures about equally large as

diameters of punctures; pubescence rather sparse; surface between punctures without microsculpture. Wings

fully developed. Abdomen with tergite 7 (fifth visible) bearing distinct whitish apical seam of palisade setae;

punctation on basal portions of tergites coarse but more or less finer than that on elytra, becoming finer

toward apical margins of tergites, middle portion of first visible tergite and apical portions of all tergites

almost impunctate, punctation in general becoming gradually finer toward apex of abdomen. Front tarsus

dilated in both sexes.

Male. First four segments of front tarsus distinctly more dilated than in female. Apical margin of stemite

7 feebly concave, apical margin of stemite 8 with wide and deep triangular emargination (Fig. 43), small

triangular area before emargination flattened and smooth. Aedoeagus (Fig. 44) large and voluminous, apical

portion of median lobe of complicated, quite characteristic shape, deeply longitudinally excavate medially,

Revision of the Tribes Quediini and Atanygnathini

213

excavation accomodating apical portion of paramere. Paramere very wide basally, of quite characteristic

shape, its extremely narrow apex slightly exceeding apex of median lobe, curved dorsally; underside of

paramere without sensory peg setae, but with subapical lateral crest at both lateral margins of apical part;

one very small seta at each side of end of crest.

Length 6. 9-8. 6 mm.

Type material. — Qiiediiis stevensi. The original series in the collection of the

British Museum (Natural History), London, contains one female specimen

(holotype) under the name of Q. stevensi. It is labelled as follows: “Type” (round

label with red margin)/ “Sikkim: Gopaldhara, Rungbong Vail. H. Stevens.”/ “H.

Stevens. Brit Mus. 1922-307/”/ “Quedius stevensi Cam. TYPE.” Right antenna

except for basal segment, and three apical segments of left hind tarsus are missing in

this specimen.

Quedius analoka. The species was described from several specimens from

western and central Nepal. For details about the original material see Smetana

1975:332. The holotype, allotype and two paratypes are deposited in the collection

Franz, Mbdling, Austria, and two paratypes (from around Ainorasha near Marpha

and from Dinguari Kola Valley) in the Canadian National Collection, Ottawa (CNC

No. 13964). The holotype of this species cannot be specifically distinguished from

the holotype of Q. stevensi; the name Q. analoka is a junior synonym of Q. stevensi.

Quedius queinneci. Coiffait (l.c.) described the species from a single female

specimen from Nepal. The holotype in the Coiffait collection. Museum National

d’Histoire Naturelle, Paris, France, is labelled as follows: “Nepal VII-82 Ganesh

Himal Salme a Ruchet”/ “1500-3400”/ “HOLOTYPE”/ “Quedius (Sauridus)

queinneci H. Coiffait 1983”. The holotype is a slightly teneral specimen that cannot

be specifically distinguished from the holotype of Q. stevensi; the name Q.

queinneci is a junior synonym of Q. stevensi. My determination label “Quedius

stevensi Cameron Smetana det. 1984” has been attached to this specimen.

Geographical distribution. — Quedius stevensi is distributed from western Nepal

eastward to Sikkim (map 1 1)

Material studied. — 67 specimens.

INDIA. Sikkim. See Type material.

NEPAL. Manang Distr. Marayandi, Thimang, 2550 m, 14- 17. IV. 80, Martens & Ausobsky (SBMF) 8;

Marayandi, above Bagarchhap, 2300-2400 m. 12- 14. IV. 80, Martens & Ausobsky (SBMF) 2. Parbat Distr.

between Chitre and Ghandrung, 2800-2900 m, 4-7. V. 80, Martens & Ausobsky (SBMF) 1; Pass N Jiri, 2600

m, 10. IV. 73, J. Martens (SBMF) 1; Pun Hill at Ghoropani Pass, 3030-3100 m, 8.X.83, Smetana & Lobl

(MHNG) 2; Below Thare Pati, 9-1 1.IV.81, Lobl & Smetana (ASCC, MHNG) 6; near Mere Dara, 3000 m,

7.IV.81, Lobl & Smetana (ASCC) 1; Malemchi, 2800 m, 18. IV. 81, Lobl & Smetana (ASCC, MHNG) 3;

above Shermantang, 2900 rn, 26.IV.81, Lobl & Smetana (MHNG) 1; Dobate Ridge NE Barahbise, 2800 m,

2.V.81; same, 3000 m, 7.V.81, all Lobl & Smetana (ASCC, MHNG) 4. Kathmandu Distr. Siwapuri Dara,

2400 m, 29. and 30. IV. 85, A. Smetana (ASCC, CNCC) 9. Khandhari Distr. Forest above Ahale, 25. III. 82,

2400 m, A. & Z. Smetana (ASCC) 1; above Tashigaon, 3500-3600 m, 6. IV. 82, A. & Z. Smetana (ASCC) 3;

“Bakan” W of Tashigaon, 3250 m, 4. IV. 82, A. & Z. Smetana (ASCC) 1; Mure Num, 1500-1900 m,

25.V.80, W. Wittmer (MHMB) 1. Lalitpur Distr. Phulcoki, 2600-2650 m, 21.III.-14.V.80, Martens &

Ausobsky (SBMF) 5; same, 2600-2650 m, 20.-22.1V. 82, A. & Z. Smetana (ASCC, CNCC) 8; same, 2550

m, 29.IV.84, Smetana & Lobl (ASCC) 2; same, 2500 m, 28-29.IV.84, Lobl & Smetana (MHNG) 2. Rasuwa

Distr. Langtang Khola Valley, 2.5 km E Syabru, 1730 m, 14. IV. 85, A. Smetana (ASCC) 1.

Quaest. Ent., 1988, 24 (2)

214

Smetana

Map 11. Distribution records for: Quedius placidus ( A ); and Q. stevensi ( • ).

Revision of the Tribes Quediini and Atanygnathini

215

Bionomics. — Qiiedius stevensi oeeurs at elevations between 1500 and 3600 m.

Most speeimens were taken by sifting leaf litter, moss and various debris in forests,

often coniferous; by sifting dead vegetation and debris at bases of large rocks on

clearings of forests; by sifting wet leaf letter and debris sprinkled by a waterfall;

some specimens were taken in pitfall traps.

Comparisons and variations. — Qiiedius stevensi is a rather distinctive species. It

is well characterized, in addition to the conspicuous aedoeagus (Fig. 44), by the

coarse punctation of elytra in combination with the chaetotaxy of the pronotum

(sublateral rows), the punctation of abdominal tergites (see description), and the

deep and wide emargination of sternite 8 in the male (Fig. 43). For a comparison

with Q. ripicola and Q. milansaar, see under the two respective species.

The punctation of the elytra varies to some extent and appears not quite as coarse

in some specimens. The female holotype is such a specimen.

15. Qiiedius (Microsaurus) ripicola Cameron

Figs. 45-49; Map 12

Qiiedius ripicola Cameron 1926:269; 1932:291

Qiiedius sodalis Cameron 1926:267; 1932;287 (syn.mn’.)

Description. — In all external characters quite similar to Q. stevensi but different as follows:

pronotum nanower, about as long as wide, less narrowed anteriorly, in some specimens about equally

narrowed both anteriorly and posteriorly; elytra at base almost as wide as pronotum at widest point;

punctation of elytra and abdominal tergites of same type as that of Q. stevensi but in general finer and

somewhat denser.

Male. First four segments of front tarsus dilated in similar way as in Q. stevensi. Apical margin of

sternite 8 with shallow and moderately wide triangular emargination (Fig. 43), triangular area before

emargination flattened and smooth. Aedoeagus (Figs. 46^9) narrow and elongate; median lobe narrowed

toward apex, slightly asymmetrical apically and with subacute apex; apical portion forming a short rounded

hook in lateral view; internal sac as in Fig. 47. Paramere narrow and elongate, narrowed anteriorly, by far

not reaching apex of median lobe, with apex obtuse, subtruncate or minutely subemarginate; four minute

setae at apical margin and one stronger and longer seta at each lateral margin well below apex, usually

accompanied, at least unilaterally, by a minute seta; underside without sensory peg seate but with some thick

minute setae.

Length 6. 8-8. 3 mm.

Type material. — Qiiedius ripicola. The collection of the British Museum

(Natural History), London, contains 1 1 conspecific specimens in the original series

of Q. ripicola. They are labelled as follows; Spec. No. 1 (d); “Type” (round label

with red margin)/ “Keyarkuli Mussoorie”/ “Dr. Cameron. 17. IV. 22”/ “Type Quedius

ripicola Dr. Cameron”/ “SYN-TYPE” (round label with blue margin). Spec. Nos.

2-f (1?, 2dd); “Chakrata Dist. Sanj Khud 6500’”/ “Dr. Cameron 29.V.22”/ “M.

Cameron. Bequest. B.M. 1955-147”. Spec. No. 5 (ci'); “Chakrata Dist. Dodora Khud

8000'”/ “Dr. Cameron 6.V.22”/ “M. Cameron. Bequest. B.M. 1955-147”. Spec. No.

6 (d); “Mohna, 5000'. Chakrata, U.P.”/ “Dr. Cameron 2.V.21.”/ “Quedius ripicola

Cam”. Spec. Nos. 7-9 (2$?, Id); “Bindal Gadh, Konain”/ “Dr. Cameron 5.V.21”/

“M. Cameron. Bequest. B.M. 1955-147”. Spec. No. 10 (?); “Gahan 7000' Simla

Hills.”/ “Dr. Cameron IX. 1921”/ “M. Cameron. Bequest. B.M. 1955-147.” Spec.

Quaest. Ent., 1988, 24 (2)

216

Smetana

No. 1 1 (ci'): “Theog 7600 Simla Hills.”/ “M. Cameron. 1 1 -IX- 1921.”/ “M. Cameron.

Bequest. B.M. 1955-147.”. Specimen Nos. 2-11 bear as last label the round label

“SYN-TYPE” with blue margin.

All males were dissected and parts were glued to plates with beetles. The first

male specimen, bearing the label “Type” (aedoeagus, genital segment and sternite 8

were glued to plate with specimen, which is slightly teneral) is hereby designated as

the lectotype of Q. ripicola; the label “Lectotype Q. ripicola Cam. Smetana des.

1983” has been attached to it.

Quedius sodalis. The collection of the British Museum (Natural History),

London, contains one female specimen under the name Q. sodalis. It is labelled as

follows: “Type”/ “Fagu 8000' Simla Hills. Dr. Cameron. 6.IX.1921.”/ “TYPE”/

“Quedius sodalis Dr. Cameron”/ “M. Cameron. Bequest. B.M. 1955-147.”

Since Cameron (1926:368) did not specify that the species was described from a

unique, the specimen is hereby designated as the lectotype of Q. sodalis’, the label

“Lectotype Q. sodalis Cam. Smetana des. 1983” has been attached to it. I am unable

to distinguish this female from larger specimens of Q. ripicola. The name Q. sodalis

should be considered a junior synonym of Q. ripicola’, my corresponding

determination label was attached to it.

Geographical distribution. — Quedius ripicola is distributed from Himachal

Pradesh and Uttar Pradesh eastward to eastern Nepal (Map 12).

Material studied. — 71 specimens.

INDIA. Himachal Pradesh. Simla Hills. Theog, 7600', 10-11.IX.21, Cameron (ASCC, BMNH) 15;

Simla, VI. 1981, de Rougemont (GDRC) 2. Uttar Pradesh. Chakrata distr. Bangar, 6000', Cameron (BMNH)

1; Bindal Gadh, Konain, Cameron (BMNH) 2; Khedar Khud, 7500', 7.V.22, Cameron (BMNH) 1; Korawa

Khud, 9100', 4.V.22, Cameron (BMNH) 3; Manjgaon, 6500', 18.V.22, 5.V.22, Cameron (BMNH) 8; Sainj

Khud, 6500', 29.V.22, Cameron (BMNH, CNCC) 2. Kumaon; N. Kumaon, Gori Riv. gorge, 5000-9000',

Champion (BMNH) 2. Mussoorie: Keyarkull, 17. IV. 22, Cameron (CNCC) 1. Garhwal: 20 km S Chamba,

1150 m, 20.IX.79,Lobl(MHNG) 1.

NEPAL. Manang Distr. Forest W Bagarchhap, 2200-2250 m, 21. and 22. IX. 84, Smetana & Lobl

(ASCC, CNCC, MHNG) 13. Khandhari Distr. Chichila nr. Ahale, 2200 m, 4.IV.84, Lobl & Smetana

(MHNG) 1; Induwa Khola Valley, 2000 m, 17. and 18. IV. 84, Smetana & Lobl (ASCC, CNCC, MHNG) 5; 2

km E Mansingma, 1900 m, 10. IV. 84, Lobl & Smetana (MHNG) 1. Chaubas, 2600 m, 5.IV.81, Lobl &

Smetana (ASCC) 1.

Bionomics. — Little is known about the habitat requirements of this species.

Cameron (1932:291) gives “margins of streams” as the habitat. The specimens from

near Bagarchhap in Nepal were taken by sifting wet moss, debris and vegetation

from close to edges of small creeks or near small waterfalls, and from the vegetation

soaked with water on huge flat rock tables in a forest. The specimen from Chaubas,

Nepal, was taken by sifting wet debris and leaf litter from around a seepage in an

oak-rhododendron forest, and the specimens from the Induwa Khola Valley and

from near Mansingma were taken by sifting wet flood-debris from sandy banks of

Induwa Khola and of a small creek.

Comparisons. — As mentioned above, Q. ripicola is quite similar in all external

characters to Q. stevensi. The difference in the shape of the pronotum is mostly

distinct; however, the difference in the punctation of the elytra and that of the

Revision of the Tribes Quediini and Atanygnathini

217

abdominal tergites is less reliable and some specimens of Q. ripicola do not

appreciably differ in this character from those of Q. stevensi. On the other hand, the

males of both species differ considerably in the development of the apical

emargination of the sternite 8 (see Figs. 43, 45), and in the shape of the aedoeagus

(see Figs. 44, 46).

Cameron (1932:191) records this species also from Darjeeling Distr. (Lopchu)

and from Sikkim. However, I failed to find any specimens of Q. ripicola from these

localities in the British Museum. These records very likely refer to another species,

most likely to Q. stevensi. Qiiedius ripicola does not seem to occur east of Nepal.

16. Quedius (Microsanrus) milansaar spec. now

Figs. 50-54; Map 12

Description. — In all external characters extremely similar to Q. ripicola and different only in shape

of aedoeagus.

Male. First four segments of front tarsus dilated in similar way as in a Q. stevensi and Q. ripicola.

Apical margin of sternite 8 with emargination similar to that of Q. ripicola (Fig. 50), triangular area before

emargination flattened and smooth. Aedoeagus (Figs. 51-54) similar to that of Q. ripicola, but larger, apical

portion of median lobe fonning large elongate hook in lateral view; paramere longer and narrower, apical

margin minutely but distinctly emarginate; four minute apical setae, inner pair situated rather far from apical

margin; two long setae at each lateral margin well below apex situated very close to each other and therefore

appearing as one seta at low magnification; underside, as in Q. ripicola, without any sensory peg setae, but

with few thick minute setae.

Length 7. 5-8. 3 mm.

Type material. — Holotype (male): “INDIA: H.P. MacleodganJ VI: 1981”/

“Rougemont”. In the collection A. Smetana, Ottawa, Canada.

Paratypes (2) (males): India: Himachal Pradesh, Nagar, VI. 1981, de Rougemont

(GDRC, MHNG) 2.

Geographical distribution. — Quedius milansaar is at present known from two

localities in Himachal Pradesh (Map 12).

Bionomics. — Nothing is known about the habitat requirements of this species;

however, it very likely lives in a similar way as described for Q. ripicola.

Comparisons. — Quedius milansaar differs from Q. stevensi, in addition to the

quite different aedoeagus, by the same set of characters given for Q. ripicola. It is

sympatric with Q. ripicola in Himachal Pradesh and can be positively distinguished

from it only by the differently shaped aedoeagus (see Figs. 46—49, 51-54).

Etymology. — The specific name is the Nepali adjective milansaar (^friendly).

17. Quedius (Microsanrus) franzi Smetana

Figs. 55-59; Map 13

Quedius franzi Smetana 1975:326

Quedius trisnlensis Coiffait 1982a:79 (syn.nov.)

Description. — Testaceorufous to rufo-brunneous or almost brunneopiceous, abdomen apically or

almost entirely, and pronotum and head variably, in general not distinctly, darkened in paler coloured

Quaest. Ent., 1988, 24 (2)

218

Smetana

Map 12. Distribution records for: Qiiedius hpicola ( • ); and Q. milansaar ( ▲ ).

Revision of the Tribes Quediini and Atanygnathini

219

specimens, occasional specimens almost entirely testaceorufous; abdomen slightly iridescent; palpi,

antennae and legs rufo-testaceous, middle and especially posterior tibiae darkened at inner margin. Head of

rounded quadrangular shape, feebly wider than long (ratio 1.13), posterior angles rounded, indistinct; eyes

rather large and convex, moderately protruding from lateral contours of head, tempora shorter than length of

eyes from above (ratio 0.72); no additional setiferous punctures between anterior frontal punctures; posterior

frontal puncture situated much closer to posteromedian margin of eye than to posterior margin of head, two

or three additional setiferous punctures posteromediad and usually one anteriad of it, posteromedian

additional punctures form with two usual punctures near posterior margin of head an irregular group of four

or five punctures; temporal puncture situated about midway between posterior margin of eye and posterior

margin of head, one additional setiferous puncture between it and posterior margin of eye; tempora with

several fine punctures; surface of head with very fine and dense microsculpture of irregular transverse waves

with numerous longitudinal connections becoming gradually more numerous anteriorly, anterior part of

clypeus therefore covered by almost meshed microsculpture. Antenna moderately long, segment 3 distinctly

longer than segment 2, segment 4 distinctly longer than wide, following three segments longer than wide,

gradually becoming shorter, segments 8-10 about as long as wide, last segment shorter than two preceding

segments combined. Pronotum somewhat wider than long (ratio 1.14), broadly arcuate basally, arcuation

flattened in middle portion; lateral margins somewhat flattened and subparallel in basal half, then distinctly

narrowed anteriorly; lateral portions of pronotum feebly and narrowly explanate posterolaterally; dorsal

rows each with three punctures; sublateral rows each with three rather fine punctures, posterior puncture

situated distinctly behind level of large lateral puncture; usually one or two additional punctures between

dorsal and sublateral rows; microsculpture of pronotum similar to that on vertex of head. Scutellum

impunctate. Elytra rather short, narrower than pronotum at widest point, at suture shorter (ratio 0.76), at

sides hardly shorter (ratio 0.94) than pronotum at midline; punctation rather fine and slightly asperate,

moderately dense, pubescence brownish, surface between punctures with extremely fine microscopic

irregularities. Wings reduced to small nonfunctional stumps reaching about half of length of elytra.

Abdomen with tergite 7 (fifth visible) lacking whitish apical seam of palisade setae; punctation of abdominal

tergites more or less finer and about equally dense to slightly sparser than that on elytra, ecoming gradually

somewhat sparser toward apex; pubescence as on elytra. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus much more dilated than in female. Stemite 8 with three long

and strong setae on each side of apical half; apical margin with moderately wide and rather shallow, arcuate

emargination (Fig. 55), triangular area before emargination llattened and smooth. Aedoeagus (Figs. 56-59)

small, median lobe subacute apically, with small hook visible in lateral view when paramere removed.

Paramere not reaching apex of median lobe, obtuse and subemarginate apically, with four setae at apical

margin and two longer bristles at each lateral margin near apex; underside of paramere with two or three

sensory peg setae on each side near apex.

Fength 5.8-7. 1 mm.

Type material. — Quediiis franzi. The species was described from three

specimens, all bearing identical locality labels as follows; “Weg v. Gosaikunde z.

Fulungmonastery’V “Zentral-Nepal Sept.-Okt. 1971 Ig. H. Franz”; the underside of

the first label bears code “Pa 170”. The male holotype and female allotype are

deposited in the Franz collection, Modling, Austria; the holotype bears red label

“HOLOTYPE Quedius franzi A. Smetana 1975”. The male paratype is deposited in

the Canadian National Collection, Ottawa (CNC No. 13963).

Quedius trisidensis. Coiffait (1982a:79) described the species from a single

female from basically the type locality of Q. franzi. The holotype is deposited in the

Senckenberg Museum, Frankfurt a.M., Federal Republic of Germany. It is labelled

as follows: “Trisuli-Tal Gosaikind, 3200 m 23.-26.IV.1973”/ “NEPAL-Expedition

Jochen Martens”/ “TYPE”/ “Holo-typus” SMF C 15109 (underside)/ “Quedius

(Microsaurus) trisulensis H. Coiffait 1979”.

The specimen is a perfectly typical female of Q. franzi, which cannot be

distinguished from the allotype of Q. franzi; the name Q. trisulensis is a junior

Quaest. Ent., 1988, 24 (2)

220

Smetana

synonym of Q. franzi. The holotype was accordingly labelled.

Geographical distvihution. — Qiiedius franzi is known only from higher

elevations (above 3000 m) in Central Nepal (map 13).

Material studied. — 38 specimens.

NEPAL. Nuwakot Distr. between Ghopte and Thare Pati, 23-26. IV. 85, 3150-3250 m, A. Smetana

(ASCC, CNCC) 14; below Thare Pati, 3300 m, 10. IV. 81, Lbbl & Smetana (ASCC) 2; Yangri Ridge, 4200

m, 21.IV.81, LobI & Smetana (ASCC, MHNG) 5. Rasuwa Distr. north slope above Syabru, 3600-3800 m,

17-19.IV.85, A. Smetana (ASCC, BMNH) 11. Aim Dugdinma b. Lughla, Khumbu, 300(M000 m, Franz

(HFCC) 2.

Bionomics. — Quedius franzi occurs above 3000 m; the original series was

collected by sifting forest floor litter in a fir forest with Rhododendron undergrowth;

other specimens were taken by sifting debris, grass roots and moss on and around

piles of rocks on a forest meadow (below Thare Pati); by sifting wet moss, fallen

leaves and fir needles in a ravine (north slope above Syabru); by sifting moist to wet

debris, leaf litter, etc., near creeks, seepages and on large rocks in a Rhododendron -

Abies forest (between Ghopte and Thare Pati); specimens from Yangri Ridge were

taken (together with Q. angnimai) by sifting debris, fallen leaves and moss under

low shrubs on a meadow (pasture) above tree line.

Comparisons and variations. — Quedius franzi is a rather conspicuous species; it

can easily be recognized by the chaetotaxy of the head and pronotum, in

combination with the shape of the pronotum, the colouration, the rather short elytra

and the fifth visible abdominal tergite lacking the whitish apical seam of palisade

setae. On the other hand, it is closely related and similar to Q. goropanus (see the

discussion under the latter species for the differences between these two species).

The chaetotaxy of the posterolateral part of the head is rather unstable; some

punctures can be missing, or additional punctures can be present (see the

description). The holotype of Q. trisulensis is a specimen with additional puncture

(bilaterally) anteriad of posterior frontal puncture. This additional puncture occurs in

almost all specimens, varies considerably in size and is rarely doubled.

18. Quedius (Microsaurus) goropanus Smetana

Figs. 60-64; Map 14

Quedius goropanus Smetana 1975:327

Description. — Plceous-black, pronotal margins or almost entire pronotum, elytra and very apex of

abdomen more or less paler, rather piceous to piceous-brown, some specimens piceous-black with black

head; abdomen slightly iridescent; palpi, antennae and legs rufo-testaceous, front tibiae slightly, middle and

posterior tibiae distinctly, darkened at inner margin. Head of rounded quadrangular shape, wider than long

(ratio 1.23), markedly narrowed behind eyes, posterior angles rounded, indistinct; eyes large and convex,

protruding from lateral contours of head, tempora distinctly shorter than length of eyes seen from above

(ratio 0.66); no additional setiferous punctures between anterior frontal punctures, however, two small round

depressions (without setae) between eyes; posterior frontal puncture situated much closer to posteromedian

margin of eye than to posterior margin of head, two setiferous punctures between it and posterior margin of

head; temporal puncture very close to posterior margin of eye, separated from margin of eye by scarcely

diameter of puncture; tempora almost lacking fine punctures; surface of head with dense and very fine

microsculpture of irregular transverse waves with numerous longitudinal connections becoming gradually

Revision of the Tribes Quediini and Atanygnathini

221

more numerous laterally and anteriorly, anterior part of clypeus therefore at least partially with almost

meshed microsculpture. Antenna moderately long, segment 3 longer than segment 2, segment 4 distinctly

longer than wide, following three segments longer than wide, gradually becoming shorter, segments 8-10

about as long as wide, last segment shorter than two preceding segments combined. Pronotum somewhat

wider than long (ratio 1.15), broadly arcuate basally, arcuation slightly flattened in middle portion; lateral

margins somewhat flattened and subparallel in basal half, then distinctly narrowed anteriorly; dorsal rows

each with three punctures; sublateral rows each with two or three punctures, posterior puncture situated

distinctly behind level of large lateral puncture; microsculpture of pronotum similar to that on vertex of

head. Scutellum impunctate. Elytra short, at base narrower than pronotum at widest point, at suture shorter

(ratio 0.71), at sides somewhat shorter (ratio 0.82) than pronotum at midline; punctation moderately coarse

and dense, slightly asperate, pubescence piceous, surface between punctures with extremely fine

microscopic irregularities. Wings reduced to small nonfunctional stumps. Abdomen with tergite 7 (fifth

visible) lacking whitish apical seam of palisade setae; punctation of abdominal tergites more or less finer

than that on elytra and about equally dense, sometimes becoming slightly sparser toward apex of abdomen.

Front tarsus dilated in both sexes.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with three

long and strong setae on each side of apical half; apical margin with wide and not deep, arcuate emargination

(Fig. 60), triangular area before emargination flattened and smooth. Aedoeagus (Figs. 61-64) rather small;

median lobe, except for small lateral lobe on each side just below middle, entirely covered by paramere;

apical portion of median lobe, when paramere removed, markedly narrowed anteriorly and with subacute

apex; internal sac as in Fig. 62. Paramere large, with apical portion wide, narrowed anteriorly and with

apical margin truncate to distinctly emarginate; one short apical seta at each lateroapical angle of paramere,

one similar short seta at each lateral margin just below apical seta, and two setae at each lateral margin well

posteriad of apical seta; underside of paramere with sensory peg setae forming two iiregular lateral groups

situated well posteriad of apex of paramere, each with 7 to 1 I peg setae.

Length 6. 9-7. 4 mm.

Type material. — The female holotype, deposited in the Franz collection,

Modling, Austria, is labelled as follows; “Umb.Goropani w. Pokhara”/

“Zentral-Nepal Sept.-Okt. 1971 Ig. H. Franz ”/ “HOLOTYPE Quedius goropanus A.

Smetana 1975”; the underside of the first label bears code “Pa 141 + 142”.

Geographical distribution. — Quedius goropanus is at present known only from

the vicinity of the Ghoropani Pass is west Nepal (Map 14).

Material studied. — 5 specimens.

NEPAL. Parhat Distr. Ridge E Ghoropani Pass, 3150 m, 7.X.83, Smetana & Lbbl (ASCC) 1; Pun Hill

at Ghoropani Pass, 3050-3100 m, 8.X.83, Smetana & Lbbl (CNCC, MHNG) 3.

Bionomics. — The holotype was found either under a stone or in rotting wood.

The specimens from Pun Hill were taken by sifting forest floor debris at the edges of

a coniferous forest; the specimen from a ridge near Ghoropani Pass was taken in a

mixed Abies, Rhododendron and Acer forest by sifting old mushrooms on fallen

trees.

Comparisons. — Quedius goropanus is related and similar to Q. franzi; however,

it differs, in addition to the differences in the shape of the aedoeagus, mainly by the

following characters: form more robust, size slightly larger, colouration in general

darker; head larger, more transverse and markedly narrowed behind eyes, with

distinctly different chaetotaxy, that is, the additional punctures on posterolateral

portion of head, which are present in Q. franzi, are absent in Q. goropanus, temporal

puncture in Q. goropanus is very close to posterior margin of the eye, whereas in Q.

franzi this puncture is situated about midway between posterior margin of the eye

and posterior margin of the head, and there is an additional puncture between it and

Quaest. Ent., 1988, 24 (2)

222

Smetana

the posterior margin of the eye.

Quedius goropanus is also closely related and quite similar to Q. tanderi and Q.

kaUo\ it can be positively distinguished from either species only by the differences

in the shape of the aedoeagus.

For a comparison of Q. goropanus with Q. angnimai see the discussion under the

latter species.

The small round depressions on the head between the eyes are present in all five

known specimens of Q. goropanus’, some of the specimens have similar additional

depressions also on the vertex of the head and/or on the pronotum behind the

posterior puncture of the sublateral rows.

19. Quedius (Microsaurus) tanderi spee.nov.

Figs. 65-68; Map 14

Deseription. — in all external characters, including general habitus and colouration, extremely

similar to Q. goropanus and different mainly by quite differently shaped aedoeagus. Form in general slightly

smaller and less robust, head in general smaller and less transverse, punctation of elytra in general finer and

denser.

Male. First four segments of front tarsus distinctly more dilated than in female. Sternite 8 with four long

and strong setae on each side of apical half; apical margin with arcuate emargination slightly narrower and

shallower than that of Q. goropanus (Figs. 60, 65), triangular area before emargination flattened and smooth.

Aedoeagus (Figs. 66-68) rather small; median lobe, except for subacute apex, entirely covered by paramere;

apical portion of median lobe, when paramere removed, short and markedly narrowed; internal sac as in Fig.

67. Paramere to great extent parallel-sided, truncate apically with apical margin slightly but distinctly

emarginate in middle and with four apical setae at each side of median emargination; underside of paramere

without sensory peg setae.

Length 6. 9-7. 4 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL (Prov.

Bagmati) Jangtang Ridge NE Barahbise, 3250 m, 5.V.81 Lobl & Smetana”. In the

collection A. Smetana, Ottawa.

Paratypes (6): same data as holotype, one specimen with elevation 3300 m

(ASCC, CNCC, MHNG) 6.

Geographical distribution. — Quedius tanderi is known only from the type

locality above Barahbise in eentral Nepal (Map 14).

Bionomics.- — The specimens of the original series were taken by sifting debris,

moss and low vegetation under bushes on a moist slope of a gully on the

northeastern slope of the Jangtang Ridge.

Comparisons. — Quedius tanderi can be positively distinguished from Q.

goropanus only by the different shape of the aedoeagus.

Two small round depressions (without setae), similar to those in Q. goropanus,

were found on the head between the eyes in three paratypes; also, some of the

paratypes have fairly large and very shallow depressions on each lateral portion of

the clypeus.

Etymology. — The specific name is the Nepali noun tanderi (=young man) in

apposition.

Revision of the Tribes Quediini and Atanygnathini

223

20. Qiiedius (Microsawus) kailo spec. now

Figs. 69-72; Map 14

Description . — In all characters, including general habitus and colouration, extremely similar to Q.

^oropaniis and different only by slightly different emargination of male stemite 8 and by differences on

aedoeagus.

Male. Sternite 8, as in Q. tauderi, with four long and strong setae on each side of apical half;

emargination of apical margin slightly shallower (Figs. 63, 69). Aedoeagus (Figs. 70-72) more robust and

somewhat larger; naiTowed apical portion of median lobe long; internal sac very similar to that of Q. tanderi

(Fig. 71). Paramere wider, with only three apical setae at each side of apical emargination, which is very

shallow.

Length 7. 1-7.3 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL -Janakpur

Thodung 3100 m 28/V/80 leg. E. Migliaccio”. In the collection A. Smetana, Ottawa,

Canada.

Geographical distribution. — Quedius kailo is at present known only from the

type locality in eastern Nepal (Map 14).

Bionomics. — Nothing is known about the habitat of the species.

Comparisons. — Quedius kailo is extremely similar to Q. tanderi and cannot be

distinguished by any external characters. Even the differences on the male sternite 8

and on the aedoeagus are of such a character that it is possible that Q. kailo actually

is only a geographically separated subspecies of Q. tanderi. This problem cannot be

solved at present; more detailed knowledge of the distribution of both Q. tanderi and

Q. kailo is needed.

The difference in the number of apical setae of the paramere in Q. tanderi (8)

and Q. kailo (6) may not be significant.

Etymology.— The specific name is the Nepali adjective kailo (brown). It refers

to the colour of the species.

21. Quedius (Microsaurus) angnimai spec. now

Eigs. 73-77; Map 14

Description . — In all characters, including general habitus and colouration, very similar to Q.

goropaniis, but different as follows: size larger, form stouter; chaetotaxy of head similar to that of Q.

goropanus\ however, one additional setiferous puncture anteromediad and occasionally another additional

setiferous puncture posteromediad of posterior frontal puncture; tempora with more numerous punctures

(often almost lacking in Q. gowpaniisY, chaetotaxy of pronotum similar to that of Q. goropanus', however,

sublateral rows with only two punctures, with posterior puncture situated before level of large lateral

puncture; elytra shorter, at suture substantially shorter (ratio 0.70), at sides distinctly shorter (ratio 0.84) than

pronotum at midline; punctation of elytra in general slightly coarser and less dense, that of abdominal

tergites m general slightly sparser.

Male. Stemite 8 (Fig. 73) with only two long and strong setae on each side of apical half, emargination

of its apical margin narrower and slightly less arcuate. Aedoeagus (Figs. 74-77) distinctly larger than that of

Q. goropanus, apical portion of median lobe narrowed, with apical margin subarcuate to broadly arcuate,

internal sac as in Fig. 75. Paramere narrow and elongate, much narrower than median lobe and distinctly not

reaching its apex, with apical margin usually variably emarginate, or occasionally subtruncate to subarcuate;

underside of paramere, as in Q. tanderi, without any sensory peg setae; four setae apically on each side, two

of them usually at apical margin on each side of emargination and two slightly below apex near lateral

Quaest. Ent., 1988, 24 (2)

224

Smetana

margin, or forming a longitudinal row of four setae near apex.

Length 7. 6-9. 5 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL (Prov.

Bagmati) below Thare Pati 3300 m, 1 1.IV.81 Lobl & Smetana”. In the collection A.

Smetana, Ottawa, Canada.

Paratypes (47): Nuwakot Distr.: same data as holotype (ASCC, CNCC, MNHG,

SMFM) 17; between Ghopte and Thare Pati, 3200 m, 23. IV. 85, A. Smetana (ASCC,

BMNH) 5; same, 3150 m, 24.IV.85, A. Smetana (ASCC) 2; same, 3150 m,

25. IV. 85, A. Smetana (ASCC) 4; same, 3200 m, 26. IV. 85, A. Smetana (ASCC,

CNCC) 6; Yangri Ridge, 4350 m, 22.IV.81, Lobl & Smetana (ASCC, MNHG) 7;

same, 4350 m, 22.IV.81, Lobl & Smetana (ASCC, CNCC) 2; same, 4500 m,

23.IV.81, Lobl & Smetana (MNHG) 1. Rasuwa Distr.: north slope above Syabru,

3800 m, 17. IV. 85, A. Smetana (ASCC) 1; same, 3800 m, 28. IV. 85, A. Smetana

(ASCC) 1; same, 3600 m, I9.IV.85, A. Smetana (CNCC) 1.

Geographical distribution. — Quedius angnimai is known from several localities

in central Nepal (map 14).

Bionomics. — The specimens from below Thare Pati were taken by sifting thick

layer of moist moss on large blocks of rock in a mixed forest {Rhododendron, Abies,

Tsuga); most specimens were taken on only two blocks, especially on the eastern

side. Specimens from Yangri Ridge were taken by sifting moss and debris under low

bushes, dead vegetation and dead grass along bases of large rocks on a pasture

above tree line. Specimens from near Syabru were sifted from floor litter and moss

under old Abies trees in a sheltered ravine; those from between Ghopte and Thare

Pati were sifted from wet moss, dead grass and other debris on a seepage slope and

along a creek on clearings in mixed Abies - Rhododendron forest.

Comparisons and variations. — Quedius angnimai can easily be distinguished

from Q. goropanus, in addition to the differences on the aedoeagus, by the presence

of the additional puncture anteromediad of the posterior frontal puncture, and by the

shorter sublateral rows of punctures on the pronotum, in combination with the other

characters mentioned in the description. It also differs by the same set of characters

from Q. tanderi and Q. kailo.

The chaetotaxy of the posterolateral part of the head is somewhat unstable;

however, the puncture situated anteromediad of posterior frontal puncture is always

present and was found doubled unilaterally in one specimen; there are often up to

three additional setiferous punctures between the posterior frontal puncture and the

two usual punctures near posterior margin of head; an additional puncture can also

be found occasionally near the temporal puncture. In one specimen, one of the

sublateral rows on pronotum has three punctures and the last puncture is situated

behind the level of the large lateral puncture.

The specimens from the elevations above 4000 m on Yangri Ridge are in general

somewhat smaller and less robust than the specimens from the lower elevations in

the forest habitats.

Far Western

Revision of the Tribes Quediini and Atanygnathini 225

Quaest. Ent., 1988, 24 (2)

lap 13. Distribution records for: Q.franzi {% ); Q. lesagei t A ); and Q. atchala ( ^ ).

Far Western

226

Smetana

Map 14. Distribution records for: Qiiedius goropanus ( ■ ); tanderi ( A ): Q. kailo ( ^ ); and Q. angnimai (•).

Revision of the Tribes Quediini and Atanygnathini

227

Etymology. — The species was named in honour of my Nepalese friend, Sherpa

Ang Nima, who accompanied me on several of my treks in the Himalaya, and who

substantially contributed to the success of these treks.

2. Subgenus DistichaUiis Casey

Type species. — Staphylinus capucinus Gravenhorst 1806, fixed by Casey

(1915:398).

The subgenus is used here in the same sense as in my revision of the North

American Quediini (Smetana 1971:153). However, see the discussion following

Quedius for the ever increasing difficulties to satisfactorily characterize and delimit

the conventional “classical” subgenera of Quedius.

Key to species of DistichaUiis

1 Elytra evenly punctate, bright metallic blue. Aedoeagus as in

Figs. 79-81. Length 7. 6-8. 4 mm

..22. Q. kasliniirensis Cameron, p. 227

r Elytra with three irregular longitudinal rows of coarse

punctures, dark metallic olive green or dark green 2

2 (T) Head not narrowed posteriorly behind eyes; eyes very large,

tempora about one-third as long as length of eyes seen from

above. Elytra shorter, at suture about as long as pronotum at

midline. Legs with coxae pale. Aedoeagus as in Figs. 83-86.

Length 6. 0-7. 8 mm .23. Q. chatter jeei Cameron, p. 229

2' Head distinctly narrowed posteriorly behind eyes; eyes smaller,

tempora about half as long as length of eyes seen from above.

Elytra longer, at suture longer (ratio 1.23) than pronotum at

midline. Legs with coxae piceous. Length 6. 8-7. 6 mm

24. Q. deceptor Cameron, p. 231

Kashmirensis Group

This group is characterized by the punctation of the elytra, which evenly covers

the entire elytral surface. It might be identical with the Marginalis Group in the

Nearctic region (see Smetana 1971:156).

The group contains one species in the Himalayan region.

22. Quedius (Distichalius) kashmirensis Cameron

Figs. 78-81; Map 15

Quedius kashmirensis Cameron 1944: 13

Quaest. Ent., 1988, 24 (2)

228

Smetana

Quedius mandra Smetana 1975:332 (syn.nov.)

Quedius deuvei Coiffait 1983a: 168 {syn.nov.)

Description. — Entirely black, including appendages, elytra bright metallic blue, pronotum and

abdomen distinctly iridescent. Head rounded, slightly wider than long (ratio 1.2); eyes large and moderately

convex, tempora less than half as long as length of eyes seen from above (ratio 0.41); two additional

setiferous punctures between anterior frontal punctures; posterior frontal puncture situated at posteromedial

margin of eye and separated from it by distance no larger than diameter of puncture, two punctures between

it and posterior margin of head; large temporal puncture situated close to posterior margin of eye, distance

not larger than diameter of puncture; tempora with some fine punctures; surface of head with fine and dense

microsculpture of transverse waves. Antenna moderately long, segment 3 longer than segment 2, segment 4

longer than wide, following segments becoming gradually shorter and wider, outer segments slightly

transverse, last segment about as long as two preceding segments combined. Pronotum broadly arcuate

basally, markedly narrowed in front, hardly wider than long (ratio 1.08); dorsal rows each with three

punctures; sublateral rows each with two or three punctures, posterior puncture situated distinctly behind

level of large lateral puncture; surface of pronotum with microsculpture similar to that on head but slightly

finer and denser. Scutellum impunctate. Elytra long, at suture hardly (ratio 1.08), at sides distinctly longer

(ratio 1.35) than pronotum at midline; punctation moderately coarse and not dense, interspaces between

punctures mostly distinctly larger than diameters of punctures; dark and long pubescence rather scarce;

surface between punctures without microsculpture. Abdomen with tergite 7 (fifth visible) bearing distinct

whitish apical seam of palisade setae; punctation of basal portions of tergites denser and slightly finer than

that on elytra, becoming gradually finer and sparser toward apical margin of each tergite, punctation in

general becoming gradually finer toward apex of abdomen. Eront tarsus dilated in both sexes.

Mcde. Eirst four segments of front tarsus more dilated than in female. Apical margin of stemite 8 with

wide and deep triangular emargination (Eig. 78), triangular area before emargination flattened and smooth.

Aedoeagus (Eigs. 79-81 ) elongate; median lobe almost parallel-sided, rather suddenly narrowed apically and

with two characteristically shaped apical plates. Paramere very narrow and elongate, not quite reaching apex

of median lobe, with arcuate apex; four setae at apex, median pair much longer than lateral pair, a pair of

setae at each lateral margin rather far below apex of paramere; sensory peg setae on underside of paramere

forming irregular long group, gradually becoming denser toward apex of paramere.

Length 7. 6-8. 4 mm.

Type material. — Quedius kashmirensis. The collection of the British Museum

(Natural History), London, contains one female specimen (holotype) under the name

Q. kashmirensis. It is labelled as follows: “Type” (round label with red margin)/

“Kashmir Gulmarg VI-VII-31 Dr. Cameron”/ “Q. kashmirensis Cam. TYPE”/ “M.

Cameron. Bequest. B.M. 1955-147”.

Quedius mandra. The species was described from one female from central Nepal

(see Smetana 1975:333). The holotype is deposited in the Franz collection, Modling,

Austria; it is labelled as follows: “Umg. Coropani w Bokhara”/ “Central Nepal

Sept. -Oct. 1971 Ig. H. Franz”/ “HOLOTYPE Quedius mandra A. Smetana 1975”.

The specimen cannot be distinguished from holotype of Q. kashmirensis', the name

Q. mandra is a junior synonym of Q. kashmirensis. My determination label

“Quedius kashmirensis Cam. Smetana det. 1984” has been attached to it.

Quedius deuvei. Coiffait (l.c.) described the species from a single female from

Nepal. The holotype in the Coiffait collection in the Museum National d’Histoire

Naturelle, Paris, France, is labelled as follows: “Nepal VII. 82 Ganesh Himal

Rupchet 3600 TD”/ “HOLOTYPE”/ “Quedius (Quediops) deuvei H. Coiffait 1983”.

The specimen cannot be specifically distinguished from the holotype of Q.

kashmirensis', the name Q. deuvei is a junior synonym of Q. kashmirensis. My

determination label “Quedius kashmirensis Cam. Smetana det. 1984” has been

Revision of the Tribes Quediini and Atanygnathini

229

attached to this specimen.

Geographical distribution. — Quediiis kashmirensis is widely distributed in the

Himalayan range, from Kashmir to eastern Nepal (Map 15).

Material studied. — 9 specimens.

INDIA. Kashmir. Yusmarg, VI.81, R. de Rougemont (ASCC, GDRC) 2. Uttar Pradesh. Kumaon,

Pindar Valley, 8000-1100', Champion (BMNH) 1; Kumaon, Ranikhet (BMNH) 1; W. Almora Divn.,

Sunderhunga, 8000-12000', VI. 1919, Champion (BMNH) 1.

NEPAL. Nepal or Milke Himal, 3500 m, III.81 (HCCC) 1.

Biononucs. — No details are known about the habitat requirements of this

species.

Comparisons. — Qiiedius kashmirensis can easily be recognized by the presence

of the two additional setiferous punctures between the two anterior frontal punctures

on the head, in combination with the long, bright metallic blue elytra. In general

habitus, it resembles the Nearctic species Q. montivagus Smetana 1971 and also

some species of the Capucinus Group (see Smetana 1971:161), occurring in the

Pacific coastal states in North America, e.g., Q. nevadensis (Casey 1915) or Q.

tetricus Smetana 1971.

Cameron (1944:13), oddly enough, assigned this species to the subgenus

Quedius, although it does not show one single character of that subgenus.

C hatter jeei Group

This group is characterized by the unequal and irregular punctation of the elytra,

which shows a distinct tendency to be arranged in longitudinal rows.

The group contains two species in the Himalayan region.

23. Quedius (Distichalius) chatterjeei Cameron

Figs. 82-86; Map 15

Quedius chatterjeei C&mQmn 1926:370; 1932:295

Description. — Head black, pronotum and abdomen piceous to piceous-black, elytra dark metallic

olive green or dark green, apical margins of abdominal tergites narrowly and paratergites variably paler;

head and pronotum feebly, abdomen distinctly, iridescent; mouthparts dark brown to piceous, antennae

piceous with bases of first two segments paler; legs, including coxae, pale testaceous, tibiae and tarsi dark

brown to piceous. Head of rounded quadrangular shape, wider than long (ratio 1.28), not narrowed behind

eyes, posterior angles rounded; eyes very large, convex, tempora about one-third as long as length of eyes

seen from above (ratio 0.28); two additional setiferous punctures (one occasionally missing) between

anterior frontal punctures; posterior frontal puncture and large temporal puncture both situated very close to

posteromedian margin of eye, sometimes actually touching it, two additional setiferous punctures behind

posterior frontal puncture at posterior margin of head; surface of head with dense and very fine

microsculpture of transverse waves. Antenna moderately long, segment 3 longer than segment 2, segments 4

and 5 longer than wide, segments 6 and 7 about as long as wide, segments 8-10 as long as wide to slightly

transverse. Pronotum slightly wider than long (ratio 1.14), widely arcuate basally and distinctly narrowed

anteriorly; dorsal rows each with three punctures; sublateral rows with two or three punctures, posterior

puncture situated behind level of large lateral puncture; surface of pronotum with microsculpture similar to

that on head. Scutellum without punctures, surface with very fine microsculpture of irregular transverse

waves. Elytra moderately long, at suture about as long as, at sides longer than pronotum at midline (ratio

Quaest. Ent., 1988, 24 (2)

230

Smetana

1.26); each elytron extremely finely and rather sparsely punctate, and with three irregular longitudinal rows

of coarser punctures, some coarser punctures also near posterolateral angles and at posterior margin, all

bearing stiff setae, epipleuron rather densely punctate and pubsecent; surface of each elytron without

appreciable microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish

apical seam of palisade setae; punctation and pubescence of tergites fine, rather dense at base of each tergite

and becoming sparser toward apex of tergite, and in general becoming gradually sparser toward apex of

abdomen. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus much more dilated than in female. Stemite 8 with deep and

rather wide triangular emargination in middle of apical margin (Fig. 82), triangular area before emargination

flattened and smooth. Aedoeagus (Figs. 83-86) quite characteristic; median lobe with two apical, ear-like,

divergent lobes; paramere long and rather wide, slightly dilated, subarcuate apical ly and completely covering

apical portion of median lobe; four setae at apical margin, inner setae close together and longer than outer

setae, a pair of setae at each lateral margin below apex; sensory peg setae on underside of paramere very

numerous, forming two rather wide lateral groups.

Length 6. 0-7. 8 mm.

Type material. — The original series of this species in the British Museum

(Natural History), London, contains 1 1 specimens. They are labelled as follows:

Spec. No. 1 (male): “Type” (round label with red margin)/ “SYN-TYPE” (round

label with blue margin)/ “Woodstock Falls. Mussoorie. Dr. Cameron. 27. III. 1921.”/

“TYPE Quedius chatterjeei Dr. Cameron”/ “M. Cameron. Bequest. B.M.

1955-147.”. Spec. Nos. 2-4: “Mossy Falls Mussoorie.”/ “Dr. Cameron.

20. III. 1921.”/ “M. Cameron. Bequest. B.M. 1955-147”. Spec. No. 5: “Muncpl.

Gardens. Mussoorie 6000'”/ “Dr. Cameron. 15.IV.22”/ “M. Cameron. Bequest. B.M.

1955-147”. Spec. No. 6: “Keyarkuli, Mussoorie.”/ “Dr. Cameron. 17/IV.22”. “Q.

chatterjeei”/ “M. Cameron. Bequest. B.M. 1955-147”. Spec. No. 7: “Dhobi Ghat

Mussoorie.”/ “Dr. Cameron. 14. IV. 22.”/ “M. Cameron. Bequest. 1955-147.” Spec.

Nos. 8 and 9: “Bangar, 6000. Chakrata. U.P.”/ “Dr. Cameron. 1.V.21”/ “Quedius

chatterjeei Cam.” Spec. No. 10: “Chakrata Dist. Jadi Gad 7000'”/ “Dr. Cameron,

9.V.22.”/ “M. Cameron. Bequest. 1955-147.” Spec. No. 11: “Chakrata Dist. Sijla

Gad 5000'”/ “Dr. Cameron. 12.V.22.”/ “M. Cameron. Bequest. B.M. 1955-147.” All

specimens (2-1 1) bear as last a round label with blue margin “SYN-TYPE”.

The first specimen (male) was dissected and the stemite 8, genital segment and

aedoeagus mounted on plate with beetle. The specimen is hereby designated as the

lectotype of Q. chatterjeei; the label “Lectotype Quedius chatterjeei Cameron

Smetana des. 1984” has been attached to it. In addition, male specimens 4 and 10

were dissected, and the aedoeagi were mounted in Canada Balsam.

Geographical distribution. — Quedius chatterjeei is distributed in the western

Himalaya, in the Mussoorie and Chakrata Districts in Uttar Pradesh, and in

Himachal Pradesh (Map 15).

Material studied. — 27 specimens.

INDIA. Himachal Pradesh. Simla, VI.81, R. de Rougemont (GDRC) 1; Jhatingri, Mandi, 6000', H.G.

Champion (BMNH) 1; Dhelu, Mandi, 4500', H.G. Champion (BMNH) 4. Uttar Pradesh. Chakrata Distr.:

Deoban, 1000', V.28, H.G. Champion (BMNH) 3; Jaunsar, VI. 1929, H.G. Champion (BMNH) 1. Mussoorie

Distr.: Mossy Falls, 22.III.32, H.G. Champion (ASCC, BMNH) 4; Dhobi Ghat, 14.IV.22, Cameron (CNCC)

1. Chaubattia, 1900-2100 m, 25.V.81, M. Brancucci (NHMB) 1.

Bionomics. — No details are known about the habitat requirements of this

species. Cameron (1932:296) gives the following habitat data: “Under dead leaves at

Revision of the Tribes Quediini and Atanygnathini

231

the margins of streams”.

Comparisons. — Quediiis chatter jeei apparently is closely related and in all

external characters extremely similar to Q. pretiosus Shaip 1874 from Japan. The

latter species differes, in addition to a few small external differences, by the

differently shaped aedoeagus.

24. Qiiediiis (Distichalius) deceptor Cameron

Map 15

Quediiis deceptor Cameron 1 944: 14

Description . — In all external characters quite similar to Q. chatterjeei but different as follows:

piceous-black, elytra dark metallic green, narrowly pale at apical margin; mouthparts, antennae and legs

piceous. Head distinctly narrowed posteriorly behind eyes, eyes smaller, tempora about half as long as

length of eyes seen from above (ratio 0.46). Antenna feebly longer, with outer segments as long as wide to

feebly longer than wide. Pronotum slightly narrower (ratio width:length = 1.20) and more narrowed

anteriorly. Elytra longer, at suture slightly (ratio 1.23), at sides distinctly (ratio 1.43) longer than pronotum at

midline; fine punctures between rows of coarser punctures coarser and more numerous. Punctation of

abdomen slightly sparser.

Male. Unknown.

Length 6. 8-7. 6 mm.

Type material. — Cameron (l.c.) described the species from a single specimen

from the Darjeeling area. The female holotype in the collection of the British

Museum (Natural History), London, is labelled as follows: “Type” (round label with

red margin)/ “Ghum district V-VI-31 Dr. Cameron”/ “Q. deceptor Cam. TYPE”/

“M. Cameron. Bequest. B.M. 1955-147”.

Geographical distribution. — Quediiis deceptor is at present known from Uttar

Pradesh (Garhwal) and from the type locality in West Bengal, in the Darjeeling area

(Map 15). It is apparently widely distributed in the Himalayan area.

Material studied. — 4 specimens.

INDIA. Uttar Pradesh. Garhwal: between Tehri and Srinagar, 900 m, 25.X.79, I. Lcibl (MHNG) 2;

above Pauri, 1900 m, 28.X.79, I. Ldbl (ASCC) 1. West Bengal. See Type material.

Bionomics. — The specimens from Garhwal were collected by sifting debris

under bushes in a ravine and by sifting leaf litter and moss in a rather dry oak forest.

Comparisons. — Quediiis deceptor can easily be distinguished from Q.

chatterjeei by the characters given in the key and in the description. Although the

male is not known at present, there is hardly any doubt about the specific

distinctiveness of Q. deceptor.

3. Subgenus Raphiriis Stephens

Type species. — Staphylinus attenuatus Gravenhorst 1806, fixed by Westwood

(1838:16).

The subgenus is used here in the same sense as in my revision of the North

American Quediini (Smetana 1971:183). However, see the discussion following

Quediiis for the ever increasing difficulties to satisfactorily characterize and delimit

Quaest. Ent., 1988, 24 (2)

232

Smetana

Map 15. Distribution records for: Q. kashmirensis ( • ); Q. chatterjeei ( A )■, Q. deceptor ( ■ ); and Q. assamensis ( ^ ).

Revision of the Tribes Quediini and Atanygnathini

233

the conventional “classical” subgenera of Quedius.

Taxonomic notes. — Quedius gardnevi is rather isolated within this subgenus,

due mainly to the medially keeled abdominal tergites and to the configuration of the

aedoeagus (see description). It is possible that this species will require the

establishment of a separate subgenus, or even genus, when the fauna of the Quediini

of the Oriental region becomes more adequately known.

Key to species of Raphinis

1 First three abdominal tergites with elevated median keel.

Aedoeagus as in Figs. 233-235. Length 6. 5-6. 8 mm

56. Q. gardneri Cameron, p. 295

r None of abdominal tergites with elevated median keel 2

2 (F) Head, pronotum and elytra bright metallic green or dark green,

bluish green or purplish green. Head with deep and coarse

punctures forming more or less distinct rugae on posterior

portion, or at least with numerous coarse punctures

posteromediad of each eye 3

2' Head and pronotum at most with metallic reflections, elytra

occasionally dark metallic green to greenish blue or metallic

blue. Head with scattered setiferous punctures 5

3 (2) Head with deep and coarse punctures forming more or less

distinct rugae on posterior portion. Dorsal rows on pronotum

formed by very coarse and deep, pit-like punctures 4

3' Head with numerous coarse punctures posteromediad of each

eye, without rugae on posterior portion. Aedoeagus as in Figs.

225-227. Length 6. 8-7. 5 mm 53. Q. hariyo spec.nov., p. 291

4 (3) Scutellum and neck impunctate. Aedoeagus as in Figs.

229-231. Length 5. 8-6. 8 mm... 54. Q. rugosus Cameron, p. 292

4' Scutellum with punctures situated in coarse transverse

depressions: neck with numerous punctures on middle portion.

Male unknown. Length 7.0 mm

55. Q. intricatus Fauvel, p. 295

5 (2’) Two to three additional setiferous punctures on each side of

head along medial margin of eye between anterior and posterior

frontal punctures. Male unknown. Length 6. 1-6.7 mm

45. Q. taruni spec, nov., p. 271

5' No additional setiferous punctures between anterior and

posterior frontal punctures 6

6 (5') Scutellum smooth, without punctures or any other sculpture.

Aedoeagus as in Figs. 88-90. Length 5.6-6.0 mm

25. Q. fluviatilis Cameron, p. 239

Quaest. Ent., 1988, 24 (2)

234

Smetana

6' Scutellum punctate, punctures varying from fine and

inconspicuous to coarse and distinct, and from just a few to

numerous, or scutellum impuncate but with fine transverse rugae 7

7 (6') Scutellum with fine transverse rugae. Surface of head and

pronotum with extremely fine, microscopic punctures.

Aedoeagus as in Figs. 200-202. Length 9.4 mm

47. Q. anomaliis Cameron, p. 274

7' Scutellum punctate. Surface of head and pronotum without

microscopic punctures 8

8 (7') Elytra with double punctation: coarse and rather irregularly

distributed punctures intermixed with distinctly finer punctures.

Aedoeagus as in Figs. 196-198. Length 10.0 mm

46. Q. durgaa spec. now, p. 273

8' Elytra with simple punctation 9

9 (8') Segment 3 of antenna distinctly longer than segment 2. Size

large, form robust. Length 7. 9-9. 6 mm 10

9' Segment 3 of antenna about as long as segment 2. Size small to

moderate, form more or less slender. Length 3. 7-6. 8 mm 14

10 (9) Legs bicoloured: femora pale yellowish, tibiae and tarsi darker.

Pubescence of elytra and abdomen golden-yellowish, forming

slightly denser patches on lateral portions of abdominal tergites.

Aedoeagus as in Figs. 221-223. Length 8. 5-9.0 mm

52. Q. aureipilis Cameron, p. 288

10' Legs more or less unicoloured, dark, femora never pale

yellowish. Pubescence of elytra and abdomen dark, evenly

covering abdominal tergites 1 1

11 (10') Elytra dark metallic green to greenish blue or metallic blue 12

11' Elytra piceous-black to black. Aedoeagus as in Figs. 204-206.

Length 8. 5-9. 6 mm 48. Q. assamensis Cameron, p. 284

12 (11) Male sternite 8 with very deep and wide emargination, fringed

by densely packed dark setae (Fig. 215). Aedoeagus very large,

paramere narrow and elongate, almost parallel-sided in middle

portion (Figs. 216, 217). Length 8. 7-9.0 mm

51. Q. kuiro spec. now, p. 288

12' Male sternite 8 with deep or moderately deep and wide

emargination, without fringe of densely packed dark setae

(Figs. 203, 207, 21 1). Aedoeagi smaller, parameres shorter and

wider, never almost parallel-sided in middle portions (Figs.

204, 208,212) 13

13 (12') Median lobe of aedoeagus in lateral view without a dent in

apical portion (Fig. 209); paramere broadly rounded apically,

shaped as in Fig. 210. Pronotum distinctly more narrowed

Revision of the Tribes Quediini and Atanygnathini

235

anteriorly than posteriorly. Length 8. 4-9. 3 mm

49. Q. himalayicus Bernhauer, p. 285

13' Median lobe of aedoeagus in lateral view with distinct dent in

apical portion! Fig. 213); paramere narrowly rounded apically,

shaped as in Fig. 214. Pronotum about equally narrowed

anteriorly and posteriorly. Length 7. 9-9.0 mm

50. Q. nilo spec. now, p. 287

14 (9') Abdominal tergite 7 (fifth visible) with whitish apical seam of

palisade setae (occasionally delicate and indistinct). Fully

winged species; elytra moderately long, at suture no more than

slightly shorter than pronotum at midline (ratio not below 0.85).

Exception: Q. satoi - see couplet 17 15

14' Abdominal tergite 7 (fifth visible) without whitish apical seam

of palisade setae. Brachypterous species; elytra short, at suture

distinctly shorter than pronotum at midline (ratio around 0.76) 26

15 (14) Pubsescence of abdominal tergites uniform, i.e. without patches

of paler and/or denser hairs on lateral portions. Aedoeagus and

male sternite 8 as in Figs. 91-94. Length 4. 9-5. 6 mm

26. Q. kaalo spec, nov., p. 243

15' Pubescence of abdominal tergites not uniform, lateral portions

of tergites with patches of paler and/or denser hairs.

(Pubescence to be observed on clean specimens from obliquely

behind in light dispersed by an translucent filter. In dirty, oily

specimens the patches cannot be properly seen and the

abdominal pubescence may appear uniform) 16

16 (15') Legs uniformly testaceous 17

16' At least tibiae of hind legs distinctly darkened; however,

usually tibiae of middle legs and femora and tibiae of hind legs

distinctly darkened 21

17 (16) Elytra short, at suture distinctly shorter than pronotum at

midline (ratio about 0.7). Whitish apical seam of palisade setae

on abdominal tergite 7 (fifth visible) delicate and indistinct.

Brachypterous, wing stumps each about reaching apical margin

of elytron. Aedoeagus and male sternite 8 as in Eigs. 151-155.

Length 4.5-6. 3 mm 37. Q. satoi spec. now, p. 260

17' Elytra moderately long, at suture about as long as pronotum at

midline. Whitish apical seam of palisade setae on abdominal

tergite 7 (fifth visible) wide and distinct. Macropterous, wings

each much longer than elytron, folded 1 8

18 (17') Punctation and pubescence of abdominal tergites very sparse,

large middle portion of each tergite (sometimes almost entire

tergite except for lateral patches) glabrous, without any

Quaest. Ent., 1988, 24 (2)

236

Smetana

18'

19 (18)

19'

20 (18')

20'

21 (16')

21'

punctation or pubescence 19

Punctation and pubescence of abdominal tergites gradually

becoming sparser both toward apex of each tergite and toward

apex of abdomen, in general more or less dense and each

tergite, without large middle portion glabrous 20

Front tarsus simple (not dilated) in both sexes. Elytra as dark as

pronotum and head, like them with distinct metallic reflections.

Punctation of elytra sparse, intervals between punctures along

transverse axis mostly several times larger than diameters of

punctures. Aedoeagus and male sternite 8 as in Figs. 141-145.

Length 4. 0^.8 mm 35. Q. simdar spec. now, p. 258

Front tarsus distinctly dilated in male. Elytra more or less paler

than pronotum and head, like them with only slight bronze

metallic reflections. Punctation of elytra more or less dense,

intervals between punctures along transverse axis mostly no

more than twice as large as diameters of punctures. Aedoeagi

and male sternites 8 as in Figs. 122-132. Length 4. 8-6.0

mm 32. Q. muscicola Cameron (pars), p. 253

Median lobe of aedoaegus with lateral outlines

characteristically bisinuate; apex of median lobe subacute

(Figs. 134, 135). Male sternite 8 with very deep and wide apical

emargination (Fig. 133). Length 4.7-5. 1 mm

33. Q. hhari spec. now, p. 256

Median lobe of aedoeagus slightly to distinctly, simply dilated

toward apex, lateral outlines therefore not bisinuate; apex of

median lobe variably rounded (Ligs. 124, 129). Male sternite 8

with moderately deep and wide emargination (Figs. 122, 123).

Length 4. 8-6.0 mm 32. Q. muscicola Cameron (pars), p. 253

First three visible abdominal tergites each with three shallow,

inconspicuous impressions at base; each impression, including

middle one, with slightly denser punctation and pubescence

than on rest of surface. Punctation and pubescence of elytra fine

and dense, interspaces between punctures along transverse axis

about as large as diameters of punctures. Scutellum with

numerous punctures (at least 15). Aedoeagus and male sternite

8 as in Figs. 1 17-121. Length 4.0-5. 2 mm

31. aureiventris Bernhauer, p. 250

First three visible abdominal tergites without shallow, in-

conspicuous impressions at base with denser punctation and

pubescence. Punctation and pubescence of elytra coarser and

distinctly sparser, interspaces between punctures along

transverse axis distinctly larger than diameters of punctures.

Revision of the Tribes Quediini and Atanygnathini

237

22 (21')

22'

23 (22')

23'

24 (23')

24'

25 (24')

25'

Scutellum with less numerous punctures (no more than 12)

Only tibiae of hind legs slightly darkened, middle legs

uniformly testaceous. Pubescence of elytra and abdomen

extensively golden-yellow. Paramere of aedoeagus reaching

apex of median lobe, slightly asymmetrical, with bisinuate

lateral margins (figs. 138, 140). Length 4.8-5.0 mm

34. Q. ekiai spec. now, p. 257

Tibiae of middle legs, tibiae and femora of hind legs and often

also tibiae of front legs distinctly darkened. Pubescence of

elytra and abdomen not extensively golden-yellow, although

yellowish pubescence may be present. Paramere of aedoeagus

not quite reaching apex of median lobe and of different shape

(Figs. 97,98, 104, 108, 113)

Male sternite 8 with three strong setae on each side (Fig. 103).

Apical portion of median lobe, when paramere removed, with

fairly long median carina; paramere of fairly characteristic

shape, with sensory peg setae on underside very numerous,

forming two long and irregular lateral rows (Fig. 106). Length

3.8^. 7 mm 28. Q. gaarho spec. now, p. 246

Male sternite 8 with two strong setae on each side (Figs. 95, 96,

107, 112). Apical portion of median lobe, when paramere

removed, with short median carina; parameres of different

shapes and with sensory peg setae on underside less numerous

(Figs. 101, 102, 111, 116)

Size relatively large, form rather robust. Pubescence of elytra

and abdominal tergites unformly dark, although forming a

patch of denser hairs on each lateral portion of each tergite.

Aedoeagus and paramere as in Figs. 113-116. Length 6. 1-7.2

mm 30. Q. paschim spec, now, p. 249

Size small, form slender. Pubescence of elytra and abdominal

tergites at least partially yellowish, particularly noticeable in

patches of denser hairs on lateral portions of tergites. Aedoeagi

and parameres of different shapes (Figs. 97-102, 108-1 1 1)

Median lobe of aedoeagus with distinct apical tooth in lateral

view (Fig. 110). Paramere elongate and narrow, with sensory

peg setae on underside forming two long rows, with peg setae

gradually becoming widely spaced in direction away from apex

of paramere (Fig. 111). Length 3. 7-4.6 mm

29. Q. daksumensis Coiffait, p. 247

Median lobe of aedoeagus without apical tooth in lateral view

(Fig. 100). Paramere shorter and wider, with sensory peg setae

on underside forming two rather short rows, with sensory peg

22

23

24

25

Quaest. Ent., 1988, 24 (2)

238

Smetana

setae crowded close to each other (Figs. 101, 102). Length

4.0-5. 1 mm 27. Q. vadhu Smetana, p. 244

26 (14') Pubescence of abdominal tergites uniform and dark, /.<?.,

without patches of paler and/or denser hairs on lateral portions.

Male sternite 8 with three strong setae on each side (Fig. 156).

Aedoeagus as in Figs. 157-160. Length 4. 3-5. 8 mm

38. Q. kanyasa Smetana, p. 262

26' Pubescence of abdominal tergites not uniform and dark, lateral

portions of tergites with patches of paler and denser hairs. Male

sternite 8 with two or three strong setae on each side (Figs. 146,

165, 170, 175, 180, 185, 190). Aedoeagi of different shapes

(Figs. 147, 166, 172, 176, 181, 186, 191) 27

27 (26') Legs uniformly testaceous. (Check also Q. naati and Q. pharak) 28

27' At least tibiae of hind legs darkened; however, usually tibiae of

middle legs and femora and tibiae of posterior legs darkened.

Occasionally legs almost uniformly testaceous (some

specimens of Q. naati) 30

28 (27) Paramere of aedoeagus relatively wide anteriorly, covering

apical portion of median lobe except for very apex (Fig. 176).

Flead with anterior portion distinctly paler in male, uniformly

dark in female, usually wider with more convex eyes (ratio

length; width 1.24). Length 4. 0-5. 5 mm

41. Q. tikta Smetana, p. 266

28' Paramere of aedoeagus narrower anteriorly, not covering most

of apical portion of median lobe (Figs. 147, 166, 172, 181, 186,

191). Head usually uniformly dark in both sexes, narrower with

less convex eyes (ratio length; width = 1 . 1 5-1 .20) 29

29 (28') Apical portion of median lobe, when paramere removed, rather

short and wide, and with wide and short median carina;

paramere lancet-shaped, moderately long, with sensory peg

setae forming moderately long rows (Figs. 182, 184). Length

4. 2-5. 8 mm 42. Q. tonglu spec. now, p. 268

29' Apical portion of median lobe, when paramere removed, long

and narrow, and with narrow and long median carina; paramere

narrow and elongate, evenly and gradually narrowed, with rows

of sensory peg setae long (figs. 187, 189). Length 4. 2-5. 8

mm 43. Q. pharak spec. now, p. 269

30 (27') Punctation and pubescence of elytra rather sparse, interspaces

between punctures along transverse axis mostly several times

larger than diameters of punctures. Pronotum narrow, usually

about as long as wide or scarcely longer than wide, not

appreciably narrowed in front. Scutellum with only a few

Revision of the Tribes Quediini and Atanygnathini

239

punctures (range 2-8, usually 3-5). Male sternite 8 with three

strong setae on each side (Fig. 190). Aedoeagus as in Figs.

191-194, paramere rather short, with sensory peg setae fonning

short rows (Fig. 194). Size small: 3. 8-4. 6 mm

44. Q. atchala Smetana, p. 270

30’ Punctation and pubescence of elytra dense, interspaces between

punctures along transverse axis no more than twice as long as

diameters of punctures. Pronotum wider, usually more or less

wider than long and narrowed in front. Scutellum with more

numerous punctures (range 8-16, usually 9-1 1). Male sternite 8

with two strong setae on each side (Figs. 146, 165, 170).

Aedoeagi and parameres of different shapes, rows of sensory

peg setae on parameres longer (Figs. 147, 150, 166, 172, 174) 31

31 (30') Median lobe of aedoeagus not narrowed, apical margin broadly

rounded; paramere short and wide (Figs. 147, 148). Average

size larger: 5. 0-6. 8 mm 36. Q. udagrci Smetana, p. 259

31' Median lobe of aedoeagus narrowed, subacute apically;

parameres elongate and more or less narrow (Figs. 166, 172,

169, 174). Average size smaller: 4. 2-5. 8 mm 32

32 (3F) Median lobe of aedoeagus evenly, almost conically narrowed

(Fig. 172). Average size larger; 4. 9-5. 8 mm

40. Q. dewar spec. now, p. 265

32' Median lobe of aedoeagus slightly dilated before narrowed

apical portion (Fig. 167). Average size smaller; 4. 2-5. 4 mm

39. Q. naati spec. now, p. 264

Fluviatilis Group

The single species of this group is characterized by the following combination of

characters; size moderately large; both head and prontotum without submicroscopic

punctures and without additional setiferous punctures; eyes large but not taking

almost entire sides of head; segment 3 of antenna slightly longer than segment 2;

scutellum impunctate; elytra with simple punctation; first four segments of front

tarsus slightly dilated in female.

25. Quedius (Raphirus) fluviatilis Cameron

Figs. 87-90; Map 16

Quedius fluviatilis Cameron 1926:369; 1932:297

Quedius kashmivicus Cameron 1932:296 (syu.uov.)

Quedius nichinaiensis Coiffait 1982a:79 (syu.uov.)

Quedius cachemiricus Coiffait 19826:277 (syu.uov.)

Quaest. Ent., 1988,24 (2)

240

Smetana

Description. — Dark brown to piceous with black head, elytral suture and apical margin of elytra,

and apical margins of abdominal tergites usually more or less paler, occasionally humeral and outer hind

angles of elytra and/or pronotum paler, testaceorufous to testaceous; palpi, antennae and legs

rufo-testaceous, antennae usually feebly darkened toward apex. Head rounded, feebly wider than long (ratio

1.15); eyes large and convex, tempora much shorter than length of eyes seen from above (ratio 0.39); no

additional setiferous punctures between anterior frontal punctures; posterior frontal puncture very cloase to

posteromedian margin of eye, one puncture between it and posterior margin of head; temporal puncture

almost touching posterior margin of eye; tempora with some fine punctures; surface of head with dense and

very fine microsculpture of transverse waves becoming somewhat confused on clypeus. Antenna moderately

long, segment 3 somewhat longer than segment 2, segments 4 and 5 longer than wide, following segments

gradually becoming shorter, outer segments about as long as wide to feebly wider than long. Pronotum

hardly wider than long (ratio 1.04), broadly rounded basally, slightly narrowed anteriorly; dorsal rows each

with three punctures; sublateral rows each with two punctures, posterior puncture situated before level of

large lateral puncture; surface of pronotum with microsculpture similar to that on vertex of head. Scutellum

impunctate. Elytra moderately long, at suture hardly shorter (ratio 0.95), at sides feebly longer (ratio 1.09)

than pronotum at midline, at base hardly narrower than pronotum at widest point; punctation fine and rather

dense, pubescence brownish, surface between punctures without microsculpture. Wings fully developed.

Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of palisade setae; punctation of

abdominal tergites finer and usually slightly denser than that of elytra, gradually becoming indistinctly

sparser toward abdominal apex; pubescence brownish. Front tarsus dilated in both sexes, slightly so in

female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side of apical half; apical margin with rather deep, subangulate emargination (Fig.

87), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 88-90) narrow and

elongate; median lobe feebly and gradually widened at about apical third and then narrowed toward subacute

apex; apical part of median lobe, when paramere removed, with small subapical hook; internal sac simple;

paramere subparallel-sided, narrower than median lobe and almost reaching apex of median lobe, with four

apical setae and two similar setae at each lateral margin below apex; underside of paramere with two

irregular longitudinal rows of 12 to 13 sensory peg setae.

Length 5. 6-6.0 mm.

Type material. — Quedius fluviatilis. The original series in the British Museum

(Natural History), London, consists of 14 specimens. They are labelled as follows:

Spec. No. 1 (male); “Type” (round label with red margin )/“SYN-TYPE“ (round

label with blue margin)/ “Chakrata Dist. Dodora Khud 8000'”/ “ Dr. Cameron.

13.V.22” / “TYPE Quedius fluviatilis Dr. Cameron/ M. Cameron. Bequest. B.M.

1955-147.” Spec. Nos. 2-8: “Chakrata Dist. Dadora Khud 8000'” / “Dr. Cameron.

6.V.22 (or) 13.V.22.”/ “M. Cameron. Bequest. B.M. 1955-147.” Spec. Nos. 9 and

10: “Chakrata Dist. Binal Gad 7000-8000'”/ “Dr. Cameron. 28.V.22.”/ “M.

Cameron. Bequest. B.M. 1955-147.” Spec. No. 11: “Chakrata Dist. Bodyar 8300'”/

“Dr. Cameron. 3-12.V.22.”/ “M. Cameron, bequest. B.M. 1955-147.” Spec. No. 12:

“Chakrata Dist. Patra Khud 8000'” / “Dr. Cameron. 15.V.22.”/ “Quedius fluviatilis

Cam.” Spec. No. 13: “Chakrata Dist. Manjgaon 6500'“ / “Dr. Cameron. 18.V.22.”/

“M. Cameron. Bequest. B.M. 1955-147.” Spec. No. 14: “Deoban, 9331' Chakrata,

U.P.” / “Dr. Cameron. 3.V.21”/ “M. Cameron. Bequest. B.M. 1955-147.”

Specimens 2-14 each bear as last label a round label “SYN-TYPE” with blue

margin.

The first specimen (male) was dissected and the stemite 8, the genital segment

and the aedoeagus were mounted on plate with beetle. The specimen is hereby

designated as the lectotype of Q. fluviatilis', the label “Eectotype Quedius fluviatilis

Revision of the Tribes Quediini and Atanygnathini

241

Cameron Smetana des. 1984” has been attached to it.

Quediiis kashmiricus. The original series in the British Museum (Natural

History), London, consists of two specimens. They are labelled as follows: Spec.

No. 1 (female): “TYPE” (round label with red margin)/ “SYN-TYPE” (round label

with blue margin)/ “Kashmir Gulmarg VI-VII-31 Dr. Cameron”/ “Q. kashmiricus

Cam. Type” / “M. Cameron. Bequest. B.M. 1959-147.” Spec. No. 2 (male):

“Kashmir Gulmarg VI-VII-31 Dr. Cameron”/ “M. Cameron. Bequest. B.M.

1959-147”/ “SYN-TYPE” (round label with blue margin).

The male specimen was dissected and the sternite 8, the genital segment and the

aedoeagus were mounted on plate with beetle. The specimen is hereby designated as

the lectotype of Q. kashmiricus-, the label “Lectotype Quedius kashmiricus Cameron

Smetana des. 1984” has been attached to it.

Neither of the two specimens can be distinguished from the lectotype of Q.

fliiviatilis. The name Q. kashmiricus is a junior synonym of Q.fluviatilis.

Quedius nichinaiensis. Coiffait (1982a:79) described the species from six

specimens from Nichinai Valley in Kashmir. I was able to study five specimens

deposited in the Senckenberg Museum, Frankfurt a.M., Federal Republic of

Germany. The male holotype is labelled as follows: “INDIA: Kashmir: Sonamarg

Himalaya-Hauptkette Nichinai-Tal, 9.VI.1976 3200-3100 m Martens & Schawaller

leg.”/ “TYPE”/ “Holo-typus” SMF 15105 (underside)/ “Quedius (Sauridus)

nichinaiensis H. Coiffait 1979”. The four paratypes (2 dd, 2??) have same locality

labels as the holotype (except “3100-3200”) and two red labels “PARATYPE”/

“Para-typus” SMF 15105 (underside).

The holotype was damaged when shipped to me. The entire abdomen was lost

and also the aedoeagus was gone from the plate, except for the basal bulbus trapped

in the glue on the plate. I managed to recover the paramere which is now glued on

the plate. Both male paratypes were dissected, the sternite 8 and the aedoeagus of

one male were mounted in Canada Balsam, and the aedoeagus of the second male

was glued to plate with beetle.

None of the five specimens can be distinguished from the lectotype of Q.

fluviatilis. The name Q. nichinaieinsis is a Junior synonym of Q. fliiviatilis. All

specimens were accordingly labelled.

Quedius cachemiricus. Coiffait (1982b:277) described the species from a single

female from Sonamarg in Kashmir. The holotype is deposited in the

Naturhistorisches Museum in Basel, Switzerland, and is labelled as follows:

“Sonamarg 17.7. 2600-2750 m”/ “Kashmir, 1976 W. Wittmer”/ “TYPE” (red label)/

“Quedius (Sauridus) cachemiricus H. Coiffait 1982”.

The holotype of Q. cachemiricus comes from the same locality as Q.

nichinaiensis-, it cannot be distinguished in any way from the lectotype of Q.

fluviatilis-, the name is a Junior synonym of Q. fluviatilis. My determination label

“Quedius fluviatilis Cam. Smetana det. 1984” has been attached to this specimen.

Quaest. Ent., 1988, 24 (2)

242

Smetana

Map 16. Distribution records for: Quedius fluviatilis )\Q. kaalo { A ); and Q. tonglu ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

243

Geographical distribution. — Qiiedius fluviatilis is distributed in the western

portion of the Himalayan range, in Kashmir, Himachal Pradesh and in northern

Uttar Pradesh (Map 16).

Material studied. — 54 specimens.

INDIA. Kashmir. Gulmarg, VI-VII-31, Cameron (BMNH) I; Yusmarg, VI-81, de Rougemont (ASCC,

GDRC) 2; Aru, X.77, H. Franz (ASCC, HFCC) 3. Himachal Pradesh. Mandi, Jhatingri, 6000', Champion

(BMNH) 5; Dharmsala, Champion (BMNH) I. Uttar Pradesh. Chakrata Distr. Mandali, 9100’ (BMNH)2;

Chutli Khud, 8000', I7.V.22, Cameron (BMNH)6; Korawa Khud, 9100’, 4.V.22, Cameron (ASCC, BMNH)

7; Deoban, 9331’, Cameron (BMNH) 3; Konain, 7800’, Cameron (BMNH) 1. Garhwal; between Tehri and

Srinagar, 900 m, 25.X.79, 1. Lbbl (MHNG) 1.

Bionomics. — Little is known about the habitat requirements of this species.

Cameron (1932:197) gives “the borders of streams” as the habitat. The specimens

from Garhwal were taken by sifting debris under bushes in a ravine.

Comparisons. — Quedius fluviatilis is rather variable in the colouration of the

body; however, it is easily recognized by the general habitus alone. It is the only

Himalayan species of Raphirus with impunctate scutellum resembling in the general

habitus the west-palaearctic species of the Q. nemoralis Baudi - Q. limhatus Heer

relationship. The only other Himalayan species of Raphirus with impunctate

scutellum [Q. taruni) differs from Q. fluviatilis considerably by the different

chaetotaxy of the head and pronotum, etc. (see under Q. taruni for details).

Muscicola Group

This species group is characterized by the following combination of characters:

size small; both head and pronotum without microscopic punctures and without

additional setiferous punctures; eyes very large, taking up almost entire sides of

head; segment 3 of antenna about as long as segment 2; scutellum punctate; elytra

with simple punctation; first four segments of front tarsus no more than moderately

dilated in female.

The group contains 19 species in the Himalayan region.

26. Quedius (Raphirus) kaalo spec.nov.

Figs. 91-94; Map 16

Description. — Piceous-black to black, head, pronotum and elytra with metallic reflections,

abdomen iridescent; palpi and antennae testaceous, legs testaceo-brunneous with indistinctly paler tarsi, hind

femora darkened except apically, hind tibiae more or less darkened medially. Head rounded, slightly wider

than long (ratio 1.20), eyes very large and convex, tempora quite short, considerably shorter than length of

eyes seen from above (ratio 0.18); no additional setiferous punctures between anterior frontal punctures;

posterior frontal puncture touching posteromedian margin of eye, one puncture between it and posterior

margin of head; temporal puncture touching posterior margin of eye; surface of head with dense and rather

coarse microsculpture of irregular transverse waves becoming slightly confused anteriorly and meshed in

middle of clypeus. Antenna moderately long, segment 3 slightly longer than segment 2, segments 4 and 5

distinctly longer than wide, segments 6 and 7 slightly longer than wide, segments 8-10 about as long as

wide, last segment about as long as two preceding segments combined. Pronotum about as long as wide,

widely arcuate basally, evenly transversely convex, moderately narrowed anteriorly; dorsal rows each with

Quaest. Ent., 1988, 24 (2)

244

Smetana

three punctures; sublateral rows each with two punctures, posterior puncture situated before level of large

lateral puncture; surface of pronotum with rnicrosculpture similar to that on head but slightly finer.

Scutellum with 1 1-14 punctures. Elytra moderately long, at base slightly narrower than pronotum at widest

point, at suture slightly shorter (ratio 0.85), at sides about as long as pronotum at midline; punctation dense,

intervals between punctures along transverse axis mostly somewhat larger than diameters of punctures;

pubescence dark; surface between punctures without rnicrosculpture. Wings fully developed. Abdomen with

tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; punctation of tergites slightly finer than

that on elytra and somewhat denser on front tergites but gradually becoming sparser toward apex of

abdomen; pubescence dark, evenly covering surface of tergites. Front tarsus dilated in both sexes,

inconspicuously so in female.

Male. First four segments of front tarsus distinctly more dilated than in female. Sternite 8 with two long

and strong setae on each side; apical margin with moderately wide and deep, obtusely triangular

emargination (Fig. 91), small triangular area before emargination flattened and smooth. Aedoeagus (Figs.

92-94) with median lobe subparallel-sided in middle portion and then narrowed into obtuse apex. Paramere

not quite reaching apex of median lobe, more or less parallel-sided, apically narrowed into obtuse apex; four

apical setae, both median setae longer than lateral setae; two setae at each lateral margin close to apex;

sensory peg setae on underside of paramere numerous, forming two irregular, rather long longitudinal rows.

Length 4. 9-5. 6 mm.

Type material. — Holotype (male): “NEPAL (Prov. Bagmati) Malemchi, 2800

m, 14.IV.81, Lobl & Smetana 24 A”. Allotype (female); “NEPAL (Prov. Bagmati)

Dobate Ridge NE Barahbise, 2700 m, 2.V.81 Lobl & Smetana”. Both holotype and

allotype in the Smetana collection, Ottawa, Canada.

Paratypes (5): NEPAL: same data as holotype (ASCC) 1 same data as allotype

but date 7.V.81 (MHNG) 1 c^. INDIA; Uttar Pradesh: Kumaon: W. Almora, H.C.

Champion (BMNH)l; W. Almora Divn., X.1918, H.G. Champion (BMNH) 2.

Geographical distribution. — Quedius kaalo is at present known from Uttar

Pradesh and from two localities in central and east-central Nepal (Map 16); it is

possibly widely distributed in the western portion of the Himalaya.

Bionomics. — The specimens from Malemchi were taken by sifting moss, leaf

litter and other debris under bushes along a stone wall between fields. The

specimens from Dobatge Ridge were taken by sifting moist to wet leaf litter and

other debris in a semideciduous forest.

Comparisons. — Quedius kaalo is well characterized among the small species of

Raphirus with punctate scutellum and partially darkened legs by the following

combination of characters, in addition to the shape of the aedoeagus: size

comparatively large, elytra moderately long, wings fully developed, abdomen with

whitish apical seam of palisade setae on tergite 7, pubescence of tergites evenly

distributed, not forming lateral patches of denser hairs.

Quedius kaalo resembles in general appearance the Holarctic species Q.

fulvicollis Stephens, or the Nearctic species Q.frigidus Smetana.

Etymology. — The specific name is the Nepali adjective kaalo (black); it refers to

the general colour of this species.

Revision of the Tribes Quediini and Atanygnathini

245

27. Quediiis (Raphirus) vcidlm Smetana

Figs. 95-102; Map 17

Quediiis vadhii Smetana 1975:334

Description. — Piceous-black to black, elytra and apical portions of abdominal tergites occasionally

slightly paler; head, pronotum and elytra with slight metallic lustre, abdomen slightly iridescent. Palpi and

antennae testaceous, legs testaceo-brunneous, front tibiae slightly, middle and hind tibiae distinctly darkened

at inner margin, posterior femora usually variably darkened. Head rounded, feebly wider than long (ratio

1.13); eyes very large and convex, tempora extremely short, considerably shorter than length of eyes seen

from above (ratio 0.21); no additional setiferous punctures between anterior frontal punctures; posterior

frontal puncture situated at posteromedian margin of eyes, one puncture between it and posterior margin of

head; surface of head with fine and not dense microsculpture of irregular transverse waves becoming

confused on clypeus. Antenna moderately long, segment 3 slightly narrower and about as long as segment 2,

segments 4-6 longer than wide, gradually becoming shorter, outer segments about as long as wide, last

segment about as long as two preceding segments combined. Pronotum about as long as wide to hardly

longer than wide (ratio 1.05), very broadly arcuate basally, evenly convex and slightly, but distinctly,

narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with two, or rarely three

punctures, posterior puncture not or rarely (when three punctures present) about reaching level of large

lateral puncture; surface of pronotum with microsculpture similar to that on head but usually slightly denser.

Scutellum punctate, punctures moderately numerous, usually no more than 12 in number. Elytra moderately

long, at suture as long as, at sides somewhat longer (ratio 1.12) than pronotum at midline; punctation

moderately coarse and not dense, interspaces between punctures along transverse axis distinctly larger than

diameters of punctures; surface between punctures without microsculpture; yellowish pubescence rather long

and sparse. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of

palisade setae; punctation of tergites distinctly finer than that on elytra, denser on basal portions of tergites

and in general becoming sparser toward apex of abdomen; first three tergites without shallow impressions;

yellowish pubescence forming inconspicuous, not quite dense, patch of denser hairs on each lateral portion

of each tergite. Front tarsus slightly dilated in both sexes, inconspicuously so in female.

Male. First four segments of front tarsus slightly more dilated than in female. Stemite 8 with two long

and strong setae on each side, apical margin with moderately deep, obtusely triangular emargination (Figs.

95, 96), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 97-102) small;

median lobe rather markedly and suddenly narrowed, apex subacute, apical portion, when paramere

removed, with fine, arcuate transverse ridge which is sometimes narrowed into structure shaped as broad V,

without apical dent in lateral view. Paramere rather short, not quite reaching apex of median lobe,

subparallel-sided in middle part, apical portion narrowed, apex broadly obtuse; with four setae at apical

margin, median setae longer than lateral setae, and with a pair of setae at each lateral margin near apex;

sensory peg setae on underside of paramere forming two short irregular rows mediad of lateral margins with

peg setae crowded close to each other.

Fength 4.0-5. 1 mm.

Type material. — The species was described from five specimens. The male

holotype and female allotype bear the following labels; “Kali-Gandaki-Tal zw. Lete

u. Tukche’V “Central-Nepal Sept.-Okt. 1971 Ig. H. Franz”, the underside of the first

label bears code “Pa 132+133”; the three female paratypes come from the same

locality as the holotype and from near Goropani west of Pokhara. The holotype,

allotype and one paratype are deposited in the Franz collection, Modling, Austria,

two paratypes are deposited in the Canadian National Collection, Ottawa (CNC No.

13968). The holotype bears red label “HOLOTYPE Quedius vadhu A. Smetana

1975”

Geographical distribution. — Quedius vadhu is known only from western and

central Nepal, both from the main Himalayan range, and from the mountain

Phulcoki in the Mahabharat Lekh range at the southern edge of the Kathmandu

Quaest. Ent., 1988, 24 (2)

246

Smetana

valley (Map 17).

Material studied. — 5 1 specimens.

NEPAL. Lalitpiir Distr. Phulcoki, 2600-2650 m, 21-22.111.80, Martens & Ausobsky (SBMF) 1;

Phulcoki, 20-22.1 V. 82, 2550-2600 m, A. & Z. Smetana (ASCC, BMNH, CNCC) 12; Phulcoki, 2600 m,

14.X.83, Smetana & Lobl (ASCC, MHNG) 6: Phulcoki, 2550-2700 m, 28-29.IV.84, Smetana & Lobl;

(ASCC) 2. Manaiig Distr. Forest W Bagarchhap, 2200-2250 m, 21 and 22. IX. 83, Smetana & Lobl (ASCC,

BMNH, MHNG) 13; Latha Manang W Bagarchhap, 22. and 23.IX.83, Smetana & Lobl (ASCC, MHNG) 7.

Mustang Distr. Thaksang, 3150-3400 m, 26. 29. IV. 80, Martens & Ausobsky (SBMF) 1. Nuwakot Distr.

Malemchi, 2800 m, I6.IV.81, Lobl & Smetana (MHNG) 1. Dobate Ridge. NE Barahbise, 2800 m, 3.V.81,

Lobl & Smetana (ASCC) 3.

Bionomics. — Quedius vadhii occurs mainly at elevations between 2500-3000 m,

mostly in forest. Most specimens were collected by sifting moist leaf litter, moss

and various debris, sometimes in rather wet habitats. Some specimens of the original

series were also found under loose bark on Finns excelsa trees.

Comparisons and variations. — Quedius vadhu and the two following species

can rather easily be recognized by the following combination of characters: size

relatively small; head, pronotum and elytra with metallic lustre; punctation of elytra

not dense; abdomen with tergite 7 with distinct whitish apical seam of palisade

setae, basal abdominal tergites without impressions, bearing dense punctation and

pubescence at bases, hairs of lateral patches of yellowish pubescence rather sparse

and inconspicuous; and tibiae distinctly darkened.

The configuration of the apical portion of median lobe adjacent to the paramere

is somewhat unstable in Q. vadhu. In typical specimens this part bears a fine

transverse ridge; however, in some populations {e.g., from Phulcoki) this ridge

becomes narrower and forms a structure shaped as a broad V; in the latter case, there

may be a trace of a hook in lateral view. Despite this variability, the combination of

the described configuration of the median lobe with the rather short paramere with

the rows of sensory peg setae short and with the peg setae crowded close to each

other, makes the distinguishing of the males of Q. vadhu from those of Q.

daksumensis and Q. gaarho relatively easy. See under the two latter species for

further discussion of the differences between these three similar species.

The five specimens of the original series of Q. vadhu were in poor condition.

They all had dirty and greasy abdomens; the patches of denser pubescence on lateral

portions of abdominal tergites were therefore obscured and were not mentioned in

the original description. With the discovery of Q. gaarho at Ghoropani Pass, it is

very likely that the female paratype of Q. vadhu from “near Goropani, W. Pokhara,

27. IX. 1971” (Smetana 1975;335) in fact belongs to Q. gaarho.

28. Quedius (Raphirus) gaarho spec.nov.

Figs. 103-106: Map 21

Description. — in all characters, including general habitus and colouration, very similar to Q. vadhu

and different by same set of small external differences as Q. daksumensis, and by characters on aedoeagus.

Male. First four segments of front tarsus more dilated than in female. Stemite 8 (Fig. 103) with three

long and strong setae on each side of apical half, emargination in middle of apical margin shallower than that

Revision of the Tribes Quediini and Atanygnathini

247

of Q. vadint, but slightly deeper than in most specimens of Q. dakswnensis (Fig. 107). Aedoeagus (Figs.

104-106) rather narrow, median lobe narrow and more or less evenly narrowed toward subacute apex, apical

portion, when paramere removed, with fairly long median carina appearing as distinct apical dent in lateral

view; paramere relatively wide, not quite reaching apex of median lobe, with same set of setae as Q. vadhu

and Q. daksiime?isis; sensory peg setae on underside very numerours, forming two long and irregular lateral

rows, with peg setae about equally spaced throughout.

Length 3. 8^. 7 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Parbat Distr.

Ridge E Ghoropani Pass 3150 m 7.X.1983 Smetana & Lobl”. Both holotype and

allotype deposited in the Smetana collection, Ottawa, Canada.

Paratypes (14): NEPAL: same data as holotype (ASCC, CNCC) 5; same data as

holotype but elevation 3100 m (ASCC, BMNH, MHNG) 5; Parbat Distr.: Ghoropani

Pass N slope, 2700 m, 6.X.1983, Smetana & Lobl (MHNG) 1; Pun Hill at

Ghoropani Pass, 3050-3100 m, 8. X. 1983, Smetana & Lobl (MHNG) 1; betw. Chitre

and Ghandrung, Chitre-side of the Pass, 2800-2900 m, 4-7. V. 80, Martens &

Ausobsky (SBML) 1. Mustang Distr.: Thaksang, 3150 m, 26-29. IV. 80, Martens &

Ausobsky (SBML) 1 .

Geographical distribution. — Quedius gaarho is known at present from the

Parbat District in Nepal, particularly from the Ghoropani Pass and its vicinity (Map

21).

Bionomics. — Quedius gaarho was collected at elevations between 2700-3150

m, mostly in mixed Rhododendron-Ahies-Acer or Ahies-Pinus excelsa forest.

Specimens were taken by sifting various mushrooms both on ground and on fallen

trees, and by sifting moss and various debris on forest floor.

Comparisons. — Quedius gaarho is very similar in all external characters to both

Q. vadhu and Q. daksumensis\ on the other hand, it can be distinguished from both

of them by the distinctly different aedoeagus (see Pigs. 97, 98, 104, 108, etc.) and by

the presence of three strong setae on each side of apical half of male stemite 8 (Pig.

103).

Etymology. — The specific name is the Nepali adjective gaarho (difficult). It

refers to the fact that the species is difficult to distinguish from the similar related

species.

29. Quedius (Raphirus) daksumensis Coiffait

Pigs. 107-1 1 1; Map 18

Quedius daksumensis Coiffait 1982b;279

Description. — In all characters, including general habitus and colouration, very similar to Q. vadhu

and different only by a few small external differences and by characters on aedoeagus. Size in general

slightly smaller, head and pronotum less voluminous and narrower, antennae in general slightly shorter and

elytra often paler, rather dark brownish piceous to occasionally brown.

Male. First four segments of front tarsus slightly more dilated than in female. Stemite 8 (Fig. 107) with

two long and strong setae on each side, emargination in middle of apical margin shallower than that of Q.

vadhu (Figs. 95, 96). Aedoeagus (Figs. 108-1 1 1 ) very similar to that of Q. vadhu and about equally long;

however, apical portion of median lobe, when paramere removed, with short carina appearing as distinct

apical dent in lateral view; paramere more elongate and narrower, with same set of setae apically as in Q.

Quaest. Ent., 1988, 24 (2)

248

Smetana

vadhu\ however, with sensory peg setae on underside forming two long rows, with peg setae gradually

becoming widely spaced in direction away from apex of paramere.

Length 3. 7-4. 6 mm.

Type material. — Coiffait (1982b:279) described the species from a single

specimen from Daksum in Kashmir. The male holotype is deposited in the

Naturhistorisches Museum in Basel, Switzerland. It is labelled as follows: “Daksum

9.-13.7.2400-2700 m’V “Kashmir 1976 W. Wittmer’V “TYPE” (red label)/“Q.

(Raphirus) daksumensis H. Coiffait 1982”.

Geographical distribution. — Quedius daksumensis is distributed from Kashmir

eastwards through central Nepal to about the Arun River Valley in eastern Nepal

(Map 18).

Material studied. — 171 specimens.

INDIA. Kashmir. See Type material.

NEPAL. Thodung-Those, 3200 m, 3-9. IV. 73, J. Martens (SBMF) 1; Ting Sang La above Barahbise,

3400 m, 13- 15. IV. 73, J. Martens (SBMF) 1; Dobate Ridge NE Barahbise, 2800 m, 2.V.81, Lobl & Smetana

(MHNG) 1; Jangtang Ridge NE Barahbise, 3250 m, 5.V.81, Lobl & Smetana (ASCC, MHNG) 6; Mere

Dara, 3200 m, 8.IV.81, Lobl & Smetana (MHNG) 1; below Thare Pati, 3300-3500 m, 10-13.IV. 81, Lobl &

Smetana (ASCC, BMNH, CNCC, MHNG) 1 18; Malemchi, 2800 m, 16.IV.81, Lobl & Smetana (ASCC) 1;

above Shermantang, 2900 m, 26.IV.81, Lobl & Smetana (ASCC) 5; Khandbari Distr. “Bakan” W of

Tashigaon, 3200-3250 m, 4-5. IV. 82, A. & Z. Smetana (ASCC) 5; above Tashigaon, 3500 m, 6. IV. 82, A. &

Z. Smetana (ASCC) 1. Nuwakot Distr. betw. Ghopte and Thare Pati, 3100-3250, 23. -26. IV. 85, A. Smetana

(ASCC, BMNH, CNCC) 17. Rasuwa Distr. Lantang Khola Valley, 2.5 km E Syabru, 1730 m, 14.IV.85, A.

Smetana (ASCC, MHNG) 13.

Bionomics. — Quedius daksumensis lives in similar habitats as Q. vadhip

however, it seems to prefer slightly higher elevations. The holotype was collected at

an elevation between 2400-2700 m; however, all other specimens studied were

taken in forest habitats close to 3000 m and up to 3500 m. Most specimens from

below Thare Pati were taken by sifting moist thick moss on large rocks in a mixed

forest (Tsuga, Abies, Rhododendron), and in a ravine with Acer growth, by sifting

moist moss at bases of large rocks on a small clearing in a forest, and by sifting

moss on large fallen trees; specimens from above Shermantang were taken by sifting

thick layers of lichen mixed with moss on large rocks. The species obviously prefers

moist moss as a habitat; however, occasionally specimens were also collected by

sifting layers of old needles and other forest floor debris under coniferous trees.

Comparisons. — Quedius daksumensis is very similar to Q. vadhw, however, it

usually can be distinguished without great difficulties by the characters mentioned

in the key and in the description, particularly by the differently shaped median lobe

and paramere of the aedoeagus. In addition, the two species seem to prefer different

habitats (see above and under Q. vadhu) and were never observed together in the

same habitat.

Quedius daksumensis is also extremely similar to Q. gaarho and can be

positively distinguished from it only by the differences on the median lobe and

paramere of the aedoeagus (see Figs. 108-11, 104-106).

Quedius daksumensis is apparently the smallest species of the genus in the

Himalayan area, followed by Q. gaarho and Q. atchala.

Revision of the Tribes Quediini and Atanygnathini

249

The two specimens from “Thodung-Those” and “Ting Sang La” were published

by Coiffait (1982a:33) under the name Q. vadhu.

30. Quediiis (Raphirus) paschim spec. now

Figs. 112-116; Map 18

Description. — Piceous-black to black, head, pronotum and elytra with slight bronze metallic

reflections, abdomen iridescent; both labial and maxillary palpi, and antennae testaceous; legs testaceous to

brunneo-testaceous, middle tibiae slightly (often indistinctly) and hind tibiae distinctly darkened medially,

hind femora darkened except for pale apex. Head rounded, slightly wider than long (ratio 1.2); relatively

small, narrower than pronotum at widest point (ratio 0.81); eyes large and convex, tempora very short,

considerably shorter than length of eyes seen from above (ratio 0.16); no additional setiferous punctures

between anterior frontal punctures; posterior frontal puncture almost touching posteromedian margin of eye,

one puncture between it and posterior margin of head, temporal puncture touching posterior margin of eye;

surface of head with dense and fine microsculpture of transverse and oblique waves with numerous

longitudinal junctions, gradually becoming almost meshed on middle portion on clypeus. Antenna

moderately long, segments 2 and 3 about equally long, segments 4-6 longer than wide, gradually becoming

distinctly shorter and wider, segments 8-10 about as long as wide, segment 11 as long as two preceding

segments combined. Pronotum about as long as wide, widely rounded basal ly and distinctly narrowed

anteriorly, evenly transversely convex; dorsal rows each with three punctures; sublateral rows with two

punctures, posterior puncture situated before level of large lateral puncture, surface of pronotum with

microsculpture finer and denser than that on head, composed of iiregular transverse waves. Scutellum with

numerous punctures (13-17). Elytra moderately long, at base slightly narrower than pronotum at widest

point, at suture feebly shorter (ratio 0.90), at sides feebly longer (ratio 1.09) than pronotum at midline;

punctation fine and dense, intervals between punctures along transverse axis no more than slightly larger

than diameters of punctures; pubescence moderately dense, dark; surface between punctures without

microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of

palisade setae; punctation of tergites fine and dense, in general denser than that on elytra, gradually

becoming sparser toward apical margin of each tergite and in general becoming sparser toward apex of

abdomen; pubescence dark, dense, forming a patch of denser hairs on lateral portions of each tergite. Front

tarsus dilated in both sexes, inconspicuously so in female.

Male First four segments of front tarsus distinctly more dilated than in female. Sternite 8 with two long

and strong setae on each side; apical margin with moderately wide and deep, obtusely triangular

emargination (Fig. 112), small triangular area before emargination flattened and smooth. Aedoeagus (Figs.

113-116) moderately large, median lobe slightly naiTowed with apex obtusely arcuate; paramere elongate

and narrow, not quite reaching apex of median lobe; four apical setae, median pair longer than lateral setae;

two setae at each lateral margin below apex; sensory peg setae on underside of paramere forming two

irregular and relatively short longitudinal lateral rows.

Fength 6. 1-7.2 mm.

Type material. — Holotype (male) and allotype (female): “INDE Garhwal (UP)

au-dessus Pauri 1900 m 28.X.79 I. Lobl”. Both holotype and allotype deposited in

the Museum d’Histoire Naturelle, Geneve, Switzerland.

Paratypes (17): INDIA: same data as holotype (ASCC, BMNH, CNCC, MHNG)

10; Uttar Pradesh: Garhwal, Mussoorie Range: 2 km E Dhanolti, 2250 m, 21.X.79,

I. Lohl (MHNG) 1; above Joshimath, 2100 m, 27.X.79, I. Ldbl (MHNG) 1.

Kumaon: Chaubattia, cca 1800 m, 12-13.X.79, I. Lobl (MHNG) 1; Chaubattia nr.

Ranikhet, cca 1800 m, 12-13.X.79, I. Lobl (MHNG) 3. NEPAL: Dobate Ridge NE

Barahbise, 2800 m, 2.V.81, Lobl & Smetana (MHNG) 1.

Geographical distribution. — Quedius paschim is at present known from several

localities in Uttar Pradesh and from one locality in east-central Nepal (Map 18); it

Quaest. Ent., 1988, 24 (2)

250

Smetana

apparently belongs to those species distributed mainly in the western portion of the

Himalayan range.

Bionomics. — Qiiedius paschim was collected at elevations of up to 2800 m by

sifting leaf litter and moss in a rather dry oak forest; by sifting leaf litter and moss

near a seepage in a mixed Rhododendron-Ahies forest; by sifting leaf litter in a

Rhododendron forest, and by sifting floor litter and debris among ferns in a forest.

Comparisons. — Quediiis paschim resembles in general habitus Q. aureiventris;

however, it differs, in addition to the characters on the aedoeagus, by the distinctly

larger size and more robust form, by the uniformly dark pubescence of the elytra

and abdomen, and by the absence of the shallow, inconspicuous impressions at the

base of each of the firest three visible abdominal tergites.

Etymology. — The specific name is the Nepali noun paschim (west). It refers to

the distributaional range of the species.

31. Quedius (Raphirus) aureiventris Bernhauer

Figs. 117-121; Map 19

Quedius aureiventris Bernhauer 1915:56 Cameron 1932:294

Quedius decipiens Cameron 1944: 14 {syn.nov.)

Description. — Piceous-black to black, elytra usually more or less paler, dark brunneous to

brunneo-piceous; head, pronotum and elytra with dark metallic reflections, abdomen iridescent; both labial

and maxillary palpi testaceous, both occasionally indistinctly darkened toward apex; antennae testaceous;

legs testaceous to brunneo-testaceous, front tibiae occasionally slightly darkened medially, middle and hind

tibiae and usually also hind femora darkened. Head rounded, slightly wider than long (ratio 1.27); eyes large

and convex, tempora very short, considerably shorter than length of eyes seen from above (ratio 0.22); no

additional setiferous punctures between anterior frontal punctures; posterior frontal puncture almost touching

posteromedian margin of eye, one puncture between it and posterior margin of head; temporal puncture

touching posterior margin of eye; surface of head with dense and very fine microsculpture of transverse

waves becoming irregular on clypeus. Antenna moderately long, segments 2 and 3 about equally long,

segments 4-6 longer than wide, gradually becoming shorter, segment 7 slightly longer than wide, segments

8-10 about as long as wide, segments 9-10 occasionally feebly transverse, segment 1 1 about as long as two

preceding segments combined. Pronotum about as long as wide, widely rounded basally and distinctly

narrowed anteriorly, evenly transversely convex; dorsal rows each with three punctures; sublateral rows each

with two punctures, posterior puncture situated slightly before level of large lateral puncture; surface of

pronotum with microsculpture similar to that on head. Scutellum with numerous punctures (range 15-19).

Elytra moderately long, at base slightly narrower than pronotum at widest point, at suture feebly shorter

(ratio 0.88) to as long as, at sides feebly longer (ratio 1.11) than pronotum at midline; punctation dense,

intervals between punctures along transverse axis mostly about as large as diameters of punctures;

pubescence dense, dark hairs intermixed with yellowish to golden-yellowish ones; surface between

punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing

whitish apical seam of palisade setae; punctation of tergites dense to very dense, finer than that on elytra,

gradually becoming sparser toward apex of abdomen; pubescense dense, dark hairs intermixed with

scattered yellowish hairs, in addition to lateral patches of yellowish pubescence on each tergite; first three

tergites each with three shallow inconspicuous impressions at base; each impression, including middle one,

with slightly denser punctation and pubescence. Front tarsus dilated in both sexes, inconspicuously so in

female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side; apical margin with moderately deep and wide, obtusely triangular

emargination (Fig. 117), small triangular area before emargination flattened and smooth. Aedoeagus (Figs.

118-121) rather small, median lobe distinctly and rather suddenly narrowed into narrow and obtusely

Revision of the Tribes Quediini and Atanygnathini

251

pointed apical portion bearing distinct dent in lateral view. Paramere nanow and elongate, not quite or just

about reaching apex of median lobe, obtusely arcuate apically; four apical setae, both median setae longer

than lateral setae; two setae at each lateral margin below apex; sensory peg setae on underside of paramere

numerous, forming two long, slightly irregular lateral rows.

Length 4. 0-5. 2 mm.

Type material — Quedius aureiventris. Bernhauer (1915:56) described the

species from specimen(s?) from “Kumaon (Binsar, 7900 Fuss ober dem Meere,

25. Mai 1912, A.D. Imms”. I was not able to locate the original material. My concept

of this species is based on three specimens from the Cameron collection in the

British Museum (Natural History), London. They are labelled as follows; Spec. No.

1 (male); “W. Almora Divn. Kumaon U.P. June’ 17 HGC” / “1084a” / red square

label/ “Quedius aureiventris. det. Bernh. Cam.”. Spec. No. 2 (male): “W. Almora

Divn. Kumaon U.P. Augt., 1917. HGC” / “Quedius aureiventris det. Bernh. Cam.”

Spec. No. 3 (male): “U. Gumti Val.W. Almora Dn. Apr. ’19 HGC” / “Quedius

aureiventris det. Bernh. Cam.” Specimens Nos 1 and 2 were dissected and the 8th

sternites, genital segments and aedoeagi were mounted on plates with beetles.

Quedius decipiens. The original material in the collection of the British Museum

(Natural History), London, contains one male specimen (holotype) under the name

Q. decipiens. It is labelled as follows: “Type” (round label with red margin/ “Ghum

dist. Mangpo V-31 Dr. Cameron” / “Q. decipiens Cam. TYPE” / “M. Cameron.

Bequest. B.M. 1955-147.” The specimen was dissected, sternite 8, genital segment

and aedoeagus were mounted on plate with beetle. It does not specifically differ

from three specimens of Q. aureiventris from Kumaon in Cameron collection (see

above). The name Q. decipiens is a junior synonym of Q. aureiventris. My

corresponding determination label was attached to the specimen.

Geographical distribution. — Quedius aureiventris is widely distributed

throughout the Himalayan range; from northern Pakistan through Kashmir, Nepal

and Darjeeling area to Bhutan (Map 19).

Material studied. — 815 specimens.

BHUTAN. Sampa-Kotoka, 1400-2600 m, 9.VI.72 (NHMB) 1.

INDIA. Kashmir. “Feronepor Nala” (BMNH) 1; locality illegible (BMNH) 1. Himachal Pradesh.

Mandi, Jhatingri, 6000', H.G. Champion (BMNH) 22. Uttar Pradesh. Chakrata Distr: Jaunsar, VI. 1929, H.G.

Champion (BMNH) 1; Manjagaon, 6500', 21.V.22, Cameron (BMNH) I; Deoban, 3.V.21, Cameron

(BMNH) 1; Jadi Gad, 7000', 9.V.22, Cameron (BMNH) 1; Patra Khud, 8000', 15.V.22, Cameron (CNCC) 1.

Dodora Khud, 8000', 13.V.22, Cameron (BMNH) 3; Korawa Khud, 9100', 4.V.22, Cameron (BMNH,

CNCC) 2. Mussoorie Distr.: Mossy Falls, 22.111.32, H.G. Champion (BMNH)3; Dhobi Ghat (BMNH) I.

Kumaon: W. Almora, H.G. Champion (BMNH) 8; Nainital Div., H.G. Champion (BMNH) 1; Nainital,

1800-2000 m, 3.V.78, W. Wittmer (NHBM) 1; Chaubattia nr. Ranikhet, cca 18000 m, 12-13.X.79, I. Lobl

(MHNB) 1; Bhim Tal, cca 1800 m, 9.X.79, I. Lobl (MHNG) 1; same, 1450-1550 m, 5.X.79, I. Lobl

(MHNG) 1. Garhwal: Rangarh, crest, 2250 m, 9.X.79, I. Lobl (MHNG) 6; 16 km off Srinagar, 550 m,

29.X.79, I. Lobl (ASCC, MHNG) 19; between Tehri and Srinagar, 900 m, 25.X.79, 1. Lobl (MHNG) 12; 22

km N Rishikesh, 450 m, 30.X.79, I. Lobl (MHNG) 4; 10 km off Chamba, 2200 m, I. Lobl (MHNG) 2;

Dhanolti, 2200 m, 20.X.79, 1. Lobl (MHNG) 4; 2 km E Dhanolti, 2250 m, 21.X.79, I. Lobl (MHNG) 1;

above Pauri, 900 m, 28.X.79, I. Lobl (MHNG) 3. West Bengal. Darjeeling Distr.: Darjeeling Hills,

Sandakpnu, 12000', 5. V. 34, H.G. Champion (BMNH) 8; Singmark-Bharapatea Bung, 10.V.75, W. Wittmer

(NHMB) 1; Ghoom-Lopchu, 2000 m, 12 and 14.X.78, Besuchet and Lobl (MHNG) 3; Ghum district,

V-VI-31, Cameron (BMNH) 2.

Quaest. Ent., 1988, 24 (2)

252

Smetana

NEPAL. Kathmandu Distr. Gokarna Forest nr. Kathmandu, 1400 m, 31. III. and 1.IV.81, Lobl &

Smetana (ASCC) 4; same, 1300 m, 9. IV. 85, A. Smetana (ASCC) 5; Siwapuri Dara, 2300-2520 m, 1 and

3.V.85, A. Smetana (ASCC) 5; above Sundarijal, 2000 m, 4.IV.8I, Lobl & Smetana (ASCC) 1; Burlang

Bhanjyang, 2600 m, 5.IV.81, Lobl & Smetana (ASCC, CNCC, MHNG) 9; Chaubas, 2600 m, 5.IV.81, Lobl

& Smetana (MHNG) 1; Chipling, 2300 m, 5.IV.81, Lobl & Smetana (ASCC) 1; Gul Bhanjyang, 2600 m,

6.IV.81, Lobl & Smetana (ASCC, MHNG) 3; Kutumsang, 2200-2400 m, 6.IV.81, Lobl & Smetana (ASCC,

BMNH, CNCC, MHNG) 27; Malemchi, 2800 m, 14.IV.81, Lobl & Smetana (ASCC, CNCC, MHNG) 26;

below Tarke Ghyang, 2600 m, 25.IV.81, Lobl & Smetana (ASCC) 2; NE Barahbise, 2500 m, 2.V.81, Lobl &

Smetana (ASCC) 1; Dobate Ridge NE Barahbise, 2800 m and 3000 m, 2 and 7.V.81, Lobl & Smetana

(ASCC, MHNG) 13; Siwapuri Dara, 2540 m, 1.V.85, A. Smetana (ASCC) 1. Lalitpur Distr. Phulcoki,

1700-2500 m, 10.V.81, I. Lobl (MHNG) 8; same, 20-21. IV.82, 2550-2600 m, A. & Z. Smetana (ASCC,

CNCC) 30; same, 14 and 16.X.83, 2600 m Smetana & Lobl (ASCC) 3; same, 28.30.IV.84, 2550-2700 m,

Smetana & Lobl (ASCC, BMNH, CNCC, MHNG) 510; same, 2000 m, 11-14.VI.76, W. Wittmer & C.

Baroni (NHMB) 1. Khandhari Distr. Forest above Ahale, 2300 m, 26. III. 82, A. & Z. Smetana (ASCC) I;

same 2200 m, 4. IV. 84, Smetana & Lobl (ASCC) 2; Forest NE Kuwapani, 2450 m, 13. IV. 82, A. & Z.

Smetana (ASCC) 2; same, 2400 m, 5. IV. 84, Smetana & Lobl (ASCC) 2; Chichila-Mure, 1900 m, 24.V.80,

W. Wittmer (NHMB) 3; below Sheduwa, 2550 m, 30.III.82, A. & Z. Smetana (ASCC) 2; above Sheduwa,

.3000 m, 1 and 2.IV.82, A. & Z. Smetana (ASCC) 3; “Bakan” W of Tashigaon, 3200 m, 3-6.IV. 82, A. & Z.

Smetana (ASCC, CNCC) 1 1; Ridge S Mansingma, 2600 m, 8. IV. 84, Smetana & Lobl (ASCC) 1; Forest S

Mansingma, 2250 m, 12. IV. 84, Smetana & Lobl (ASCC) 2; Induwa Khola Valley, 2050 m, 17. IV. 84,

Smetana & Lobl (ASCC) 3. Mustang Distr. Lete, 2550 m, 2.X.83, Smetana & Lobl (ASCC) 4. Rasuwa

Distr. 1.5 km NE Bhargu, 2000 m, 12. IV. 85, A. Smetana (ASCC) 2. Chapuri, 16. III. 79, de Rougemont

(GDRC) 1; Aharbal, VI.81, de Rougemont (GDRC) 1; Phalkuwa, 2500 m, IV. 84, de Rougemont (GDRC) I.

PAKISTAN. Murree-Abbotta, 2200-2500 m, 13.VI.77, Wittmer & Brancucci (NHMB) 1; Chhangla

Gall, 7000’, Hazara, 12.VI.74, C. Baroni Urbani (NHMB) 1.

Bionomics. — Quedius aureivenths is a rather common species, occurring from

low elevatons of 550 m to middle elevations of over 3000 m ; however, most

specimens were taken at elevations between 2100-2280 m. Specimens were

collected mainly in forest habitats by sifting leaf litter and other forest floor debris,

by sifting moss on rocks or on standing live trees or fallen dead trees, and by sifting

various decaying plant matter. The species is extremely common on the mountain

Phulcoki near Kathmandu in the spring in and under decaying last years vegetation

at bases of large rocks on forest clearings or in ditches along the road to the

microwave tower on the top of Phulcoki. It is worth mentioning that it almost

completely disappears from these habitats on Phulcoki in the fall after the monsoon

(only very few specimens were found).

Comparisons and variations. — Quedius aureiventris can be easily recognized

among the winged, small species of Raphiriis by the type of pubescence of the elytra

and abdomen, particularly by the presence of three shallow, inconspicuous

impressions at the bases of the first three visible abdominal tergites with slightly

denser punctation and pubescence, in combination with the distinctly, partially

darkened legs (see the description for details). None of the species that resemble Q.

aureiventris (e.g., Q. daksumensis, Q. vadhu or Q. paschim) has the basal

impressions on the abdominal tergites.

The density of the punctation and pubescence of the abdominal tergites varies to

some extent in this species.

The holotype of Q. decipiens has atypically coloured antennae; the right antenna

is dark from segment 6 and the left antenna is dark already from segment 2, but

Revision of the Tribes Quediini and Atanygnathini

253

segments 2-5 of the latter have irregular pale spots. This colouration is either

artificial or of teratological nature.

32. Quedius (Raphirus) muscicola Cameron

Figs. 122-132; Map 20

Quedius muscicola Cameron 1932:295

Quedius doherQi C&meron 1932:297 (syn.nov.)

Quedius heterogaster Cameron 1944: 14 {syiuwv.)

Description. — Piceous-black, elytra usually more or less paler, dark rufo-brunneous to

brunneopiceous; head, pronotum and elytra with faint to slight metallic bronze reflections, abdomen

iridescent; both labial and maxillary palpi, antennae and legs uniformly testaceous. Head rounded, slightly

wider than long (ratio 1.25); eyes large and convex, tempora very short, considerably shorter than length of

eyes seen from above (ratio 0.20); no additional setiferous punctures between anterior frontal punctures;

posterior frontal puncture touching posteromedian margin of eye, one puncture between it and posterior

margin of head; temporal puncture touching posterior margin of eye; surface of head with fine and dense

microsculpture of transverse and oblique waves gradually changing into meshes on middle of clypeus.

Antenna moderately long, segments 2 and 3 about equally long, segments 4-7 longer than wide, gradually

becoming shorter, segments 8-10 about as long as wide, segment 11 as long as two preceding segments

combined. Pronotum about as long as wide, widely rounded basally and distinctly nan'owed anteriorly,

evenly transversely convex; dorsal rows each with three punctures; sublateral rows each with two punctures,

posterior puncture situated slightly before level of large lateral puncture; surface of pronotum with

microsculpture of transverse waves similar to those on head. Scutellum with numerous punctures (range

9-15). Elytra moderately long, at base slightly naiTOwer than pronotum at widest point, at suture about as

long as, at sides slightly longer (ratio 1.16) than pronotum at midline; punctation fine and dense; intervals

between punctures along transverse axis about as large as diameters of punctures; pubescence dense, golden

yellowish; surface between punctures without microsculpture. Wings fully developed. Abdomen with tergite

7 (fifth visible) bearing whitish apical seam of palisade setae; punctation and pubescence of tergites in

general more or less dense, gradually becoming sparser both toward apex of each tergite and toward apex of

abdomen; pubescence golden yellowish, forming distinct patch of dense hairs on either lateral portion of

each tergite. Front tarsus dilated in both sexes, inconspicuously so in female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side; apical margin with moderately wide and not deep obtusely triangular

emargination (Figs. 122, 123), small triangular area before emargination flattened and smooth. Aedoeagus

(Figs. 124-132) moderately large; median lobe with apical portion rather variable; mostly very slightly

widened before rounded apical margin, or less frequently variably dilated apically; when paramere removed,

with short median carina forming a fine hook in lateral view. Paramere not quite reaching, about reaching or

slightly exceeding apex of median lobe, elongate and in general feebly narrowed and with subacute apex;

four setae at apical margin, median two setae somewhat longer than lateral ones, two similar setae at each

lateral margin below apex; sensory peg setae on underside of paramere numerous, forming two irregular,

moderately long rows.

Length 4. 8-6.0 mm

Type material. — Quedius muscicola. The collection of the British Museum

(Natural History), London, contains five syntypes under the name Q. muscicola.

They are labelled as follows: Spec. No. 1 (?): “Type” (round label with red margin)/

“SYN-TYPE” (round label with blue margin)/ “Ghum district V- VI-31 Dr.

Cameron” / “Quedius muscicola Cam. TYPE” / “M. Cameron. Bequest. B.M.

1955-147”. Spec. No. 2 (?): Ghum dist. Tiger Hill 8,500-10,000 ft. Dr. Cameron” /

“M. Cameron. Bequest. B.M. 1955-147” / “SYN-TYPE” (round label with blue

margin). Spec. No. 3 (?): same labels as No. 2. Spec. No. 4 (c7): “Moss” / “Ghum

dist. V-VI-31 Dr. Cameron” / “M. Cameron. Bequest. B.M. 1955-147” /

Quaest. Ent., 1988, 24 (2)

254

Smetana

“SYN-TYPE” (round label with blue margin). Spec. No. 5 (d'): same labels as Spec.

No. 2 (teneral specimen).

The male specimen No. 4 was dissected, the aedoeagus (paramere separated) and

the genital segment were mounted on plate with beetle. The specimen is hereby

designated as the lectotype of Q. muscicola; the label “Lectotype Quedius muscicola

Cameron Smetana des. 1984” has been attached to it.

Quedius dohertyi. The Cameron collection in the British Museum (Natural

History), London, contains one male specimen (holotype) under the name Q.

dohertyi. It is labelled as follows; “Type” (round label with red margin)/ “64518” /

“Doherty” / “Fry Coll. 1905.100” / “Birmah Ruby Mes” / “Q. dohertyi Cam.

TYPE”.

The specimen (broken in two pieces: head & prothorax and elytra & abdomen)

was remounted, the sternite 8, and the aedoeagus were mounted in Canada Balsam,

and the genital segment on plate with beetle.

The specimen cannot be distinguished from specimens of Q. muscicola with

sparsely punctate abdomen. The name Q. dohertyi is a junior synonym of Q.

muscicola; my corresponding determination label has been attached to the specimen.

Quedius heterogaster. The collection of the British Museum, (Natural History),

London, contains two conspecific syntypes under the name Q. heterogaster. They

are labelled as follows: Spec. No. 1 (9): “Type” (round label with red margin) “Dr.

Cameron”. 1 4-22. V. 22” / “Chakrata Dist. Kanasar 7050'”/ “Q. heterogaster Cam.

TYPE” / “M. Cameron. Bequest. B.M. 1955-147.” Spec. No. 2 (d): “SYN-TYPE”

(round label with blue margin)” “Chakrata Dist. Kanasar 7050'”/ “Dr. Cameron.

14-22.V.22”/“M. Cameron. Bequest. B.M. 1955-147.”

The second (male) specimen was dissected, the aedoeagus (paramere separated)

and the genital segment were mounted on plate with beetle. The specimen is hereby

designated as the lectoptype of Q. heterogaster; the label “Lectotype Quedius

heterogaster Cameron Smetana des. 1984” has been attached to it.

Neither specimen can be distinguished from the lectotype of Q. muscicola. The

name Q. heterogaster is a junior synonym of Q. muscicola. My corresponding

determination labels were attached to both specimens.

Geographical distribution. — Quedius muscicola is widely distributed

throughout the Himalayan range from northern Himachal Pradesh and Uttar Pradesh

through Nepal eastwards through West Bengal (Darjeeling area) to Burma (Map

20).

Material studied. — 108 specimens.

BURMA. See the Type material of Q. dohertyi.

INDIA. Himachal Pradesh. Kulu, Parbatti V., 6000-8000', H.G. Champion (BMNH) 3. Uttar Pradesh.

Chakrata Distr. Korawa Khud, 9100', 4.V.22, Cameron (BMNH) 3; Patra Khud, 8000', 15.V.22, Cameron

(BMNH) 2. Kumaon: W. Almora, H.G. Champion (BMNH 3; W. Almora Divn., VI. 1917, H.G. Champion

(BMNH) 1. Garhwal: 10 km E Chamba, 2200 m, 2. and 20.X.79, I. Lbbl (MHNG) 2; between Tehri and

Srinagar, 900 m, 25.X.79, I. Lobl (MHNG) 1; 10 km E Dhanolti, 2450 m, 21.X.79, I. Lobl (MHNG) 1;

above Pawn, 1900 m, 28.X.79, 1. Lobl (ASCC, MHNG) 6; 6 km E Dhanolti, 2300 m, 21.X.79, I. Lobl

(MHNG) 1; Dhanolti, 2200 m, 20.X.79 (MHNG) 3; 2 km E Dhanolti, 2250 m, 21.X.79, I. Lobl (MHNG) 2;

Revision of the Tribes Quediini and Atanygnathini

255

4 km S Bhatwari, 1400 m, 2I.X.79 (MHNG) 1. West Bengal. Darjeeling Distr. Ghoom-Lopchu, 200 m.

14.X.78, Besuchete & Lobl (MHNG) 2; 13 km N Ghoom, 1500 m, 15.X.78, Besuchet & Lobl (ASCC,

MHNG) 7; Ghoom, 1500 m, 15.X.78, Besuchet & Lobl (MHNG) 1; Algarah, 1800 m, 9.X.78, Besuchet &

Lobl (MHNG) 1; Singmari-Bharapatea Bung, 10.V.75, W. Wittmer (NHMB) 1; Darjeeling (BMNH) 1;

Ghum Distr., V-VI-31, Cameron (BMNH) 4; Ghum Distr., Tiger Hill, 8500-10000', V-VI-31, Cameron

(BMNH, CNC0 4; Lebong, 1600-1800 m, 8.V.75, 'W. Wittmer (NHMB) 1.

NEPAL. Kaski Distr. above Dhumpus, 2100 m, 8-10.V.80, Martens & Ausobsky (SBMF) 4. Khandhan

Distr. Forest above Ahale, 2300 m, 26.111.82, A. & Z. Smetana (ASCC) 2; Chichila S., Ahale, 2200 m, 4 and

24.IV. 84, Fobl & Smetana (MHNG) 2; Chichila-Mure, 1900 m, 24.V.80, W. Wittmer (NHMB) 2; Forest NE

Kuwapani, 2450-2500 m, 28. III., 13. IV. and 14.IV.82, A & Z Smetana (ASCC) 3; same, 2350-2400 m,

5. IV. 84, Smetana & Fobl (ASCC, CNCC, MHNG) 7; same, 2250 m, 6.IV.84, Smetana & Lobl (ASCC) 1;

same, 2400-2450 m, 29.IV. 84, Smetana & Lobl (ASCC) 2; 2 km E Mansingma, 1900 m, 19.IV. 84, Lobl &

Smetana (ASCC, MHNG) 4; Pass NE Mangmaya, 2300 m, 6.IV.84, Fobl & Smetana (ASCC, MHNG) 2;

Ridge NE Mangmaya, 2800 m, 7.IV.84, Smetana & Lobl (ASCC) 1; Ridge S Mansingma, 2800 m, 7. IV. 84,

Smetana & Lobl (ASCC) 1; Forest S Mansingma, 2300 m, 12. IV. 84, Smetana & Lobl (ASCC) 1; Induwa

Khola Valley, 200-2050 m, 16.1V. 84, Smetana & Lobl (ASCC) 2. Lalitpiir Distr. Phulcoki, 2600 m,

12.VI.76, Wittmer & Baroni (NHMB) 1; same, 1700 m, 10.V.8 1 , Lobl (MHNG) 1; same, 2550 m, 21.1 V.81,

A. & Z. Smetana (ASCC) 1; same, 2500-2550 m, 28-29.IV.84, Lobl & Smetana (ASCC, MHNG) 7; 2 km S

Godavari, 1700 m, 12. IX. 83, Smetana & Lobl (ASCC) 1. Rasiiwa Distr. 1.5 km NE Bhargu, 2000 m,

12.IV. 85, A. Smetana (ASCC) 1. Gul Bhanjyang, 2600 m, 6.IV.81, Lobl & Smetana (ASCC) 1; Dobate

Ridge NE Barahbise, 2700-2800 m, 3 and 7.V.81, Lobl & Smetana (ASCC, MHNG) 2.

Bionomics. — Quediiis muscicola typically occurs at lower elevations, from

about 900 m to 2800 m. Specimens were collected in forest habitats near creeks or

dry creek beds (in dry season), near seepages or other similar habitats by sifting

moist to wet moss on large rocks and fallen trees, less frequently also by sifting wet

forest floor debris, leaf litter etc. Some specimens were also taken by sifting flood

debris along small creeks (after rainstorms) and by sifting debris under large ferns

around a seepage.

Comparisons and variations. — Quedius muscicola can fairly easily be

recognized among the fully winged small species of Raphirus by the rather pale

colouration (the elytra are usually paler than the rest of the body and all appendages

are uniformly testaceous), by the fairly long elytra with mostly dense punctation and

golden yellowish pubescence, and by the golden yellowish pubescence of the

abdomen forming distinct lateral patches of denser hairs.

Quedius muscicola is a rather variable speeies, particularly in the density of the

punctation and pubescence of the abdominal tergites, and in the shape of the

aedoeagus (see Figs. 124-132); however, there is no correlation between various

shapes of the aedoeagus and the density of the abdominal punctation. Specimens

from the western portion of the Himalaya (Himachal Pradesh, Uttar Pradesh and

eastwards to about central Nepal) have the abdominal tergites densely punctate and

pubescent and also the elytra are rather finely and densely punctate. East of central

Nepal specimens with more or less sparsely punctate and pubescent abdominal

tergites start to appear and may be dominant in some populations; in some

specimens the tergites appear almost impunctate and glabrous except for small

lateral patches of dense punctures and hairs (such specimens occur, e.g., in the

Induwa Khola valley in eastern Nepal or in Darjeeling area in West Bengal, and the

holotype of Q. dohertyi from Burma also belongs to this form) and also the

Quaest. Ent., 1988, 24 (2)

256

Smetana

punctation of the elytra tends to be more or less coarser and sparser. Such specimens

were never found west of eastern Nepal. If a densely punctate and pubescent

specimen from the western portion of the distributional range is compared with an

eastern specimen with an almost glabrous abdomen, it seems to be impossible that

they could be conspecific. However, there is a continuous intergradation between

these two extremes and there are no consistent differences in the shape of the

aedoeagus (see above). I am therefore reasonably confident that we deal here with

only one species. Nevertheless, should the specimens with almost glabrous

abdomianl tergites prove to be specifically different in the future, the name Q.

dohertyi should be used for them.

Quedius satoi resembles Q. nniscicola\ however, the former species differs, in

addition to the distinctly different shape of the aedoeagus (see Figs. 124-132,

152-155), by the wider head, by the shorter elytra, by the wings reduced to

nonfunctional stumps (see under Q. satoi) and by the very delicate, sometimes

hardly visible whitish apical seam of palisade setae on the fifth visible tergite. Both

species often occur together in eastern Nepal.

For comparison with Q. hhari see under that species.

33. Quedius (Raphiriis) hhari spec. now

Figs. 133-136; Map 21

Description. — in all characters, including general habitus, colouration and pubescence very similar

to Q. nmscicola and differing mainly by characteristic shape of aedoeagus and different emargination of

male stemite 8. Punctation of abdominal tergites moderately dense and almost evenly distributed; however,

in a few specimens sparse and leaving some medioapical areas of tergites almost impunctate.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 (Fig. 133)

with two long and strong setae on each side; emargination of apical margin much deeper and wider than that

of Q. muscicola (Figs. 122, 123). Aedoeaugs (Figs. 134-136) quite characteristic; median lobe with two

dilatations, one larger in about middle and one smaller below apex, lateral outlines of median lobe therefore

characteristically bisinuate; apex of median lobe subacute. Paramere not reaching apex of median lobe, with

subacute apex; four setae at apex, all four almost equally long and strong, two similar setae at each lateral

margin just below apex; sensory peg setae on underside of paramere numerous, forming two rather long,

slightly irregular rows.

Length 4.7-5. 1 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL, Khandbari

District” / “For. above Ahale 2400 m 25. III. 82 A. & Z. Smetana”. Both holotype

and allotype in the Smetana collection, Ottawa, Canada.

Paratypes (9): same data as holotype but 2300 m and date 26.III.82 (CNCC) 1;

Khandbari Distr. Kuwapani, 2100 m, 28. III. 82, A. & Z. Smetana (ASCC) 1; Induwa

Khola Valley, 2000 or 2050 m, 16.IV.84, Smetana & Ldbl (ASCC, BMNH) 4; Val.

Induwa Kola, 2000 m, 16. and 18. IV. 84, Lobl & Smetana (MHNG) 3.

Geographical distribution. — Quedius hhari is known from two localities in the

foothills of the main Himalayan range in eastern Nepal (Map 21).

Bionomics. — The specimens of the original series were taken by sifting wet

moss on rocks (Forest above Ahale), by sifting soaking wet moss around a small

Revision of the Tribes Quediini and Atanygnathini

257

forest spring, or by sifting moss peiTnanently sprinkled by water from a small

waterfall (Induwa Khola Valley).

Comparisons. — Specimens of Q. hhari are in general somewhat smaller than the

average specimens of Q. muscicola, with shorter antennae, more parallel-sided

pronotum, and shorter and more densely punctate elytra; however, the shape of the

aedoeagus is necessary for positive determination.

Etymology.- — The specific name is the Nepali adjective bhari (full); it refers to

the double dilatation of the median lobe of the aedoeagus.

34. Quedius (Raphirus) eklai spec.nov.

Figs. 137-140; Map 21

Description . — in all characters very similar to Q. muscicola but different as follows: average size

slightly smaller; colouration similar but median face of hind tibiae darkened; head and pronotum slightly

narrower, latter less distinctly nanowed anteriorly than in most specimens of Q. muscicola: elytra somewhat

shorter, at suture feebly shorter (ratio 0.86), at sides about as long as pronotum at midline, pubescence of

elytra and abdomen extensively golden yellow.

Male. First four segments of front tarsus not appreciably more dilated than in female. Sternite 8 (Fig.

137) with two long and strong setae on each side, emargination of apical margin about same as that of Q.

muscicola (Figs. 122, 123). Aedoeagus (Figs. 138-140) fairly small. Median lobe vaguely constricted in

middle portion, with subacute apex; apical portion, when paramere removed, with fine and short median

Carina forming a fine hook in lateral view. Paramere reaching apex of median lobe, slightly asymmetrical,

with bisinuate lateral margins; four short apical .setae and two equally short setae at each lateral margin not

far below apex of paramere; sensory peg setae on underside of paramere forming two slightly irregular and

rather short lateral rows.

Length 4. 8-5.0 mm.

Type material. — Holotype (male) and allotype (female): Nepal; “172 Parbat

Dist., zwischen Chitre und Ghandrung, Chitre-Seite des Passes, Tsuga-Rhodod.

2800-2900 m Martens & Ausobsky leg. 4/7 Mai 80”. In the Senckenberg Museum,

Frankfurt a.M., West Germany (holotype) and in the Smetana collection, Ottawa,

Canada (allotype).

Geographical distribution. — Quedius eklai is at present known only from the

type locality in western Nepal (Map 21).

Bionomics. — Both specimens were taken by sifting in a moist Tsuga-

Rhododendron forest but no real details are known.

Comparisons. — Quedius eklai can be distinguished from Q. muscicola, and also

from Q. hhari, in addition to the differences in the shape of the aedoeagus, by the

characters given in the key and in the description. There is a slight chance that the

somewhat asymmeterical shape of the paramere is of a teratological nature;

however, I had a similar feeling about the shape of the median lobe when I saw the

first male of Q. hhari, until I later found an additional series of males with

identically shaped median lobe. The short, equally long apical setae on the paramere

also seem to be characteristic of Q. eklai. For these reasons, I decided to consider

the two specimens as a separate species; however, more males are needed to confirm

its status.

Quaest. Ent., 1988, 24 (2)

258

Smetana

Etymology. — The specific name is the Nepali word eklai (alone).

35. Quedius (Raphirus) sundar spec.nov

Figs. 141-145; Map 17

Description. — Black, head, pronotum and elytra with dark metallic reflections, abdomen iridescent;

palpi, antennae and legs entirely pale testaceous. Head rounded, feebly wider than long (ratio 1.16); eyes

very large and convex, temples very short, considerably shorter than length of eyes seen from above (ratio

0.20); no additional punctures between anterior frontal punctures; posterior frontal puncture touching

posteromedian margin of eye, one puncture between it and posterior margin of head, temporal puncture

touching posterior margin of eye; surface of head with dense and very fine microsculpture or irregular

transverse waves becoming almost meshed on clypeus. Antenna moderately long, segments 2 and 3 about

equally long, segments 4-6 longer than wide, gradually becoming shorter and slightly wider, segments 7-9

about as long as wide, segment 10 feebly transverse, last segment about as long as two preceding segments

combined. Pronotum about as long as wide, widely arcuate basally, evenly transversely convex, feebly to

slightly narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with two punctures,

posterior puncture situated before level of large lateral puncture; surface of pronotum with microsculpture

similar to that on head but finer. Scutellum with 3 to 7 punctures. Elytra moderately long, at base about as

wide as pronotum at widest point, at suture about as long as, at sides slightly longer than pronotum at

midline (ratio 1.25); punctation moderately coarse, sparse intervals between punctures along transverse axis

mostly several times larger than diameters of punctures; pubescence sparse, short and yellowish; surface

between punctures without microsculpture but with some microscopic inegularities. Wings fully developed.

Abdomen with tergite 7 (fifth visible) with very distinct whitish apical seam of palisade setae; punctation of

tergites extremely fine and very sparse, large middle portion of each tergite across its entire width glabrous,

without any punctation or pubescence; pubescence in general very sparse, golden-yellow, forming lateral

patches of denser and longer hairs on each tergite. Front tarsus not dilated in either sex.

Male. Sternite 8 (Fig. 141 ) with two long and strong setae on each side, apical margin with rather wide

and deep obtusely triangular emargination; pubescence of sternite in general sparse. Aedoeagus (Figs.

142-145) rather small; median lobe with apical margin obtusely arcuate, apical portion in lateral view with

minute dent. Paramere relatively wide, just about reaching apex of median lobe, apex narrowly arcuate; four

setae at apical margin, median setae longer than lateral ones; two unequally long setae at each lateral margin

Just below apex; sensory peg setae on underside of paramere not numerous, forming two somewhat irregular

longitudinal rows, each with 7-8 tubercles.

Length 4. 0^,8 mm.

Type material. — Holotype (male): “NEPAL, Khandbari District” / “For. NE

Kuwapani 2500 m 18. III. 82 A. & Z. Smetana”. Allotype (female): same data as

holotype but 2450 m and date 13. IV. 82. Both holotype and allotype in the Smetana

collection, Ottawa, Canada.

Paratypes (9): same data as holotype (ASCC) 1; same data as allotype (CNCC)

1; same data as holotype, but 2500 or 2550 m and date 14. IV. 82 (ASCC, CNCC) 3;

For. above Ahale, 2300 m, 26.III.82, A. & Z. Smetana (ASCC, BMNH) 2;

Khandbari Distr.: Induwa Khola Valley, 2050 m, 24. IV. 84, Lobl & Smetana

(MFING) 1; Chichila s/Ahale, 2200 m. 24.IV. 84, Lobl & Smetana (MHNG) 1.

Geographical distribution. — Quedius sundar is known from a few localities in

the foothills of the main Himalayan range in eastern Nepal (Map 17).

Bionomics. — The specimens of the original series were typically found by

sifting moist to wet moss on rocks near creeks or forest seepages.

Comparisons and variations. — Quedius sundar is very easy to recognize mainly

by the sparse and moderately coarse punctation of the elytra and the extremely fine

Revision of the Tribes Quediini and Atanygnathini

259

and very sparse punctation of the abdominal tergites, leaving large middle portion of

each tergite glabrous (see the description for details), in combination with the dark

metallic reflections of dorsal surface of the body, the entirely pale testaceous

appendages, and the sparse golden-yellow pubescence of abdominal tergites forming

patch of dense hairs on either lateral portion of each tergite.

Specimens of Q. muscicola with extremely sparse punctation of the abdominal

tergites may superficially resemble Q. simdar, however, they differ, in addition to

the differently shaped aedoeagus (see Figs. 124-132, 142-145) and to the larger

size, by the characters given in the key: see couplet 19.

The pubescence of elytra is very indistinct in some specimens.

Etymology. — The specific name is the Nepali adjective sundar (beautiful); it

refers to the general appearance of this species.

36. Quedius (Raphirus) udagra Smetana

Figs. 146-150; Map 21

Quedius udagra Smetana 1975:235

Description. — Piceous to piceous-black with black head, apical margins of abdominal tergites

sometimes paler, occasionally also elytra and pronotum slightly paler. Abdomen feebly iridescent; palpi,

antennae and legs testaceous, posterior tibiae distinctly, middle tibiae indistinctly to slightly darkened at

inner margin. Head rounded, slightly wider than long (ratio 1.16); eyes very large and convex, temples

extremely short, considerably shorter than length of eyes seen from above (ratio 0.15); no additional

setiferous punctures between anterior frontal punctures; posterior frontal puncture situated at posteromedian

margin of eye, one puncture between it and posterior margin of head; surface of head with fine and dense

microsculpture of transverse waves becoming irregular toward clypeus and forming here and there irregular

meshes. Antenna moderately long, segment 3 slightly narrower and about as long as segment 2, segments 4

and 5 distinctly longer than wide, following segments gradually becoming shorter, outer segments

indistinctly to slightly longer than wide, last segment almost as long as two preceding segments combined.

Pronotum about as long as wide to feebly wider than long (ratio 1.09), widely arcuate basally, evenly convex

and moderately narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with two or

three punctures, posterior puncture not or about reaching (if three punctures present) level of large lateral

puncture; surface of pronotum with microsculpture of transverse waves usually slightly denser and finer than

those on head. Scutellum with 8-16 (usually 9-1 1) punctures. Elytra short, at suture distinctly (ratio 0.76), at

sides slightly shorter (ratio 0.86) than pronotum at midline; punctation fine and dense, intervals between

punctures along transverse axis usually no more than twice as large as diameters of punctures, pubescence

pale brownish; surface between punctures without microsculpture. Wings reduced to very small

nonfunctional stumps reaching about two thirds of length of elytra. Abdomen with tergite 7 (fifth visible)

lacking whitish apical seam of palisade setae: punctation of tergites usually feebly finer than that on elytra,

slightly denser on basal portions of tergites and in general becoming gradually sparser toward apex of

abdomen; same applies to light brownish pubescence, forming inconspicuous but distinct patch of denser

and slightly paler hairs on either lateral portion of each tergite. Front tarsus dilated in both sexes,

inconspicuously so in female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side; apical margin with moderately deep, obtusely triangular emargination (Fig.

146), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 147-150) rather

wide, median lobe slightly narrowed toward apex and then feebly dilated and broadly rounded apically;

apical part, when paramere removed, with fine median carina forming a fine hook in lateral view. Paramere

not reaching apex of median lobe, spatulate; with four setae at apical margin, both median setae longer than

lateral setae, and with a pair of unequally long setae at each lateral margin near apex; sensory peg setae on

underside of paramere numerous, forming two very irregular longitudinal rows.

Quaest. Ent., 1988, 24 (2)

260

Smetana

Length 5. 0-6. 8 mm.

Type material. — The species was described from four specimens from Phulcoki

near Kathmandu; the male holotype and female allotype bear the locality label:

“Phulchoki b. Kathmandu, Nepal, Ig. Franz” (with code “Pa 151” on the underside)

and the label “HOLOTYPE (or ALLOTYPE) Quedius udagra A. Smetana 1975”.

The two paratypes bear same locality label as holotype, except code “Pa 15”. The

holotype and allotype are deposited in the Eranz collection, Modling, Austria; the

paratypes are deposited in the Canadian National Collection, Ottawa (CNC No.

13965).

Geographical distribution. — Quedius udagra is known only from the mountain

Phulcoki in the Mahabharat Range at the southern edge of the Kathmandu valley

(Map 21).

Material studied. — 105 specimens.

NEPAL. Lalitpur Distr. Phulcoki, 2500 m, 10.V.81, L Lobl (MHNG) 3; 2550-2650 m, 20-22.1V. 82, A.

& Z. Smetana (ASCC, CNCC) 21; 2550-2650 m, 13.-16.X.83, Smetana and Lobl (ASCC, BMNH, CNCC)

41; 2400-2550 m, 28-30.lV.84, Lobl and Smetana (ASCC, BMNH, CNCC, MHNG) 40.

Bionomics. — Quedius udagra occurs near the top of the mountain. The

specimens of the original series were collected at about 2700 m by sifting fallen

leaves on large limestone rocks. Additional specimens were collected slightly lower,

mainly by sifting old dead vegetation, various debris and fallen leaves at bases of

large rocks on clearings in the forest. Some specimens were also taken by sifting

small piles of decaying vegetation along the road to the microwave tower.

Comparisons. — Quedius udagra can easily be distinguished from the other

small brachypterous species of Raphirus with patches of denser pubescence on

abdominal tergites and with darkened posterior tibiae by its rather large size and

robust form, in combination with the characteristic shape of the median lobe and

paramere of the aedoeagus (Pigs. 147, 148, 150). The aedoeagus of Q. udagra,

particularly the apical portion of the median lobe, resembles that of Q. satov,

however, the latter species differs, in addition to some small differences on the

aedoeagus (Pigs. 147-150, 152-155), mainly by the presence of a delicate whitish

apical seam on the fifth visible abdominal tergite, by the uniformly testaceous legs

and by the distinctly more sparsely punctate abdominal tergites.

37. Quedius (Raphirus) satoi spec.nov.

Pigs. 151-155; Map 22

Description. — in all external characters similar to Q. udagra but different as follows: size in

general slightly smaller and form less robust; colouration in general paler, elytra and often also pronotum

(usually less than elytra) brownish to brownish-red; abdomen with apical margins of tergites usually

distinctly paler, abdomen often extensively pale, reddish-brown to almost testaceo-rufous, particularly

anteriorly; anterior portion of head in males more or less paler than in females; palpi, antennae and legs

uniformly testaceous. Antenna with segments 4 and 5 usually slightly shorter. Punctation of elytra in general

sparser and coarser. Wings reduced to nonfunctional stumps reaching to about apical margin of elytron.

Punctation of abdominal tergites much sparser, middle portions of tergites to variable extent with only

scattered punctures or almost impunctate; pubescence paler, yellowish to golden-yellowish; tergite 7 (fifth

Revision of the Tribes Quediini and Atanygnathini

261

visible) with very delicate, sometimes hardly visible whitish apical seam of palisade setae.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 (Figs. 151)

with two long and strong setae on each side; emargination of apical margin usually somewhat deeper and

sharper than that of Q. iidagva (Fig. 146). Aedoeagus (Figs. 152-155) quite similar to that of Q. iidagra but

slightly less robust with apex of median lobe less widely rounded; paramere somewhat narrower, rows of

sensory peg setae on underside in general more regular and each with peg setae less numerous.

Length 4. 5-6. 3 mm

Type material. — Holotype (male) allotype (female): “NEPAL, Khandbari

District” / “For. NE Kuwapani 2450 m 13. IV. 82 A. & Z. Smetana”. Both holotype

and allotype in the Smetana collection, Ottawa, Canada.

Paratypes (161): Nepal: Nuwakot Distr.: Gul Bhanjang, 2600 m, 6.IV.81, Ldbl &

Smetana (ASCC, MHNG) 8; above Shermantang, 2900 m, 26.IV.81, Ldbl &

Smetana (ASCC, MHNG) 10. Khandbari Distr.: same data as holotype (ASCC,

CNCC) 13; same data but 2500 m and date 28.III.82 (ASCC, BMNH, CNCC) 18;

same data but date 11. IV. 82 (ASCC) 6; same data but 2500 m and date 14.IV. 82

(ASCC, BMNH, CNCC) 21; same data but 2400 m and date 5. IV. 84, Smetana &

Ldbl (ASCC, BMNH) 6; same data but 2400 m and date 24. IV. 84, Smetana & Ldbl

(ASCC, CNCC) 8; Foret N-E Kuwapani, 2350 m, 5.IV.84, Ldbl & Smetana

(MHNG) 7; same but 2250 m and date 24. IV. 84 (MNHG) 5; above Sheduwa, 3000

m, 31.III.-1.IV.82, A. & Z. Smetana (ASCC, BMNH, CNCC) 13; same, but date

2. IV. 82 (ASCC) 2; above Tashigaon, 3100 m, 7. or 8. IV. 82, A. & Z. Smetana

(ASCC) 2; Pass NE Mangmaya, 2300 m, 6.IV.84, Smetana & Ldbl (ASCC) 5; Col

N-E Mangmaya, 2300 m, 6. IV. 84, Ldbl & Smetana (MHNG) 3; Ridge S

Mansingma, 2800 m, 7. IV. 84, Smetana & Ldbl (ASCC) 1; Forest S Mansingma,

2300 m, 12. or 13.IV.84, Smetana & Ldbl (ASCC, BMNH) 7; Induwa Khola Valley,

2000 m, 14. or 16.IV.84, Smetana & Ldbl (ASCC, CNCC) 6; same but 2800 m and

date 15.IV.84 (ASCC) 1; Val. Induwa Kola, 2100 m, 17.IV.84, Ldbl & Smetana

(MHNG) 4. Forest S Mansingma, 2200 m, 11. IV. 84, Smetana & Ldbl (ASCC,

CNCC) 8; Foret S Mansingma, 2200 m. 1 1.IV.84, Ldbl & Smetana (MHNG) 7.

Geographical distribution. — Quedius satoi is distributed from central to eastern

Nepal (Map 22).

Bionomics. — Quedius satoi is a rather common species, particularly in eastern

Nepal, typically occurring at lower elevations between 2000-3000 m in the

evergreen or semideciduous broadleaved forest. Specimens were taken by sifting

various forest-floor litter, such as fallen leaves, dead decaying vegetation and other

debris, also by sifting moist moss along bases of large rocks or on large rocks or

fallen trees. Some specimens were also taken by sifting of debris under

Rhododendron bushes, or in forest clearings by sifting of moss and debris under

large ferns.

Comparisons and variations. — The density of punctation of the abdominal

tergites varies distinctly in this species; however, in most specimens the middle

portions of tergites are only sparingly punctate; specimens with middle portions of

tergites impunctate or to the contrary more than sparingly punctate are rare.

Quaest. Ent., 1988, 24 (2)

262

Smetana

The similarity of the aedoeagus of Q. satoi with that of Q. udagra (see Figs.

152-155, 147-150) is remarkable, nevertheless there is no doubt that two separate

species are involved. In addition to the morphological differences (see above), both

species differ in their distributional ranges. Quedius udagra is apparently endemic to

the higher elevations of the mountain Phulcoki in the Mahabharat Range at the

southern edge of the Kathmandu Valley; Q. satoi, on the other hand, is widely

distributed in the foothills of the main Himalayan range in central and eastern Nepal.

Quedius satoi resembles also Q. tikta, particularly due to the sexually dimorphic

colouration of the head (paler anteriorly in the males); however, Q. tikta can easily

be distinguished, in addition to the differently shaped aedoeagus (see Figs. 152-155,

176-179), by the in general somewhat smaller size, by the distinctly more densely

punctate and pubescent elytra and the abdominal tergites, and by the absence of the

whitish apical seam of palisade setae on fifth visible tergite. Also, Q. tikta occurs at

higher elevations, from about 2800 m to nearly 5000 m (see under Q. tikta for

details); the two species were never found living together in the same habitat.

Quedius satoi also resembles Q. muscicola. For a comparison, see the discussion

under the latter species.

38. Quedius (Raphirus) kanyasa Smetana

Figs. 156-164; Map 22

Quedius kanyasa Smetana 1975:338

Quedius lama Coiffait 1982b:278 (syn.nov.)

Quedius dhaulagirensis Coiffait 1982a:80 (syn.nov.)

Description. — Dark reddish-brown with darker abdomen, apex and apical margins of tergites more

or less paler and with head black to almost uniformly piceous-black. Abdomen feebly iridescent; palpi and

antennae testaceous, legs testaceous to rufo-testaceous, middle and posterior tibiae distinctly darkened at

inner margin, occasionally also posterior femora feebly darker. Head rounded, slightly wider than long (ratio

1.20); eyes very large and convex, temples extremely short, considerably shorter than length of eyes seen

from above (ratio 0.16); no additional setiferous punctures between anterior frontal punctures; posterior

frontal puncture situated at posteromedian margin of eye, one puncture between it and posterior margin of

head; surface of head with fine and dense microsculpture of transverse waves becoming irregular toward

clypeus and forming here and there irregular meshes. Antenna slender and moderately long, segment 3

slightly narrower and about as long as segment 2, segments 4 and 5 distinctly longer than wide, following

segments gradually becoming shorter, outer segments about as long as wide, last segment about as long as

two preceding segments combined. Pronotum about as long as wide, widely arcuate basally, evenly convex

and only moderately narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with

two punctures, posterior puncture not reaching level of large lateral puncture; surface of pronotum with

microsculpture of transverse waves slightly denser and finer than those on head. Scutellum with 7-11

punctures. Elytra short, at suture distinctly (ratio 0.70), at sides somewhat (ratio 0.90) shorter than pronotum

at midline; punctation fine and dense, pubescence brownish; surface between punctures without

microsculpture. Wings reduced to very small nonfunctional stumps reaching about two thirds of length of

elytron. Abdomen with tergite 7 (fifth visible) lacking whitish apical seam of palisade setae; punctation of

tergites slightly finer than that of elytra, slightly denser on basal portions of tergites and in general becoming

sparser toward apex of abdomen; same applies to brownish pubescence which does not form patches of paler

and denser hairs on lateral portions of tergites. Front tarsus dilated in both sexes, inconspicuously so in

female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with three

long and strong setae on each side; apical margin with rather deep, obtusely triangular emargination (Figs.

Revision of the Tribes Quediini and Atanygnathini

263

156, 161), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 157-160,

162-164) with median lobe rather strongly narrowed into subacute apex; apical part, when paramere

removed, with distinct median carina forming a small hook in lateral view. Paramere narrow and elongate,

not quite reaching apex of median lobe, subparallel-sided in middle part, apex variably arcuate to broadly

rounded; with four setae at apical margin, both median setae longer than lateral setae, and with a pair of

unequally long setae at each lateral margin near apex; sensory peg setae on underside of paramere forming

two irregular and variable longitudinal rows.

Length 4. 3-5. 8 mm.

Type material. — Quedius kanyasa. The species was described from three

specimens; the male holotype bears the following labels: “Dzunda Khola-Tal b.

Talphi 3000-3500 m” / “Gebiet von Jumla Westnepal, Ig. H. Franz” / “HOLOTYPE

Quedius kanyasa A. Smetana 1975”; the underside of first label bears code “Pa

193”. The female allotype bears the labels: “b. Maharigaon 3000-3500 m” / “Gebiet

von Jumla Westnepal, Ig. H. Franz” / “ALLOTYPE Quedius kanyasa A. Smetana

1975”; the underside of first label bears code “Pa 211-212”. The female paratype

bears same locality labels as the allotype, except code “202-212”. The holotype and

allotype are deposited in the Franz collection, Modling, Austria, the paratype is

deposited in the Canadian National Collection, Ottawa (CNC No. 13966).

Quedius lama. Coiffait (1982b:278) described the species from three specimens

from Nepal. I was able to study the male holotype and one female paratype

deposited in the Naturhistorisches Museum, Basel, Switzerland. They are labelled as

follows: Holotype:“Padmara-Khari Lagna Bumro 3400-2750 m” / “Nepal, 1977

28.V. W. Wittmer” / “TYPE” (red label)/ “Q. (Raphirus) lama Coiffait 1982”.

Paratype: first two labels as in holotype + third label “PARATYPE” (red). Both

holotype and paratype are indistinguishable from the male holotype of Q. kanyasa\

my determination label “Quedius kanyasa Smet. Smetana det. 1983” was attached to

both specimens. I have not seen the male paratype from Lake Rara, deposited in the

collection Coiffait, Museum d’Histoire Naturelle, Paris, France.

Quedius dliaulagirensis Coiffait (1982a:80) described the species from a single

specimen. The male holotype, deposited in the Senckenberg Museum, Frankfurt

a.M., Federal Republic of Germany, is labelled as follows: “siidl. Dhaulagiri

Dhorpatan 3000-3200 m 7.25.V.1973” / “NEPAL-Expedition Jochen Martens”

“TYPE” / “Holo-Type” / “SMF C 15103” (both red labels)/ “Quedius (Raphirus)

dhaulagirensis H. Coiffait 1979”. The specimen agrees externally completely with

the holotype of Q. kanyasa. The aedoeagus is larger and slightly different in a few

details (see Figs. 162-164), but the internal sacs are basically identical (though in

different positions in the two holotypes). Sternite 8 in dhaulagirensis is damaged

in the apical portion (Figs. 161), but there is no reason, why the emargination could

not be as deep as in kanyasa.

I do not see any reason why Q. dhaulagirensis should be retained as a separate

taxon. The name Q. dhaulagirensis is a junior synonym of Q. kanyasa', my

determination label “Quedius kanyasa Smet. Smetana det. 1983” was attached to

this specimen.

Quaest. Ent., 1988, 24 (2)

264

Smetana

Geographical distribution. — Quedius kanyasa is known from western Nepal,

eastward to Ghoropani Pass area (Map 22).

Material studied. — 49 specimens.

NEPAL. Lake Rara, 2920 m, 2. VI. 77, Wittmer (NHMB) 2; Gebiet des Rarasees H. Lranz (HLCC) 4;

Padmara-Khari Lagna-Bumro, 2750-3400 m, 28.V.77, Wittmer (NHMB) 1. Parhat Distr. Ghoropani Pass,

2850 m, 9.x. 83, Smetana and LobI (ASCC) 2; Pun Hill at Ghoropani Pass, 3050-3100 m, 8.X.83, Smetana

and Lbbl (ASCC, BMNH, CNCC, MHNG) 34.

Bionomics. — The holotype of Q. kanyasa was collected in the Dzunda Khola

valley in an alder forest with intermixed spruce by sifting forest floor detritus and

moss near the river banks; both allotype and paratype of Q. kanyasa were found in a

gorge forest near Sinemoro by sifting forest floor detritus. The specimens from Pun

Hill were taken on a pasture by sifting moss and debris under low bushes; the

specimens from Ghoropani Pass were taken by sifting rotting weeds and other debris

at the edges of primitive fields.

Comparisons. — Quedius kanyasa can easily be distinguished among the

brachypterous species of Raphirus with darkened tibiae, in addition to the characters

on the aedoeagus, by the uniform punctation of the abdominal tergites and by the

presence of three strong setae on each side of the male sternite 8 (Fig. 156);

however, quite rarely, the pubescence on the abdominal tergites may have the

tendency to form quite indistinct patches of vaguely denser hairs laterally.

The presence of three long and strong setae on each side of the male sternite 8 is

a rare character among the small species of Raphirus; in addition to Q. kanyasa, it at

present is known to occur only in Q. gaarho and Q. atchala; in the latter species, the

most median pair of setae is distinctly shorter than the two more lateral pairs (see

Figs. 103, 190). The male holotype of Q. dhaulagirensis bears three pairs of strong

setae on the sternite 8; they match those of Q. kanyasa.

39. Quedius (Raphirus) naati spec.nov.

Figs. 165-171; Map 22

Description. — in all characters very similar to Q. kanyasa, but different as follows: colouration less

variable than that of Q. kanyasa, dark reddish-brown specimens with darker abdomen and black head do not

occur in Q. naati', most specimens are piceous to piceous-black with usually darker head and with elytra and

apical margins of abdominal tergites occasionally paler; middle tibiae usually uniformly pale and posterior

tibiae only slightly darkened, rarely almost uniformly pale. Pubescence of abdominal tergites forming

inconspicuous patch of paler and slightly denser pubescence on either lateral portion of each tergite. Size in

general slightly smaller and form more slender.

Male. Lirst four segments of front tarsus distinctly more dilated than in female. Sternite 8 with two long

and strong setae on each side; apical margin with moderately deep, obtusely triangular emargination (Lig.

165), small triangular area before emargination flattened and smooth. Aedoeagus (Ligs. 166-169) similar to

that of Q. kanyasa, but narrower and slightly smaller, paramere almost touching lateral margins of median

lobe at one point; median lobe slightly dilated before narrowed apical portion and, when paramere removed,

with fine median carina forming a small hook in lateral view . Paramere narrow and elongate, not quite

reaching apex of median lobe, arcuate apically; with four setae at apical margin, median setae longer than

lateral setae; sensory peg setae on underside of paramere forming two long and fairly regular longitudinal

rows.

Revision of the Tribes Quediini and Atanygnathini

265

Length 4. 2-5. 4 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL (prov.

Bagmati) Jangtang Ridge NE Barahbise, 3250 m 5.V.81 Ldbl & Smetana”. In the

Smetana collection, Ottawa, Canada.

Paratypes (22); same data as holotype (ASCC, CNCC, MNHG, SMPM) 22.

Geographical distribution. — Quedius naati is at present known only from one

locality in central Nepal (Map 22).

Bionomics. — All specimens of the original series were taken by sifting of moist

moss, debris and low vegetation under bushes in a small valley with a brook.

Comparisons. — Quedius naati can rather easily be distinguished from Q.

kanyasa by the characters given in the key and in the description. The aedoeagus of

Q. naati is very similar to that of Q. kanyasa; however, it again differs in a few

details on the median lobe and on the paramere as well (see the description and Pigs.

157-160, 166-169). Also, both species are apparently allopatric: Q. kanyasa seems

to be restricted to western Nepal, whereas Q. naati is known at present from

east-central Nepal.

Quedius naati also resembles Q. dewar; however, it differs, in addition to the

characters on the aedoeagus, by the smaller size and less robust form and by the

slimmer legs with first four segments of male front tarsus less dilated.

Etymology. — The specific name is the Nepali noun naati (grandson) in

apposition.

40. Quedius (Raphirus) dewar spec.nov.

Figs. 170-174; Map 17

Description. — In all external characters very similar to Q. kanyasa, but different as follows: form

slightly stouter with wider head and pronotum, colouration less variable than that of Q. kanyasa, all

specimens piceous-black, elytra with distinct metallic reflections and abdomen more iridescent. Elytra

slightly shorter, especially at suture, and in general more dilated posteriorly. Pubescence of abdominal

tergites paler and forming inconspicuous patch of paler and slightly denser pubescence on either lateral

portion of each tergite.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side; apical margin with moderately deep, obtusely triangular emargination (Fig.

170). Aedoeagus (Figs. 172-174) elongate, median lobe evenly, almost conically narrowed, with subacute

apex and, when paramere removed, with fine median carina forming a small hook in lateral view. Paramere

narrow and elongate, not quite reaching apex of median lobe, arcuate apically; with two setae at apical

margin, and with a pair of unequally long setae at each lateral margin near apex; sensory peg setae on

underside of paramere forming two long irregular rows.

Length 4. 9-5. 8 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL (Prov.

Bagmati) Mere Dara 3200 m, 8. IV. 81 Lobl & Smetana”. In the Smetana collection,

Ottawa, Canada.

Paratypes (5); same data as holotype (MHNG) 1; same data as holotype but

3100-3300 m and date 7.IV.81 (CNCC) 1; Malemchi, 2800 m, 16.IV.81, Lobl &

Smetana (ASCC) 1; Dhorpatan, 3000 m, 20.V.73, J. Martens (SMFM) 1;

Qiiaest. Ent., 1988, 24 (2)

266

Smetana

Kathmandu Distr., Siwapuri Dara, 2520 m, 1.V.85, A. Smetana (ASCC) 1.

Geographical distribution. — Quedius dewar is at present known from western

and central Nepal (Map 17).

Bionomics. — The specimens from Mere Dara were taken on northern slopes in

Rhododendron growths by sifting wet fallen leaves, debris and moss. The specimen

from Malemchi was taken by sifting soaking wet debris, fallen leaves and moss

among large rocks permanently sprinkled by water from a waterfall in a shady

ravine.

Comparisons. — Quedius dewar can easily be distinguished from Q. kanyasa by

the characters given in the key and in the description. It differs from Q. naati, in

addition to the differences in the shape of the aedoeagus, by the larger size and more

robust form, by the stronger legs with four first segments of the male front tarsus

much more dilated than those of Q. naati. The shape of the median lobe of the

aedoeagus of Q. dewar is quite characteristic (Figs. 172, 173).

Etymology. — The specific name is the Nepali noun dewar (husband’s younger

brother) in apposition.

41. Quedius (Raphirus) tikta Smetana

Figs. 175-179, 388-395, 421; Map 23

Quedius tikta Smetana 1975:339

Description. — Pale brown to piceous, with darker head; males paler: pale brown with pronotum

usually more or less appreciably darkened in middle and with piceous head with anterior portion

reddish-brown to brown; females darker: more or less piceous with elytra sometimes feebly paler, pronotum

uniformly piceous and head uniformly piceous-black. Abdomen iridescent; palpi, antennae and legs

uniformly testaceous to testaceobrunneous. Head rounded, noticeably wider than long (ratio 1.24); eyes very

large and convex, temples extremely short, considerably shorter than length of eyes seen from above (ratio

0.13); no additional setiferous punctures between anterior frontal punctures; posterior frontal puncture

situated at posteromedian margin of eye, one puncture between it and posterior margin of head (Fig. 421);

surface of head with dense and very fine microsculpture of irregular transverse waves with scattered

longitudinal connections becoming gradually more numerous anteriorly, front part of head and especially

clypeus therefore covered with meshed microsculpture. Antenna slender, segment 3 slightly narrower and

about equally long as segment 2 (Fig. 388), segments 4 and 5 very distinctly longer than wide, segments 6-8

longer than wide, gradually becoming shorter and wider, segments 9-10 slightly to feebly longer than wide,

last segment about as long as two preceding segments combined. Pronotum about as long as wide, widely

arcuate basally, evenly convex and only slightly narrowed anteriorly; dorsal rows each with three punctures;

sublateral rows each with two or three punctures, posterior puncture not or about reaching (when three

punctures present) level of large lateral puncture; surface of pronotum with microsculpture of transverse

waves distinctly denser and somewhat finer than that on head. Scutellum puncate on apical half (Fig. 390),

number of punctures variable. Elytra short, at suture distinctly (ratio 0.76), at sides somewhat (ratio 0.88)

shorter than pronotum at midline; punctation dense and fine, pubescence brownish; surface between

punctures without microsculpture. Wings reduced to very small narrow nonfunctional stumps reaching about

two thirds of length of elytron. Abdomen with tergite 7 (fifth visible) lacking whitish apical seam of palisade

setae (Fig. 391); punctation of tergites about same as that on elytra, slightly denser and finer on bases of

tergites and in general becoming sparser toward apex of abdomen; same applies to brownish pubescence,

forming distinct patch of denser yellowish hairs on either lateral portion of each tergite. Front tarsus dilated

in both sexes, inconspicuously so in female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with two long

and strong setae on each side; apical margin with moderately deep, obtusely triangular emargination (Fig.

Revision of the Tribes Quediini and Atanygnathini

267

175), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 176-179) narrow

and elongate, median lobe strongly narrowed, subacute to acute apically; apical part, when paramere

removed, with fine longitudinal median carina forming a small hook in lateral view. Paramere elongate,

relatively wide and covering apical portion of median lobe except for very apex, obtusely rounded to

subemarginate apically, with four very small setae at apical margin, median setae longer and stronger than

lateral setae, and a pair of similar setae close to apex; minute setae, present on lateral margins in most

species of Raphirus, are relatively long and numerous, especially on anterior portions of lateral margins;

sensory peg setae on underside of paramere forming two relatively short longitudinal rows along midline of

paramere.

Length 4. 0-5. 5 mm.

Type material. — The species was described from three specimens, all bearing

identical locality labels: “Zw. Tare-Pati u. Gosaikund” / “Zentral-Nepal Sept. -Okt.

1971 Ig. H. Franz”, the underside of the first label bears code “Pa 163”. The male

holotype and allotype are deposited in the Franz Collection, Modling, Austria; the

holotype bears red label “HOLOTYPE Quedius tikta A. Smetana 1975”. The female

paratype is deposited in the Canadian National Collection, Ottawa (CNC No.

13967).

Geographical distribution. — Quedius tikta is known at present only from the

main Himalayan range in Central Nepal (Map 23).

Material studied. — 304 specimens.

NEPAL. Kathmandu Distr. Siwapuri Dara, 2450 m, 30.IV. 85, A. Smetana (ASCC) 1. Nuwakot Distr.

near Mere Dara, 3000 m, 7.IV.81, Lbbl and Smetana (ASCC) 1; Mere Dara, 3200 m, 8.IV.81, Lbbl and

Smetana (ASCC, BMNH, CNCC, MHNG) 25; below Thare Pati, 3300-3500 m, 9-12.IV.81, Lobl and

Smetana (ASCC, BMNH, CNCC, MHNG) 90; between Ghopte and Thare Pati, 3100-3250, 23.26.IV.85, A.

Smetana (ASCC, BMNH, CNCC, MHNG) 51;. Malemchi, 2800 m, 16.IV.81, Lobl and Smetana (ASCC) I;

Yangri ridge, 4150-4800 m, 21-24.IV.81, Lobl and Smetana (ASCC, BMNH, CNCC, MHNG) 102;

Thodung, 3200 m, 3-9. IV. 73, J. Martens (SBMF) 1. Rasiiwa Distr. north slope above Syabru, 3600-3800 m,

17. 19.IV.86, A. Smetana (ASCC,BMNH, CNCC) 28.

Bionomics. — Quedius tikta seems to be the most abundant brachypterous

species in central Nepal. It occurs from forest habitats at about 2800 m all the way

to the alpine zone close to 5000 m. Most specimens were collected by sifting moist

moss (on large rocks or at their bases, or on fallen trees), leaf litter and forest floor

debris in forest habitats, or higher up by sifting leaf litter, other debris and moss

under bushes, and by sifting thick layers of moss, lichens and other low vegetation

in the alpine zone.

Comparisons. — Quedius tikta can easily be recognized among the flightless

small Raphirus species, in addition to the characters on the aedoeagus, by the

distinct sexual dimorphism in the colouration of the body, particularly by the

bicoloured head in the male (see the desciption). This phenomenon is quite

conspicuous, but not unique; it occurs also in Q. durgaa and in some specimens of

Q. tonglu.

The three specimens of the original series had dirty and greasy abdomens; the

patches of denser paler pubescence on lateral portions of tergites were therefore

obscured and were not mentioned in the original desciption.

Quaest. Ent., 1988, 24 (2)

268

Smetana

42. Qiiedius (Raphirus) tonglii spec. now

Figs. 180-184; Map 16

Description. — in all characters, including general habitus, colouration and pubescence very similar

to Q. tikta, but different as follows: colouration (both sexes) same as in paler males of Q. tikta, abdomen

particularly pale in most specimens, but head in males without appreciably paler anterior portion. Head in

general smaller and narrower (ratio width: length=1.15) and with less convex eyes. Pronotum in general

somewhat narrower and elytra indistinctly shorter. Punctation and pubescence of abdominal tergites about

equal to that of average specimens of Q. tikta.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 (Fig. 180) not

appreciably different from that of Q. tikta. Aedoeagus (Figs. 181-184) rather small; apical portion of median

lobe, when paramere removed, rather short and wide and with wide, short median carina. Paramere not

reaching apex of median lobe nor lateral margins of median lobe, moderately long and lancet-shaped; four

setae at apical margin, middle two setae much longer than two lateral setae, two short setae at each lateral

margin below apex; sensory peg setae on underside of paramere forming moderately long, irregular lateral

rows.

Length 4. 2-5. 8 mm.

Type material. — Holotype (male) and allotype (female): “INDIA W. Bengal

Darjeeling distr. Tonglu 2700 m 16.X.78 Besuchet-Lobl”. Both holotype and

allotype in the Museum d’histoire naturelle, Geneve, Switzerland.

Paratypes (119): same data as holotype (ASCC, BMNH, MHNG) 19; same data,

but elevation 3100 m (ASCC, CNCC, MHNG) 62; Darjeeling distr.. Tiger Hill,

2200-2300 m, 13.X.78, Besuchet-Lobl (ASCC, BMNH,CNCC, MHNG) 30; same

data but elevation 2500-2600 m and date 18.X.78 (MHNG) 8.

Geographical distribution. — Quedius tonglu is at present known from Tonglu

and Tiger Hill in the Darjeeling area in West Bengal (Map 16).

Bionomics. — Quedius tonglu was collected at elevations from 2200 to 3100 m;

at Tonglu on the northern slope by sifting forest floor litter and fallen leaves under

old trees and higher up (3100 m) under bushes. At Tiger Hill, the species was taken

by sifting forest floor litter and moss on a southern slope.

Comparisons. — Quedius tonglu can be distinguished from Q. tikta by the

characters given in the key and in the description. In some specimens (both sexes)

the head is barely noticeably paler around each antennal insertion and in rare male

specimens the pale colouration is distinct and developed in the same way as in the

males of Q. tikta.

Quedius tonglu also resembles Q. pharak\ however, the latter species differs, in

addition to the characters on the aedoeagus, by the slightly more robust form, by the

wider pronotum, by the more sparsely punctate elytra and by the darker coloured

abdomen with punctation and pubescence of tergites usually sparser and with lateral

patches of pale hairs less distinct.

Etymology. — The specific name is the geographic name Tonglu (type locality)

in apposition.

Revision of the Tribes Quediini and Atanygnathini

269

43. Quedius (Raphirus) pharak spec.nov.

Figs. 185-189; Map 23

Description. — in all characters, including habitus, colouration and pubescence very similar to Q.

tikta, but different as follows: colouration same as in females of Q. tikta, i.e. head uniformly dark in both

sexes; abdomen in most specimens piceous-black with apical margins of tergites paler. Microsculpture on

head and pronotum slightly coarser; head in general less transverse with less convex eyes; pronotum in

general wider and stouter. Punctation and pubescence of elytra and abdominal tergites sparser, lateral

patches of pale hairs less distinct. Size in general slightly larger.

Male. First four segments of front tarsus distinctly more dilated than in female. Sternite 8, including

apical emargination, same as that of Q. tikta (see Figs. 175, 185). Aedoeagus (Figs. 186-189) very similar to

that of Q. tikta: however, median lobe, when paramere removed, with stronger and longer median carina

forming a stronger hook in lateral view. Paramere nanower, reaching to about lateral margins of apical

portion of median lobe, rows of sensory peg setae on its underside long.

Length 4. 2-5. 8 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL, Khandbari

District” / “above Tashigaon 3500 m 6.IV.1982 A. & Z. Smetana”. In the Smetana

Collection, Ottawa, Canada.

Paratypes (11): same data as holotype (ASCC, CNCC) 2; same data, but

elevation 3550 m (CNCC) 1; same data but elevation 3600 m (ASCC) 2; Khandbari

Distr., “Bakan” W of Tashigaon, 3200 m, 5. IV. 82, A. & Z. Smetana (ASCC) 2;

Dobate Ridge NE Barahbise, 3000 m, 7.V.8I, Ldbl & Smetana (ASCC, MHNG) 4.

Geographical distribution. — Quedius pharak is at present known from two

areas in east-central and eastern Nepal (Map 23).

Bionomics. — Quedius pharak was collected at elevations from 3000 to 3500 m

in forest habitats. All specimens from near Barahbise were taken by sifting moist

moss on large rocks in an old deciduous forest with dominant old oak trees. The

specimens from above Tashigaon were collected by sifting deep layers of moist

needles (moldy at some spots) and other debris under a huge Abies tree; by sifting

deep layers of rotting and moldy leaves under Rhododendron bushes; by sifting wet

leaves and other debris under a small forest waterfall, and by treading wet grass in a

depression (clearing) in a forest.

Comparisons. — Quedius pharak is very similar to Q. tikta; however, it can

rather easily be distinguished from it by the characters given in the key and in the

description. The species were never found together. Quedius tikta seems to be more

western and does not reach eastward even the Barahbise area; Q. pharak, on the

other hand, seems to be an eastern species, distributed from Barahbise area

eastwards to the Arun River valley and perhaps even further east.

For comparison with Q. tonglu, see the discussion there.

Two specimens of the original series have the hind tibiae feebly darkened

medially.

Etymology. — The specific name is the Nepali adjective pharak (different).

Quaest. Ent., 1988, 24 (2)

270

Smetana

44. Quedius (Raphirus) atchala Smetana

Figs. 190-194; Map 13

Quedius atchala Smetana 1975:341

Description . — Piceous-black to black, head and pronotum with feebly, elytra with distinct metallic

reflections, abdomen slightly iridescent; palpi, antennae and legs testaceous, middle and posterior tibiae

distinctly darkened at inner margin. Head rounded, slightly wider than long (ratio 1.15), eyes very large and

convex, temples extremely short, considerably shorter than length of eyes seen from above (ratio 0.15); no

additional setiferous punctures between anterior frontal punctures; posterior frontal puncture situated at

posteromedian margin of eye, one puncture between it and posterior margin of head; surface of head with

fine and moderately dense microsculpture of irregular transverse waves becoming confused anteriorly,

especially on clypeus. Antenna rather short, segment 3 narrower but about equally long as segment 2,

segments 4 and 5 slightly longer than wide, following segments gradually becoming shorter and wider,

segments 9 and 10 about as long as wide, last segment feebly shorter than two preceding segments

combined. Pronotum nanow, about as long as wide to scarcely longer than wide, widely arcuate basally,

evenly convex, hardly narrowed anteriorly, with lateral margins often almost parallel-sided in posterior half;

dorsal rows each with three punctures; sublateral rows each with two or three punctures, last puncture not or

about reaching level of large lateral puncture (if three punctures present); surface of pronotum with

microsculpture similar to that on head but slightly coarser. Scutellum with only a few punctures (2-8,

usually 3-5). Elytra short, at suture distinctly (ratio 0.75), at sides slightly (ratio 0.90) shorter than pronotum

at midline; punctation rather sparse, intervals between punctures along transverse axis mostly several times

larger than diameters of punctures; pubescence pale brownish, surface between punctures without

microsculpture but with some extremely fine microscopic irregularities. Wings reduced to small

nonfunctional stumps reaching about two thirds of length of elytron. Abdomen with tergite 7 (fifth visible)

lacking whitish apical seam of palisade setae; punctation of tergites slightly finer than that on elytra, slightly

denser on basal portions of tergites and gradually becoming slightly sparser toward apex of abdomen; same

applies to light golden-brownish pubescence, forming inconspicuous but distinct patch of denser and slightly

paler hairs on either lateral portion of each tergite. Front tarsus dilated in both sexes, inconspicuously so in

female.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with three

long and strong setae on each side; most median pair of setae distinctly shorter than two more lateral pairs;

apical margin with rather shallow and wide, obtusely triangular to almost arcuate emargination (Fig. 190),

small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 191-194) with median

lobe subparallel-sided to vaguely constricted in middle portion, with subacute apex; apical portion, when

paramere removed, with fine median carina forming a fine tooth in lateral view. Paramere not quite reaching

apex of median lobe, almost parallel-sided to feebly nan'owed, arcuate apically; with four setae at apical

margin, both median setae longer than lateral setae, and with two or three unequally long setae at each lateral

margin near apex; sensory peg setae on underside of paramere numerous, forming two irregular longitudinal

rows.

Fength 3. 8-4. 6 mm.

Type material. — The species was described from a single specimen from the

Gosaikund Lakes area in Central Nepal (see Smetana 1975:341). The holotype is

deposited in the Franz collection, Modling, Austria.

Geographical distribution. — Quidius atchala is known from a few localities in

Central Nepal (Map 13).

Material studied. — 68 specimens.

NEPAF. Nuwakot Distr. below Thare Pati, 3500 m, 12.IV.81, Fobl and Smetana (ASCC, CNCC) 14;

Yangri Ridge, Yangri, 4150 m, 21 and 24. IV. 81, Fobl and Smetana (ASCC, BMNH, MHNG) 14; between

Ghopte and Thare Pati, 3250 m, 23. IV. 85, A. Smetana (ASCC) 1. Rasuwa Distr. north slope above Syabru,

3600-3800 m, 17-1 9.1 V.85, A. Smetana (ASCC, CNCC, MCZC) 34; Gosaikund, Farabina Pass, 4000^100

m, 20. IV. 85, A. Smetana (ASCC) 1; Fangtang Khola Valley, 2.5 km E Syabru, 1730 m, 14. IV. 85, A.

Smetana (ASCC) 2.

Revision of the Tribes Quediini and Atanygnathini

271

Bionomics. — Quedius atchala occurs at elevations above 3000 m, from the

upper forest zone to the alpine zone. Specimens were sifted from moist moss at

bases of rocks under Acer sp. trees, from forest floor litter and moss under old Abies

trees in a sheltered ravine, from wet moss, dead grass and other debris on a seepage

slope in a mixed Ahies-Rhododendron forest, and in the alpine zone from moss,

lichens and debris under low vegetation. The specimens taken near Syabru at

relatively low elevation come from soaking wet moss in a small waterfall.

Comparisons. — Quedius atchala can easily be distinguished from the other

small brachypterous species of Raphirus with patches of denser pubescence on

abdominal tergites and with darkened posterior tibiae by its small size in

combination with the narrow and rather parallel-sided pronotum and the rather

sparse punctation of elytra.

One of the specimens studied has no punctures on scutellum.

Taruni Group

The single species belonging to this group is characterized by the following

combination of characters; size small; head with additional punctures on each side

along median margin of eye between anterior and posterior frontal punctures, and

with two punctures between posterior frontal puncture and posterior margin of head;

eyes large but not taking almost entire sides of head; segment 3 of antenna as long

as segment 2; pronotum with each of dorsal rows with four punctures and with

additional, characteristically located punctures (see the description of Q. taruni for

details); scutellum impunctate; elytra with simple punctation.

45. Quedius (Raphirus) taruni spec.nov.

Map 24

Description. — Piceous black, elytra brownish-piceous with indistinctly paler humeri and with

apical margin paler, abdomen feebly iridescent; palpi piceous, antennae brownish testaceous with two basal

segments darker, legs piceous with darkened tibiae and paler tarsi. Head round, slightly wider than long

(ratio 1.14); eyes large and convex, tempora much shorter than length of eyes seen from above (ratio 0.28);

no additional punctures between anterior frontal punctures; however, with two or three additional setiferous

punctures on each side along medial margin of eye between anterior and posterior frontal punctures;

posterior frontal puncture situated close to posteromedian margin of eye, separated from it by distance no

larger than diameter of puncture, two punctures between it and posterior margin of head; temporal puncture

almost touching posterior margin of eye, one additional puncture at posterior margin between it and

posterior frontal puncture; surface of head with moderately dense microsculpture of extremely fine, often

rudimentary transverse waves. Antenna short, segment 2 shorter than segment 1, segment 3 about equally

long but somewhat thinner than segment 2, segments 4 and 5 longer than wide, segment 5 somewhat shorter

than segment 4, segment 6 as long as wide, segments 7-10 wider than long, gradually becoming wider, last

segment about as long as two preceding segments combined. Pronotum rather narrow, about as long as wide,

broadly rounded basally, lateral margins somewhat arcuate and indistinctly narrowed anteriorly; dorsal rows

each with four punctures; one of punctures at posterior margin of pronotum situated somewhat away from

posterior margin, forming with four punctures of dorsal row a row of five punctures extended from anterior

to posterior margin of pronotum; sublateral rows each with three or four punctures, posterior puncture

Quaest. Ent., 1988, 24 (2)

272

Smetana

situated behind level of large lateral puncture, one additional puncture between dorsal and sublateral row on

each side; surface of pronotum with microsculpture similar to that on head. Scutellum large, impunctate,

with extremely fine microsculpture of rudimentary transverse waves. Elytra long, at base indistinctly

narrower than pronotum and slightly widened posteriorly; at suture slightly longer (ratio 1.11), at sides

distinctly longer (ratio 1.29) than pronotum at midline; punctation fine, superficial and sparse, interspaces

between punctures considerably larger than diameters of punctures, pubescence fine and short, dark; surface

between punctures without microsculpture but with numeropus microscopic irregularities. Wings fully

developed. Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of palisade setae;

punctation of tergites finer and somewhat denser than that of elytra, slightly denser on basal portion of each

tergite and in general becoming gradually sparser toward apex of abdomen; pubescence fine, dark. First four

segments of front tarsus slightly dilated.

Male. Unknown.

Length 5.0-6. 1 mm.

Type material. — Holotype (female): “NEPAL Rasuwa Dis. north slope above

Syabru 3800 m 18. IV. 85 A. Smetana”. In the Smetana collection, Ottawa, Canada.

Paratypes (20): Nepal: same data as holotype but date 17. IV. 85 (ASCC, BMNH)

3. Rasuwa Dis.: Gosaikund, Larabina Pass, 4000^100 m, 20. IV. 85, A. Smetana

(ASCC) 1; Gosaikund Lakes, 4200-4300 m, 20.IV.85, A. Smetana (ASCC, CNCC,

MHNG) 8; same, 4450 m, 21. IV. 85, A. Smetana (ASCC) 2. “Himalaya-Expedition

Prof. Dr. H. Janetschek 1961 nach Nepal” / “loc. 213 III” (IZUI, NHMW) 6. (See

the discussion for details on the material collected by Dr. Janetschek).

Geographical distribution. — Qiiedius taruni is known from several localities in

central Nepal (Map 24).

Bionomics. — The specimens collected by myself were taken by sifting fallen

leaves and other debris under various low bushes in subalpine zone (north slope

above Syabru) and by sifting moss, lichens and debris under low vegetation in the

alpine zone (Larabina Pass and Gosaikund Lakes). The specimens collected by

Janetschek were taken from “Moos-und Llechtenheiden mit Zwerg-rhododendron”

(Scheerpeltz 1976a:43 - see the discussion).

Comparisons. — Quedius taruni is a conspicuous species due to the presence of

the additional punctures on the head between the anterior and posterior frontal

punctures and due to the chaetotaxy of the pronotum, particularly by the presence of

four punctures in each of the dorsal rows. It is the only Quedius species known to

me to have this combination of the chaetotaxy on the head and pronotum; it shares

the character of the additional punctures between the anterior and posterior frontal

punctures on the head with the species of the genera Indoquedius and Bolitogyrus,

with some species of the genus Heterothops and with some Nearctic species of the

subgenus Raphirus (Q. prostans Horn 1878 and Q. seriatus Horn 1878).

The locality “213 III” on the specimens collected by Janetschek reads as follows:

“Raldurje, SE above base camp Yaral bei Tangpoche, cca 4400 m, 9.V.1961”.

Although all 21 known specimens of this species are females, which is quite

unusual in the genus Quedius, I do no believe that this species is parthenogenetic. I

expect that extensive collecting in suitable habitats, perhaps at a different time of the

year (all available specimens were collected in spring), will produce males.

Revision of the Tribes Quediini and Atanygnathini

273

This is the species published by Scheerpeltz (1976a:43) under the name of

Qiiedius ripicola.

Etymology. — The specific name is the Nepali noun taruni (young woman). It

refers to the fact that I only had female specimens for study.

Durgaa Group

The single species of this group is characterized by the following combination of

characters: size large; both head and pronotum without microscopic punctures and

without additional setiferous punctures, eyes very large, taking almost entire sides of

head; antennal segment 3 longer than segment 2; scutellum punctate; elytra with

double punctation (see the description of Q. durgaa for details) but without

longitudinal rows of larger punctures.

46. Quedius (Raphirus) durgaa spec. now

Figs. 195-198; Map 18

Description. — Piceous-black with indefinitely paler pronotum, head gradually becoming

rufo-testaceous anteriorly, labrum rufo-testaceous, apical margins of abdominal tergites indefinitely paler;

abdomen slightly iridescent; mouthparts testaceous; antennae testaceous, gradually becoming paler toward

apex; legs brunneotestaceous with piceous femora. Head rounded, wider than long (ratio 1.21); eyes large

and convex, tempora considerably shorter than length of eyes seen from above (ratio 0.25); no additional

setiferous punctures between anterior frontal punctures; posterior frontal puncture situated very close to

posteromedian margin of eye, separated from it by distance about equal to diameter of puncture, one

puncture between it and posterior margin of head; temporal puncture almost touching posterior margin of

eye; tempora impunctate; surface of head with extremely fine microsculpture of irregular, rudimentary

waves. Antenna moderately long, hardly incrassate toward apex, segment 3 somewhat longer than segment

2, following segments longer than wide, gradually becoming shorter, last segment slightly shorter than two

preceding segments combined. Pronotum slightly wider than long (ratio 1.15), broadly rounded basally,

distinctly narrowed anteriorly; dorsal rows each with three punctures; sublateral rows each with two

punctures, posterior puncture situated at about level of large lateral puncture, one distinct impression in

continuation of each sublateral row, situated well behind level of large lateral puncture; large lateral

puncture situated rather far from lateral pronotal margin, separated from it by distance equal to about three

diameters of pucture; surface of pronotum with microsculpture of dense and extremely fine, rudimentary

waves. Scutellum with a few indistinct punctures. Elytra rather short, at base narrower than pronotum at

widest point, at suture, slightly shorter (ratio 0.85), at sides as long as pronotum at midline; punctation

coarse and rather irregularly distributed, consisting of intermixed coarse and fine punctures; pubescence

absent, surface between punctures without microsculpture. Wings fully developed. Abdomen with tergite 7

(fifth visible) with distinct whitish apical seam of palisade setae; punctation of abdominal tergites finer than

coarse elytral punctures, slightly denser at base of each tergite and gradually becoming slightly sparser

toward apex of abdomen, pubescence piceous, short; surface between punctures without appreciable

microsculpture.

Male. First four segments of front tarsus strongly dilated. Stemite 8 with moderately wide and deep

triangular emargination (Fig. 195), triangular area before emargination flattened and smooth. Aedoeagus

(Figs. 196, 198) large, median lobe strongly narrowed into rather long and naiTow apical portion. Internal

sac with four characteristic elongate structures, two middle ones with long apical spines. Paramere long and

wide, covering entire median lobe except for very tip, gradually narrowed into arcuate apex; six unequally

long setae at apical margin and two very long and strong setae slightly below them; underside of paramere

with two long, irregular, lateral rows of sensory peg setae gradually becoming larger away from apex of

paramere.

Quaest. Ent., 1988, 24 (2)

274

Smetana

Length 10.0 mm.

Type material. — Holotype (male): “India W. Bengal Darjeeling dist. 13 km. N.

Ghoom 1500. 15.X.78 Besuchet Lobl. In the Museum d’Histoire Naturelle de

Geneve, Geneve, Switzerland.

Geographical distribution. — Quedius durgaa is at present known only from the

type locality in the Darjeeling district (Map 18).

Bionomics. — The holotype was taken by sifting moss and fallen leaves under

bushes.

Comparisons. — Quedius durgaa is a distinctive species, well characterized, in

addition to the shape of the aedoeagus, by the double punctation of the elytra, in

combination with the large size, and the colouration of the head.

Etymology. — The specific name is the name of one of the Hindu Goddesses.

Anomalus Group

The single species of this group is characterized by the following combination of

characters: size large; head and pronotum with microscopic punctures but without

additional setiferous punctures; eyes very large, taking almost entire sides of head;

antennal segment 3 somewhat longer than segment 2; scutellum impunctate but with

fine transverse rugae; elytra each with three longitudinal rows of larger punctures.

47. Quedius (Raphirus) anomalus Cameron

Figs. 199-202; Map 20

Quedius anomalus C&memn 1926:370; 1932:295

Description. — Black, with palpi, antennae towared apex and legs piceous. Head rather small,

rounded, slightly wider than long (ratio 1.21), posterior angles rounded; eyes very large and moderately

convex, tempora much shorter than length of eyes seen from above (ratio 0.25); no additional setiferous

punctures between anterior frontal punctures; posterior frontal puncture situated very close to posteromedian

margin of eye, separated from it by distance smaller than diameter of puncture, one additional setiferous

puncture between it and posterior margin of head; surface of head with fairly dense punctation of extremely

fine, microscopic punctures becoming gradually sparser toward vertex, and with extremely fine

microsculpture of rudimentary transverse waves. Antenna moderately long, segment 3 longer than segment

2, following segments all longer than wide, gradually becoming shorter, penultimate segment slightly longer

than wide, last segment shorter than two preceding segments combined. Pronotum rather wide, slightly

wider than long (ratio 1.13), widely arcuate basally and strongly narrowed anteriorly; dorsal rows each with

three punctures, sublateral rows with two punctures, both situated close to anterior margin of pronotum,

much in front of level of large lateral puncture; surface of pronotum with microscopic punctures and

microsculpture similar to those on head; however, microscopic punctures somewhat finer and sparser and

microsculpture still finer and less apparent than on head. Scutellum large, impunctate, but with some very

fine transverse rugae, simulating punctation. Elytra long, at base somewhat narrower than pronotum at

widest point; at suture as long as, at sides longer than pronotum at midline (ratio 1.16); punctation fairly

deep, moderately coarse and dense, each elytron with three longitudinal rows of larger punctures, one along

suture, one rather irregular lateral row and a discal row between them reduced to two punctures on posterior

half of elytron; surface between punctures without microsculpture. Wings fully developed. Abdomen with

tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; punctation on first abdominal tergites

about equal to that on elytra, coarser and denser on basal portion, gradually becoming much finer toward

apex of abdomen, fifth visible tergite quite finely punctate; pubescence dark.

Revision of the Tribes Quediini and Atanygnathini

275

Male. First four segments of front tarsus strongly dilated. Sternite 8 with wide and deep, obtusely

triangular emargination in middle of apical margin (Fig. 199), small triangular area before emargination

Fattened and smooth. Aedoeagus (Figs. 200-202) very large; median lobe entirely covered by paramere,

strongly narrowed into rather acute apical part; paramere large, long and wide, rather strongly narrrowed,

apical margin obtusely arcuate; three setae at each lateral margin just below apex; sensory peg setae on

underside of paramere moderately numerous, forming two irregular longitudinal rows.

Length 9.4 mm (abdomen slightly extended).

Type material. — The collection of the British Museum (Natural History),

London, contains one male specimen under the name Q. anomalus. It is labelled as

follows: “Type” (round label with red margin)/ “Dhobi Ghat, Mussoorie. Dr.

Cameron. 14. IV. 22” / “TYPE Quedius anomalus Dr. Cameron” / “M. Cameron.

Bequest. B.M. 1955-147”.

The specimen was dissected and the 8th sternite, genital segment and aedoeagus

were mounted in Canada Balsam. The specimen is hereby designated as the

lectotype of Q. anomalus', the label “Lectotype Quedius anomalus Cameron

Smetana des. 1984” has been attached to it.

Geograplueal distribution. — Quedius anomalus is at present known only from

the type locality in Uttar Pradesh (Map 20).

Material studied. — The lectotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Quedius anomalus is a distinctive species; it can readily be

recognized by the following combination of characters: head with very large but

only moderately convex eyes, surface of head and pronotum with microscopic

punctures; pronotum strongly narrowed in front; scutellum impunctate but with

some very fine transverse rugae; elytra long, with longitudinal rows of coarser

punctures in addition to usual punctation; abdominal punctation moderately coarse

on front tergites but becoming much finer toward apex. Quedius anomalus cannot

be confused with any other species occurring in the Himalayan area.

Himalayicus Group

This species group is characterized by the following combination of characters:

size large; both head and pronotum without microscopic punctures and without

additional setiferous punctures; eyes very large, taking almost entire sides of head;

antennal segment 3 distinctly longer than segment 2; scutellum punctate; elytra with

simple punctation; first four segments of front tarsus strongly dilated in both sexes.

The group contains five species in the Himalayan region; they all resemble in

general habitus the species of the subgenus Quedius (not represented by any species

in the Himalaya). However, they differ immediately by the bilobed labrum and by

the very large and convex eyes taking most of the lateral portions of the head.

Quaest. Ent., 1988, 24 (2)

276

Smetana

Map 17. Distribution records for: Quedius vadhu ( • ); 0. sundar ( A ); and Q. dewar { ■ ).

277

Revision of the Tribes Quediini and Atanygnathini

Qiiaest. Ent., 1988, 24 (2)

Map 18. Distribution records for: Q. daksiimensis (% )\Q. paschim ( A ); durgaa ( ■ ); and Q. aweipilis ( ^ ).

278

Smetana

Map 19. Distribution records for: Quedius aureivenths.

279

Revision of the Tribes Quediini and Atanygnathini

Quaest. Ent., 1988, 24 (2)

Map 20. Distribution records for: Qiiedius miiscicola ( • ); and Q. anomalus ( A ).

280

Smetana

Map 21. Distribution records for: Qiiedius gaarho (% )\Q. bhari ( A); Q. eklai ( ■ ); and Q. iidagra ( ^ ).

Far Western

Revision of the Tribes Quediini and Atanygnathini 28 1

Quaest. Ent., 1988, 24 (2)

Map 22. Distribution records for: Quedius satoi (%)\Q. kanyasa { A ); and Q. naati ( ■ ).

282

Smetana

Map 23. Distribution records for: Quedius tikta ( • ); and Q. pharak ( A ).

Revision of the Tribes Quediini and Atanygnathini 283

Quaest. Ent., 1988, 24 (2)

Map 24. Distribution records for: Quediiis tanmi ( • ); and Q. hariyo ( A ).

284

Smetana

48. Quedius (Raphirus) assamensis Cameron

Figs. 203-206: Map 15

Quedius assamensis Cameron 1932:293

Description. — Piceous-black to black, abdomen slightly iridescent, apical margins of tergites and

apex of abdomen occasionally indistinctly paler; both labial and maxillary palpi brunneotestaceous, antennae

piceous with first two segments and most of third segment brunneotestaceous, first segment occasionally

partially darkened; legs brunneous with paler tarsi, coxae, femora (front femora usually indistinctly) except

for apices, and medial portions of tibiae darkened to almost black. Head rounded, wider than long (ratio

1.33); eyes large and convex, tempora considerably shorter than length of eyes seen from above (ratio 0.21 );

no additional punctures between anterior frontal punctures; posterior frontal puncture situated very close to

posteromedian margin of eye, separated from it by distance no larger than diameter of puncture, one

additional setiferous puncture between it and posterior margin of head; temporal puncture separated from

posterior margin of eye by distance equal to diameter of puncture; surface with fine and dense

microsculpture of irregular transverse waves with numerous longitudinal junctions and therefore almost

meshed here and there and gradually becoming more or less meshed on central portion of clypeus. Antenna

rather short, segment 3 longer than segment 2, following segments longer than wide, gradually becoming

shorter and slightly wider, segments 9 and 10 feebly longer than wide to almost as long as wide, last segment

shorter than two preceding segments combined. Pronotum feebly wider than long (ratio 1.09), widely

rounded basally and distinctly narrowed anteriorly, evenly transversely convex; dorsal rows each with three

fine punctures; sublateral rows with two punctures, posterior puncture situated distinctly before level of large

lateral puncture, sometimes sublateral rows with third puncture (unilaterally or bilaterally) situated at about

level of large lateral puncture; surface of pronotum with very fine and dense microsculpture of transverse

waves, in general finer and denser than that on head. Scutellum punctate. Elytra rather short, at base slightly

narrower than pronotum at widest point, at suture slightly shorter (ratio 0.83), at sides about as long as

pronotum at midline; punctation dense and moderately coarse, interspaces between punctures about equal to

diameters of punctures; pubescence black. Wings fully developed. Abdomen with tergite 7 (fifth visible)

bearing distinct whitish apical seam of palisade setae; punctation finer and denser than that of elytra,

becoming slightly sparser toward apical margin of each tergite and in general becoming less dense toward

apex of abdomen; pubescence black, dense. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus distinctly more dilated than in female. Apical margin of sternite

6 inconspicuously concave in middle of apical margin; apical margin of sternite 7 broadly and shallowly,

arcuately emarginate apically, slightly flattened and smooth before emargination; apical margin of sternite 8

with wide, moderately deep obtusely triangular emargination, emargination with cluster of darker setae at

each lateral comer (Fig. 203), without smooth area before emargination. Aedoeagus (Figs. 204-206) very

large and voluminous; median lobe wide, suddenly narrowed into obtusely rounded apical portion, with two

lateral sclerites; paramere very large and wide, covering most of median lobe, evenly arcuately narrowed,

with apex arcuate; four apical setae, inner pair much larger than lateral pair, two long and strong setae at

each lateral margin way below apex of paramere; underside of paramere with rather long, irregular row of

sensory peg setae along each lateral margin.

Length 8. 5-9. 6 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains five specimens under the name Q. assamensis-, but probably only

the first male represents the original material. It is labelled as follows: “Type”

(round label with red margin)/ “S.S. Chatterjee 12. III. 1924” / “Naga Hills 4000'

Assam” / “Q. assamensis Cam. TYPE” / “M. Cameron. Bequest. B.M. 1955-147”.

The specimen was dissected and the aedoeagus (the paramere was separated) was

mounted in Canada Balsam. The specimen is hereby designated as the lectotype of

Q. assamensis-, the label “Lectotype Quedius assamensis Cameron A. Smetana des.

1984” has been attached to it.

Revision of the Tribes Quediini and Atanygnathini

285

The remaining four specimens belong to Q. aureipilis (see there for details); they

were accordingly labelled.

Geographical distribution. — Qiiedius assamensis is distributed mainly in the

eastern portion of the Himalayas, from central Nepal to Nagaland, however, its

distributional range extends westwards to Kumaon, Uttar Pradesh (Map 15).

Material studied. — 26 specimens.

INDIA. Nagaland. See Type material. Uttar Pradesh. Kumaon, Haldwani Distr., H.G. Champion

(BMNH)5.

NEPAL. Kathmandu Distr. Gokarna Forest, 1400 m, 31. Ill and 1.IV.81, Ldbl & Smetana (ASCC) 3.

Khandhari Distr. Khandbari, 1700 m, 23.111.82, A. & Z. Smetana (ASCC) 1; Pangma, 1700 m, 4.IV.84,

Smetana & Lobl (ASCC, CNCC) 4; Pangma-Ahale, 1600-2000 m, 4. IV. 84, Lobl & Smetana (MHNG) 2;

Arun Valley at Num main bridge, 1050 m, 20-21. IV. 84, Smetana & Lobl (ASCC, MHNG) 7. Sindhnpalchok

Distr. 4 km S Tarang Marang, 900 m, 28.IV.81, Lobl & Smetana (ASCC, MHNG) 2. Dumahan, IV. 1984, de

Rougemont (GDRC) 1.

Bionomics. — Quedius assamensis occurs at lower elevations under 2000 m. The

specimens from Nepal were collected by sifting leaf litter and various debris in

moist to wet habitats, such as seepages etc. both in forests and in open disturbed

habitats. The specimens from Arun valley were taken, together with Q. nilo, by

sifting thin layer of soaking wet leaves and other debris on a sandy bank of a creek.

Comparisons. — Quedius assamensis is in all external characters also similar to

Q. chinensis Bernhauer 1915 (I do not know the male of the latter species). Quedius

chinensis differs mainly by the microsculpture on both the head and pronotum; it is

almost entirely meshed on the head and tends to form irregular meshes on the

pronotum (based on one “cotype” from Bemhauer’s collection).

49. Quedius (Raphirus) himalayicus Bernhauer

Figs. 207-210; Map 25

Quedius himalayicus Bemhauer 1915:55; Cameron 1932:293

Description. — Black, elytra dark metallic green to greenish-blue, abdomen slightly iridescent;

appendages piceous-black to black, all tarsi reddish-brown. Head rounded, wider than long (ratio 0.73); eyes

large and convex, tempora considerably shorter than length of eyes seen from above (ratio 0.31); no

additional punctures between anterior frontal punctures; posterior frontal puncture situated very close to

posteromedian margin of eye, separated from it by distance smaller than diameter of puncture, one additional

setiferous puncture between it and posterior margin of head; temporal puncture almost touching posterior

margin of eye; surface of head with very fine, dense microsculpture of irregular transverse waves with

numerous longitudinal junctions and therefore forming rudimentary meshes here and there and becoming

more or less meshed on central portion of clypeus. Antenna rather short and only slightly thickened toward

apex, segment 3 longer than segment 2, following four segments longer than wide, gradually becoming

shorter and wider, outer segments feebly longer than wide to equally long as wide, last segment slightly

shorter than preceding two segments combined. Pronotum feebly wider than long (ratio 1.15), widely

rounded basally and distinctly narrowed anteriorly, evenly transversely convex; dorsal rows each with three

fine punctures; sublateral rows each with two punctures, posterior puncture situated slightly behind level of

large lateral puncture, posterior puncture missing bilaterally in several specimens; surface of pronotum with

extremely fine, dense microsculpture of transverse waves, in general more superficial and finer than that on

head. Scutellum punctate. Elytra moderately long, at base only slightly narrower than pronotum at widest

point, at suture about equally long, at sides feebly longer than pronotum at midline (ratio 1.10); punctation

moderately coarse, dense, interspaces between punctures about equal to diameters of punctures; pubescence

black. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing distinct whitish apical seam of

Quaest. Ent., 1988, 24 (2)

286

Smetana

palisade setae; punctation finer and somewhat denser than that on elytra, becoming sparser toward apical

margin of each tergite and in general becoming less dense toward apex of abdomen; pubescence black,

dense. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus more dilated than in female. Apical margin of sternite 7 slightly

arcuately emarginate apically, slightly flattened and smooth before emargination; apical margin of sternite 8

with moderately deep and wide, arcuate emargination (Fig. 207), small triangular area before emargination

flattened and smooth. Aedoeagus (Figs. 208-210) large and voluminous, median lobe suddenly narrowed

into a short, obtuse apical portion entirely covered by apical portion of paramere. Paramere moderately long,

broadly constricted in middle portion, more or less broadly rounded apically; four apical setae, inner pair

much longer than lateral pair; two unequally long setae at each lateral margin way below apex of paramere;

underside of paramere with two rather long and slightly irregular lateral rows of sensory peg setae.

Length 8. 4-9. 3 mm.

Type material. — The female holotype (see Bernhauer 1915:55) in the Bemhauer

collection in the Field Museum of Natural History, Chicago, Illinois, is labelled as

follows; “Bhowali Kumaon” / “17.6.1912” / “For Zool Coll” / “56” / “himalayicus

Bernh. Typus unic.” / “Chicago NH Mus. M. Bernhauer Collection”. The specimen

is in fair shape: the left antenna is missing, only two basal segments remain from the

right antenna and left hind tarsus is missing.

Geographical distrihution. — Qiiedius himalayicus is widely distributed

throughout the Himalayan range, from Uttar Pradesh eastward to Darjeeling area

and Nagaland (Map 25).

Material studied. — 87 specimens.

INDIA. Nagaland. Naga Hills, Laimatak (BMNH) 1. Himachal Pradesh. Murree Hills, VI. 1888,

Thobba (BMNH) 1; Naggar, Kuly, 5000', Champion (BMNH) 1; Jibhi, Seraj, 6000', V.1926, Champion

(BMNH) 2; Jhatingri, Mandi, 6000', Champion (BMNH) I; Simla, VI.81, R. de Rougemont (GDRC) 1;

Macleodganj, VI.81, R. de Rougemont (GDRC) I. Uttar Pradesh. Chakrata Dsitr; Korawa Khud, 9100',

4.V.22, Cameron (BMNH) 1; Jadi Gad, 7000', 9.V.22, Cameron (BMNH) 1; Manjgaon, 6500', 18.V.22,

Cameron (BMNH) 1. Mussoorie Distr.: Dhobi Ghat, 14. IV. 22, Cameron (BMNH) 4; Mossy Falls, 20,24.

and 27.III.21, 22.III.32, Cameron (BMNH, USNM) 11. Kumaon: W. Almora, Champion (BMNH) 22; W.

Almora Divn., VI. and VIII. 1917, III. 1918, Champion (BMNH) 7; Ranikhet, Champion (BMNH) 1.

Garhwal: between Tehri and Srinagar, 900 m, 25.X.79, I. Lobl (MHNG) 1; 22 km N Rishikesh, 450 m,

30.x. 79, 1. Lobl (MHNG) 1. West Bengal. Rungbong Vail., Gopaldhara, H. Stevens (BMNH) 22; Darjeeling

Distr., Gopaldhara, 3440^720', 10 and 15.XI.19, H. Stevens (BMNH) 3; Namsoo, XI.1920, H. Stevens

(BMNH) 1.

NEPAL. Nnwakot Distr. above Shermanthang, 2900 m, 26.IV.81, Lobl & Smetana (ASCC) 1; 4 km S

Tarang Marang, 900 m, 28.IV.81, Lobl & Smetana (ASCC) 1.

Bionomics. — Little is known about the habitat requirements of Q. himalayicus.

The specimens from Nepal were taken among debris on moist bottom of a dried out

pond (above Shermanthang) and by sifting soaking wet leaf litter and other debris at

base of a small waterfall (near Tarang Marang).

Comparisons. — Quedius himalayicus, and the two following species, can easily

be recognized by the following combination of characters: fairly large size, body

black with metallic green to greenish-blue or metallic blue elytra, head with very

large and convex eyes (see the description), scutellum punctate, elytra and abdomen

densely punctate. On the other hand, it is not easy to distinguish these three species

and males are necessary for a positive identification (see under Q. nilo and Q. kuiro

for details).

Revision of the Tribes Quediini and Atanygnathini

287

Quedius himalayicus also resembles Q. assamensis', however, the latter species

differes, in addition to the characters on the aedoeagus, by the non-metallic black

elytra. Another similar species, Q. aureipilis, differs from Q. himalayicus

conspicuously by the pale yellowis femora and by golden-yellowish pubescence of

the elytra and abdomen.

50. Quedius (Raphirus) uilo spec. now

Figs. 211-214; Map 25

DesCI'iptiou. — in all external characters very similar to Q. himalayicus but different as follows:

elytra almost always metallic blue, only rarely withy greenish tint; head in general slightly more transverse

with slightly larger and more convex eyes (ratio length of temporal length of eyes seen from above =0.23);

antenna slightly longer, with outer segmetns usually somewhat longer than wide; pronotum in general

narrower (ratio width; length = 1.07) and in most specimens about equally narrowed anteriorly and

posteriorly; punctation of elytra and abdomen in general slightly coarser and less dense.

Male. First four segments of front tarsus more dilated than in female. Sternites 7 and 8 with

emarginations similar to Q. himalayicus, that of stemite 8 deeper (for sternite 8 see Fig. 211). Aedoeagus

(Figs. 212-214) similar to that of Q. himalayicus', however, apical portion of median lobe less abruptly

narrowed and in general wider, apical portion with distinct dent in apical portion. Paramere much wider with

apical portion more elongately naiTOwed and narrowly rounded apically, apical setae closer together, setae at

lateral margins shorter, rows of sensory peg setae on underside of paramere start farther from apex of

paramere and are more irregular.

Length 7. 9-9. 9 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Dis. Arun Valley at Num main bridge 1050 m 20.IV. 1984 Smetana & Lobl. In the

Smetana collection, Ottawa, Canada.

Paratypes (20): same data as holotype, date 20. or 21. IV. 84 (ASCC, BMNH,

CNCC, MNHG) 20.

Geographical distribution. — Quedius uilo is at present known only from the

type locality in eastern Nepal (Map 25).

Bionomics. — The specimens from the Arun Valley in Nepal were taken,

together with Q. assamensis, by sifting thin layer of soaking-wet leaves and other

debris on a sandy bank of a creek.

Comparisons and variations. — Although very similar in all external characters

to Q. himalayicus, Q. nilo can be distinguished without much difficulty by the

characters mentioned in the description, particulrly by the different shape of the

pronotum.

For a comparison with Q. kuiro see the discussion under the latter species.

Etymology.— The specific name is the Nepali adjective nilo (blue); it refers to

the colouration of the elytra of this species.

Quaest. Ent., 1988, 24 (2)

288

Smetana

51. Quedius (Raphinis) kuiro spec.nov.

Figs. 215-217; Map 25

Description. — in all external characters very similar to Q. himalayicus but different as follows:

head with slightly more convex eyes, tempora slightly longer (ratio length of tempora: length of eyes seen

from above = 0.29), surface of head and pronotum with finer microsculpture, slightly iridescent, especially

that of pronotum; antenna longer with outer segments longer than wide; elytra in general longer, bright

metallic blue.

Male. First four segments of front tarsus more dilated than in female. Apical margin of stemite 7 with

moderately wide and deep arcuate emargination and slightly flattened in front of emargination; apical

margin of sternite 8 with wide and very deep, obtusely triangular emargination fringed by densely packed

dark setae (Fig. 215). Aedoeagus (Figs. 216, 217) very large; median lobe abruptly narrowed into rather

short and narrow apical portion entirely covered by paramere. Paramere narrow and elongate, almost

parallel-sided in middle portion, with narrowly arcuate apex; four apical setae, median pair much longer and

stronger than lateral pair, lateral pair distinctly shifted below apex of paramere; two unequally long setae at

each lateral margin way below apex of paramere; underside of paramere with two long rows of sensory peg

setae, diverging posteriorly.

Length 8. 7-9.0 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Distr. Induwa Khola Valley 2000 m 17. IV, 84 Smetana & Lobl. In the Smetana

collection, Ottawa, Canada.

Geographical distribution. — Quedius kuiro is at present known only from the

type locality in eastern Nepal (Map 25).

Bionomics. — The two specimens of the original series were taken by sifting thin

layer of moist leaf litter and other debris on sandy or gravelly banks of the Induwa

Khola river.

Comparisons. — Quedius kuiro is in all characters similar to Q. nilo, but differs

from it, in addition to the characters on the aedoeagus, by the less transverse head

with slightly longer tempora, by the still longer antenna, the shape of the pronotum

(same as in Q. himalayicus), the more densely punctate elytra and, in the male sex,

by the secondary sexual characters on the abdominal stemites, particularly by the

wide and very deep emargination of stemite 8, fringed by densely packed dark setae.

The last character is quite characteristic for this species.

Etymology. — The specific name is the Nepali noun kuiro (fog) in apposition. It

was applied in the memory of the turbulent, stormy, foggy and wet weather that

made our collecting in the Induwa Khola Valley so challenging and frustrating at

times, but after all quite rewarding and successful.

52. Quedius (Raphinis) aureipilis Cameron

Figs. 218-223; Map 18

Quedius aureipilis Cameron 1932;294

Description. — in all characters similar to Q. himalayicus but different as follows: piceous-black,

maxillary palpi brunneous, antennae piceous with each first segment brunneous, femora pale yellowish,

tibiae piceous to piceous-black, tarsi brunneous; abdomen iridescent; pubescence of elytra and abdomen

golden-yellowish, forming slightly denser patches on lateral portions of abdominal tergites. Head slightly

smaller with somewhat shorter tempora (ratio length of tempora: length of eyes seen from above = 0.17):

Revision of the Tribes Quediini and Atanygnathini

289

pronotum distinctly narrower and about equally narrowed anteriorly and posteriorly, elytra narrower. Hatter

and more parallel-sided, usually slightly more densely punctate.

Male: First four segments of front tarsus more dilated than in female. Apical margin of sternite 7 feebly,

indistinctly concave in middle; apical margin of sternite 8 with moderately wide and deep, obtusely

triangular emargination (Fig. 218), small triangular area before emargination flattened and smooth.

Aedoeagus (Figs. 221-223) with median lobe narrowed into moderately long, obtusely triangular apical

portion, entirely covered by paramere; apical portion of median lobe with small dent in lateral view.

Paramere rather long, almost parallel-sided in middle portion, nanowed into subacute apex; four apical

setae, two middle ones distinctly longer than lateral ones; two unequally long setae at each lateral margin

below apex; underside of paramere with two irregular, lateral rows of sensory peg setae.

Length 8. 5-9.0 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains three specimens under the name Q. aureipilis. They are labelled as

follows: Spec. No. 1 (female): “Type” (round label with red margin)/ “Naga Hills

Laimatak Assam (12)” / “Q. aureipilis Cam. TYPE”. Spec. No. 2 (female): “Naga

Hills Laimatak Assam (12)”/ “M. Cameron. Bequest B.M. 1955-147” /

“SYN-TYPE” (round label with blue margin). Spec. No. 3 (male): same labels as

Spec. No. 2. The male specimen No. 3 was dissected and the aedoeagus was

mounted in Canada Balsam. The male specimen is hereby designated as the

lectotype of Q. aureipilis', the label “LECTOTYPE Quedius aureipilis Cameron

Smetana des. 1984” has been attached to it.

Geographical distribution . — Quedius aureipilis is at present known only from

the Naga Hills in Nagaland (Map 18).

Material studied. — 9 specimens.

INDIA. Nagaland. Naga Hills, Laimatak (ASCC, BMNH, CNCC) 6.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Quedius aureipilis differs readily from all remaining species of

the Himalayicus Group by the pale yellowish femora and by the golden-yellowish

pubescence of the elytra and abdomen.

Intricatus Group

This species group is characterized by the following combination of characters:

small to medium sized species; head, pronotum and elytra bright metallic; head and

pronotum with numerous punctures that often are very coarse and deep and may

form rugae on head; segment 3 of antenna longer than segment 2; scutellum

impunctate or punctate; elytra with simple punctation or with very coarse punctation

forming rugae; first four segments of front tarsus strongly dilated in both sexes.

The group contains at present three species in the Himalayan region and another

species {Q. multipunctatus Sharp 1889) in Japan.

Quaest. Ent., 1988, 24 (2)

290

Smetana

Map 25. Distribution records for: Q. himalayicus (% )\Q. nilo ( A ); and Q. kuiro ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

291

53. Quediiis (Raphirus) hariyo spec.nov.

Figs. 224-227; Map 24

Description. — Head, pronotum and elytra bright metallic green, abdomen black, slightly iridescent;

both labial and maxillary palpi and antennae entirely testaceous, first segment of latter occasionally slightly

darkened; legs testaceous with darker tibiae, outer face of front tibiae and inner face of both middle and

posterior tibiae black and often also apices of middle and posterior femora darkened. Head rounded, slightly

wider than long (ratio 1.25); eyes large and convex, tempora considerably shorter than length of eyes seen

from above (ratio 0.23); two fine setiferous punctures between anterior frontal punctures; posterior frontal

puncture situtated very close to posteromedian margin of eye and separated from it by distance no larger

than diameter of puncture; temporal puncture almost touching posterior margin of eye; group of fairly coarse

punctures anteriad, mediad and posteriad of posterior frontal puncture; surface of head with scattered,

extremely fine punctures appearing somewhat denser and coarser on posterior portion of head, and with fine

and dense microsculpture of iiregular transverse waves with tendency to form here and there incomplete

rudimentary meshes. Antenna moderately long, segment 3 longer than segment 2, segments 4-7 longer than

wide, gradually becoming shorter, segments 8-10 about as long as wide, last segment slightly shorter than

two preceding segments combined. Pronotum about as long as wide, widely arcuate basally and distinctly

narrowed anteriorly; dorsal rows long and irregular, each consisting of 7-9 punctures; sublateral rows very

long and irregular, each with 6-8 punctures, posterior puncture situated considerably behind level of large

lateral puncture; minute, shaiply engraved longitudinal impression close to each lateral margin in about

middle; surface of pronotum with scattered extremely fine punctures on middle portion and with fine and

dense microsculpture of transverse waves. Scutellum impunctate, with rudimentary microsculpture. Elytra

rather long, at suture feebly (ratio 1.13) at sides slightly (ratio 1.22) longer than pronotum at midline;

punctation dense and moderately coarse, interspaces between punctures about equal to slightly larger than

diameters of punctures; pubescence yellowish; surface between punctures without microsculpture. Wings

fully developed. Abdomen with tergite 7 (fifth visible) bearing very distinct whitish apical seam of palisade

setae; punctation of tergites extremely fine, rather sparse on front tergites and gradually becoming even

sparser toward apex of abdomen; pubescence dark on middle portions but becoming golden-yellowish

toward lateral portions of tergites. Front tarsus strongly dilated in both sexes.

Male. First four segments of front tarsus more dilated than in female. Stemite 8 with three strong and

long setae on each side in apical half; apical margin with moderately deep and wide, obtusely triangular

emargination (Fig. 224), small triangular area before emargination flattened and smooth. Aedoeagus (Figs.

225-227) large and elongate, median lobe in general slightly narrowed toward apex, slightly swollen around

middle and more or less broadly arcuate apically; internal sac as in Fig. 226. Paramere long and strong, not

quite reaching apex of median lobe, slightly narrowed toward arcuate apex; four apical setae, two middle

ones much longer and stronger than lateral ones, two long and strong setae at each lateral margin below

apex; sensory peg setae on underside of paramere numerous, forming two irregular and rather wide groups,

each with 22-24 peg setae.

Fength 6. 8-7. 5 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL, Khandbari

District” / “above Sheduwa 3000 m 2. IV. 82 A. & Z. Smetana”. In the collection A.

Smetana, Ottawa, Canada.

Paratypes (5): Nepal: same data as holotype (ASCC) 1; same data as holotype

but date 31.III.-1.IV.84 (CNCC) 1; Khandbari Distr., For. NE Kuwapani, 2450 m,

13. IV. 82, A. & Z. Smetana (BMNH) 1; Prov. Bagmati, Gul Bhanjyang 2600 m,

6.IV.81, Lobl & Smetana (MHNG, ASCC) 2.

Geographical distribution. — Quedius hariyo is known from a few localities in

central and eastern Nepal (Map 24).

Bionomics. — All specimens of the original series were taken in broad-leaved

semideciduous forests in elevations between 2450-3000 m by sifting moist moss on

fallen trees.

Quaest. Ent., 1988, 24 (2)

292

Smetana

Comparisons. — Quedius hariyo is a very conspicuous species due to the bright

metallic green colour of the forebody in combination with the chaetotaxy of the

head and pronotum. The colouration of the body and the development of the

aedoeagus, including the armature of the internal sac, suggests close relationship of

Q. hariyo to Q. rugosus\ however, the latter species differs immediately by the very

rough sculpture of the forebody. Also, the habitat requirements of the two species

differ (see under Q. rugosiis).

Quedius hariyo is in all external characters very similar to Q. multipunctatus

from Japan; however, the Japanese species differs as follows: palpi, antennae and

legs uniformly rufo-testaceous, extremely fine punctures on head much less

numerous, elytra longer and punctation of abdominal tergites denser.

Etymology. — The specific name is the Nepali adjective hariyo (green). It refers

to the colouration of the species.

54. Quedius (Raphirus) rugosus Cameron

Figs. 228-231; Map 26

Quedius ruiyosKS Cameron 1921:273; 1932:298

Quedius mussoorieusis Cameron 1932:299 (syn.uor.)

Description. — Head, pronotum and elytra brilliant metallic green, dark green, bluish green or

purplish-green, abdomen black, slightly iridescent; both labial and maxillary palpi testaceous, or partially

darkened or almost entirely piceous; antennae testaceous, or first segment piceous with metallic reflections,

or antennae brunneopiceous with first 2-3 segments darker and with metallic reflections; legs piceous to

piceous-black, tibiae with metallic reflections, femora (except for darker apices) and coxae pale testaceous.

Head rounded, slightly wider than long (ratio 1.21); eyes very large and convex; tempora considerably

shorter than length of eyes seen from above (ratio 0.20); surface of head covered with deep and coarse

punctures forming more or less distinct rugae on posterior portion of head and gradually becoming finer and

well isolated toward clypeus, clypeus to variable extent impunctate, small impunctate area present also on

vertex; punctation in general obscuring usual setiferous punctures that can only be traced by presence of

long setae; surface between punctures with very fine microsculpture of rudimentary transverse waves and/or

incomplete meshes. Antenna moderately long, segment 3 longer than segment 2, segments 4-6 longer than

wide, gradually becoming shorter, outer segments feebly longer than wide to indistinctly transverse, last

segment shorter than two preceding segments combined. Pronotum about as long as wide, fairly strongly

margined laterally and basally; widely arcuate basally, either equally arcuately narrowed both anteriorly and

posteriorly, or more distinctly narrowed anteriorly; dorsal rows very long and more or less irregular, convex

toward midline, each formed by 7-10 very coarse, pit-like punctures; lateral portions each with a group of

very coarse punctures similar to those in dorsal rows and with variable number of coarse and fine punctures,

same fine punctures usually also present anteriorly between dorsal rows, posterior area between dorsal rows

usually without any punctures; surface of pronotum with microsculpture similar to that on head. Scutellum

impunctate, with extremely fine rudimentary microsculpture. Elytra moderately long, at base about as wide

as pronotum at widest point, at suture about as long as pronotum at midline, at sides somewhat longer (ratio

1.25); punctation very coarse and deep, confluent, to variable extent forming transverse and/or oblique

rugae, deflexed lateral portion of each elytron with more or less fine, simple punctation; pubescence

golden-yellow, occasionally intermixed with dark hairs and with a group of whitish hairs on lateral portion

of each elytron; surface between rugae without microsculpture. Wings fully developed. Abdomen with

tergite 7 (fifth visible) bearing very distinct whitish apical seam of palisade setae; punctation of tergites

extremely fine, dense on basal portions and gradually becoming sparser toward apical margin of each tergite:

surface with extremely fine, dense microsculpture of incomplete transverse waves; pubescence dark on

middle portion but becoming golden-yellow on lateral portions of each tergite, golden-yellow hairs usually

also present on apical margin of tergites 1-4. Front tarsus strongly dilated in both sexes.

Revision of the Tribes Quediini and Atanygnathini

293

Male. First four segments of front tarsus more dilated than in female. Stemite 8 with two strong and

long setae on each side in apical half, apical margin with moderately wide and deep, almost arcuate

emargination (Fig. 228), small triangular area before emargination flattened and smooth. Aedoeagus (Figs.

229-231) large and elongate, median lobe subparallel-sided to slightly narrowed toward broadly arcuate

apex; internal sac as in Fig. 230. Paramere strong, narrowed toward apex, not quite reaching to slightly

exceeding apex of median lobe, obtuse apically; four apical setae, two middle ones much longer and stronger

than lateral ones, two fairly long and strong setae at each lateral margin below apex; sensory peg setae on

underside of paramere numerous, forming two irregular, medially only vaguely separated, longitudinal

groups, each with 14-23 peg setae.

Length 5. 8-6. 8 mm.

Type material. — Quedius rugosus. The collection of the British Museum

(Natural History), London, contains 7 specimens under the name Q. rugosus\

however, only first three belong to the original series. They are labelled as follows;

Spec. No. 1: “Type H.T.” (round label with red margin)/ “SYN-TYPE” (round label

with blue margin)/ “Lebong 5000 ft. IX. 1908 H.M.L.” / “Quedius rugosus Cam.

TYPE” / “M. Cameron. Bequest. B.M. 1955-147.” Spec. No. 2: “Sikkim N. India

1920.258” / “Lebong 5000 ft. IX. 1908 H.M.L. (this label is on pin with text down)/

“Quedius rugosus Cam.”. Spec. No. 3: “W. Almora, Kumaon. India. H.G.C.” /

“SYN-TYPE”.

The first specimen (male) was dissected and the tergite and sternite 8, the genital

segment and aedoeagus were mounted on plate with beetle. The specimen is in fair

shape (someone tried to dissect it): left antenna missing except for basal segment,

right antenna broken off after second segment (rest glued on plate), right posterior

tarsus missing except for first segment, abdomen separated and in two pieces and

8th sternite badly damaged. The specimen (the other two are in worse condition) is

hereby designated as the lectotype of Q. rugosus; the label “Lectotype Quedius

rugosus Cameron Smetana des. 1984” has been attached to it.

Only fragments of the second specimen remain; the elytra with both pairs of

middle and posterior legs (tarsus of left middle leg missing) on a minuten pin. There

is a plate under the elytra, bearing the head and thorax of a small brownish quediine

unknown to me, with small eyes and only one fine puncture in dorsal rows of the

pronotum.

The third specimen (male) from W. Almora is missing the head and prothorax,

including all appendages (except for right front leg). Someone dissected it: the 8th

stemite and genital segment are rather badly damaged and glued on plate, and the

aedoeagus with the paramere separated are glued on plate.

Quedius mussooriensis. The collection of the British Museum (Natural History),

London, contains two female specimens under the name Q. mussooriensis. They are

labelled as follows: Spec. No. 1; “Type” (round label with red margin)/

“SYN-TYPE” (round label with blue margin)/ “Arni Gad, Mussoorie” / “Dr.

Cameron. 13.IV.22” / “Q. mussooriensis Cam. TYPE” / “Cameron. Bequest. B.M.

1955-147.” Spec. No. 2: “Ami Gad, Mussoorie.” / “Dr. Cameron. 28.V.21” / “M.

Cameron. Bequest. B.M. 1955-147.” / “SYN-TYPE” (round label with blue

margin).

Quaest. Ent., 1988, 24 (2)

294

Smetana

The first specimen is hereby designated as the lectotype of Q. mussooriensis; the

label “Lectotype Quedius mussooriensis Cameron Smetana des. 1984” has been

attached to it. It cannot be specifically distinguished from the lectotype of Q.

rugosus. The name Q. mussooriensis is a junior synonym of Q. rugosus\ my

corresponding determination label was attached to the specimen.

Geographical distribution. — Quedius rugosus is widely distributed throughout

the Himalaya; from Himachal Pradesh through Nepal and the Darjeeling area to

northern Burma (Map 26).

Material studied. — 25 specimens.

BURMA. Mishmi Hills, Chhagion, 3350', 25.11.35, M. Steele (BMNH) 1.

INDIA. Nagaland. Naga Hills, 9000' (BMNH) 1. Uttar Pradesh. Kumaon: W. Almora, Champion

(BMNH) 3; W. Almora Dvn., May 1919, Champion (ASCC, BMNH) 5; Haldwani Dist., Champion

(BMNH) 2; Gori Riv. Gorge, 9000', Champion (BMNH) 1. West Bengal. Ghum Distr., Rongdong Valley,

V-VI.31, Cameron (BMNH, CNCC) 2; Ghum Distr., Mangpo, V. 1931, Cameron (BMNH) 1.

NEPAL. Chapiiri. 16.111.79, de Rougemont (ASCC, GDRC) 2; Nagarkot, X.80, de Rougemont (GDRC)

1; Khandhari distr. 2 km E Mansingma, 1900 m, 19. IV. 84, Lobl & Smetana (MHNG) 2.

Bionomics. — The habits of Q. rugosus, and possibly of additional not yet

described species of this group from southeastern Asia, can be summarized as

follows (de Rougemont in litt.): the species are very hygrophilous, found

exclusively near running water, either in the spray zones of cascades or in wet moss

under waterfalls or near the waterline of boulders. They are very active insects,

taking immediately to the wing when their habitat is disturbed, or running very

quickly for a short distance before taking off. In Thailand, several specimens of a

possibly undescribed species were observed hovering aroung a boulder in

mid-stream, alighting and running over its surface before taking to the wing and

repeating the process.

The coarse sculpture and green or bluish metallic reflection of the forebody are

analogous to that of many species of Dianous which inhabit the same biotope, and

undoubtedly serve to conceal the beetles amid the glistening droplets of water from

spray on moss and lichens in that habitat.

The species must compete to a certain extent with Dianous species for prey, but

it is possible that their greater mobility enables them to capture certain species

which evade the slower Dianous.

Comparisons. — Originally, I considered Q. mussooriensis, which tended to be

more brightly metallic green with antennae uniformly testaceous and with the

sculpture in general less coarse, as a distinct species. However, as the number of

specimens studied increased, it became apparent that it is impossible to specifically

distinguish Q. mussooriensis from Q. rugosus. The colouration of antennae, the tint

of the mettalic surface and the coarseness, location and number of the punctures on

the head and pronotum and of the rugae on the elytra varies to great extent without

being correlated in any way with each other. Also, the aedoegai are essentially

identical (including the development of the internal sac) and the number of the peg

setae on the underside of paramere varies in such a way, that any correlation with

some of the external characters is impossible. For these reasons I decided to

Revision of the Tribes Quediini and Atanygnathini

295

recognize only one species, Q. rugosus.

55. Qiiedius (Raphirus) inthcatus Fauvel

Map 26

Quedius iutricatits¥?L\iyQ\ 1895:274; Cameron 1932:298

Description. — Very similar to Q. rugosus but different as follows: antennae always uniformly

testaceous. Head with coarse, dense and deep punctation forming rugosities except for small smooth area on

vertex. Neck with numerous punctures on middle portion . Pronotum without appreciable rows of punctures,

entire surface of pronotum covered with coarse rugulose punctation except for two small smooth areas on

each side of midline at about middle. Scutellum with punctures situated in coarse transverse depressions.

Male. Unknown.

Length 7.0 mm.

Type material. — The Fauvel collection in the Institut Royal des Sciences

Naturelles de Belgique, Bruxelles, contains one female specimen under the name Q.

intricatus. It is labelled as follows: “Carin Ghecu 1300-1400 m L. Fea II-III.88” /

“intricatus Fvl.” / “Syntype” / “R.I. Sc. N.B. 17479 Quedius Coll, et det. A. Fauvel.”

The speciment is hereby designated as the lectotype of Q. intricatus', the label

“Lectotype Quedius intricatus Fauvel Smetana des. 1983” has been attached to it.

Geographical distribution. — Quedius intricatus is at present known only from

the type locality in Burma (Map 26).

Material studied. — The Lectotype.

BURMA. See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species;

however, it almost certainly lives in a similar way to Q. rugosus.

Comparisons. — Quedius intricatus is a poorly known species due to lack of

specimens available for study. However, it can easily be distinguished from the

related species, Q. rugosus, by the characters given above, particularly by the

punctate neck and by the sculpture of the scutellum.

Gardneri Group

The single species of this group, Q. gardneri, is unique because of the

configuration of the abdominal tergites 3-6, each bearing a longitudinal median

keel, and by the paramere of the aedoeagus divided in two fairly long branches (Fig.

233).

See the discussion following the description of Raphirus for a discussion of Q.

gardneri.

56. Quedius (Raphirus) gardneri Cameron

Figs. 232-235; Map 26

Quedius gardneri Cameron 1932:292

Quedius coeruleus Coiffait 1977:241 {syn.now)

Quaest. Ent., 1988, 24 (2)

296

Smetana

Description. — Piceous-black, elytra greenish-blue, abdomen feebly iridescent; maxillary and labial

palpi, antennae and legs piceous. Head narrow, about as long as wide, distinctly narrower than pronotum at

widest point (ratio 0.68), strongly narrowed posteriorly behind eyes, posterior angles absent; eyes rather

large and moderately convex, tempora distinctly shorter than length of eyes seen from above (ratio 0.55); no

additional punctures between anterior frontal punctures; posterior frontal puncture situated close to

posteromedian margin of eye and separated from it by distance about equal to diameter of puncture, two

additional setiferous punctures between it and posterior margin of head; surface of head with very dense and

extremely fine microsculpture of transverse waves. Antenna slender, moderately long, segment 3 narrower

and slightly longer than segment 2, segment 4 distinctly, segment 5 somewhat longer than wide, following

segments gradually becoming shorter, outer segments about as long as wide. Pronotum narrow, as long as

wide, widely arcuate basally and strongly narrowed anteriorly; dorsal rows each with three punctures;

sublateral rows each with two punctures, posterior puncture situated at or slightly behind level of large

lateral puncture; surface of pronotum with microsculpture similar to that on head. Scutellum large, without

punctures, surface with rather dense and very fine microsculpture of irregular transverse waves. Elytra long,

at suture feebly (ratio 1.07), at sides distinctly longer (ratio 1.31) than pronotum at midline; punctation fine

and not dense, interspaces between punctures much larger than diameters of punctures; pubescence

yellowish; surface between punctures without microsculpture. Wings fully developed. Abdomen with tergite

7 (fifth visible) bearing whitish apical seam of palisade setae; first three tergites distinctly impressed at base,

impressions divided by obtusely elevated median keel that becomes vague on third tergite; basal impressions

of first three tergites with rather dense and coarse punctation, remainder of tergal surface with only scattered

very fine punctures, median portions impunctate, impunctate area most extensive on first two tergites; tergite

five finely and not densely punctate, punctation coarser and denser at base; pubescence piceous, with

yellowish hairs here and there. Front tarsus dilated in both sexes.

Male. First four segments of front tarsus distinctly more dilated than in female. Stemite 8 with wide and

deep, acutely triangular emargination in middle apical margin (Fig. 232), small triangular area before

emargination Fattened and smooth. Aedoeagus (Figs. 233-235) moderately large; median lobe elongate,

constricted in middle portion, apical portion with obtuse apex, face adjacent to paramere with two strongly

sclerotized, acutely pointed lamellae curved medially; paramere robust, divided in two fairly long branches

separated at base by obtusely angulate arc; apices of branches not reaching apex of median lobe; underside

without any sensory peg setae but with two pairs of minute setae.

Length 6. 5-6. 8 mm.

Type material. — Quedius gardneri. The collection of the British Museum

(Natural History). London, contains one female specimen under the name of Q.

gardneri. It is labelled as follows: “Type” (round label with red margin)/

“Lepchajagat, 7000' Darjeeling, Bengal. J.C.M. Gardner. 13. IX. 1929.” / “495” /

“Quedius gardneri Cam. TYPE” / “M. Cameron Bequest. B.M. 1955-147.” The

specimen is in perfect shape. It is hereby designated as the lectotype of Q. gardneri-,

the label “Lectotype Quedius gardneri Cameron A. Smetana des. 1984” has been

attached to it.

Quedius coeruleus. Coiffait (1977:241) described the species from two males

from Bhutan. I was able to study the holotype deposited in the Naturhistorisches

Museum in Basel, Switzerland; it is labelled as follows: “Nobding 41 km O Wangdi

Ph. 2800 m” / “Nat. Hist. Museum Basel-Bhutan Expedition 1972” / “HOLOTYPE”

/“Quedius (Sauridus) coeruleus H. Coiffait 1976”.

The specimen was dissected and the aedoeagus and the tergite and sternite 8

were mounted in Canada Balsam. Except for the different sex, the holotype is in all

details identical with the lectotype of Q. gardneri. The name Q. coeruleus is a junior

synonym of Q. gardneri-, my corresponding determination label was attached to the

specimen.

Revision of the Tribes Quediini and Atanygnathini

297

The paratype of Q. coerideus from Dorjula (see Coiffait, l.c.) is deposited in the

collection Coiffait, Museum National d’Histoire Naturelle, Paris, France (not seen).

Geographical distribution . — Quedius gardneri is distributed in the eastern

Himalaya; it is at present known from one locality in the Darjeeling area and from

two localities in Bhutan (Map 26).

Material studied. — 3 specimens.

BHUTAN. Holotype of Q. coerideus (see Type material); Dorjula, 3100 m, 6. VI. 72 (HCCC) 1 (see

Coiffait 1977:241).

INDIA. West Bengal. Lectotype of Q. gardneri (see Type material).

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Quedius gardneri is a distinctive species, due to the

conspicuous sculpture of the basal abdominal tergites (see description), in

combination with the narrow head which is strongly narrowed posteriorly behind the

eyes, the narrow pronotum which is strongly narrowed anteriorly and the

greenish-blue elytra with yellowish pubescence. The aedoeagus is quite

characteristic, due to the paramere with two apical branches.

The species cannot be confused with any other species of Quedius from the

Himalayan area.

See under Raphirus for some additional information.

57. Quedius (suhgenus ?) Uneipennis Cameron

Figs. 236-242; Map 26

Quedius Uneipennis Cameron 1932:290

Description. — Piceous-black, head, pronotum and abdomen slightly iridescent, elytra, meso- and

mestastemum, first two abdominal tergites, base of third tergite, apical portion of fifth tergite and

conesponding stemites, and rest of tip of abdomen reddish; labial and maxillary palpi dark testaceous; first

four antennal segments piceous with paler apical portions, remaining segments testaceous; legs

rufo-testaceous. Head strongly transverse (ratio width: length = 1.52), strongly narrowed posteriorly behind

eyes, posterior angles completely rounded; eyes rather large, convex, moderately protruding from lateral

contours of head, tempora distinctly shorter than length of eyes seen from above (ratio 0.69); no additional

punctures between anterior frontal punctures, posterior frontal puncture almost touching posteromedian

margin of eye, two minute punctures on each side near inner margin of eye between anterior and posterior

frontal puncture; two rather fine punctures posteromediad of posterior frontal puncture near posterior margin

of head; temporal puncture situated slightly closer to posterior margin of head than to posterior margin of

eye, one additional setiferous puncture between it and posterior margin of eye; clypeus with extremely fine

punctures; surface of head polished, without appreciable microsculpture, however, with sparsely distributed

microscopic punctures. Last segment of maxillary palpus distinctly longer than penultimate segment, slighty

attenuate basally. Antenna moderately long, segment 3 slightly longer than segment 2, segment 4 slightly

longer than wide, segments 5 and 6 as long as wide, outer segments slightly transverse, last segment about as

long as two preceding segments combined. Pronotum wider than long (ratio 1.18), only slightly transversely

convex, widest at about apical third, strongly narrowed both anteriorly and posteriorly, lateral margins

slightly sinuate posteriorly, basal margin subtruncate medially; dorsal and sublateral rows absent; lateral

puncture almost touching lateral margin of pronotum; surface of pronotum similar to that of head. Scutellum

punctate, punctures unequally spaced. Elytra moderately long, at base only slightly narrower than pronotum

at widest point, at suture about as long as, at sides somewhat longer than pronotum at midline (ratio 1.21);

each elytron with sutural and discal longitudinal row of punctures, sutural row situated in feebly impressed

groove, discal row irregular, epipleuron coarsely punctate, pubescent and with numerous long setae; surface

of elytra without appreciable microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible)

Quaest. Ent., 1988, 24 (2)

298

Smetana

bearing whitish apical seam of palisade setae; punctation of abdominal tergites fine and moderately dense,

gradually becoming slightly finer toward apex of abdomen, somewhat denser on basal portion of tergites;

middle portions of first two tergites impunctate except basally; pubescence black. Front tarsus dilated in both

sexes.

Male. First four segments of front tarsus much more dilated than in female. Apical margin of stemite 8

with moderately wide and shallow, arcuate emargination, small triangular area before emargination flattened

and smooth, apical margin of sternite laterad of central emargination with long and strong setae (Fig. 236).

Both tergite 10 and sternite 9 of genital segment emarginate apically (Figs. 237-238). Aedoeagus (Figs.

239-242) small, median lobe asymmetrical apically, in lateral view with small tooth far below apex;

paramere small, wide basally, evenly narrowed, apical portion naiTowly incised, with four very long apical

setae and two somewhat shorter lateral setae on each side below apex; underside of paramere with two

longitudinal rows of sensory peg setae, each row with ten peg setae, gradually becoming larger and paired

toward base of paramere. Internal sac as in Fig. 240.

Length 10.0 mm.

Type material. — Quediiis Uneipennis was described from specimens from

Burma (Cameron 1932:290). I was able to study two specimens deposited in the

collection of the British Museum (Natural History), London. They are labelled as

follows: Spec. No. 1 (male): “Type” (round label with red margin)/ “Doherty” (label

upside down)/ “Birmah Ruby Mes” / “Fry Coll. 1905.100.” (label upside down)/

“Quedius lineipennis Cam. TYPE.” Spec. No. 2 (female): “64412” “Doherty” /

“Birmah Ruby M” / “Fry Coll. 1905-100.” / “SYN-TYPE” (round label with blue

margin). The male specimen was dissected, and the aedoeagus, sternite and genital

segment mounted in Canada Balsam. The specimen is hereby designated as

lectotype of Q. lineipennis', the label “Lectotype Quedius lineipennis Cameron

Smetana des. 1983” has been attached to it.

Geographieal distribution. — Quedius lineipennis is at present known only from

the type locality in Burma (Map 26).

Material studied. — See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species. I

suspect that it lives under the bark of dead trees.

Comparisons. — Quedius lineipennis is quite easily distinguished by many

characters, in particular by the transverse head, its chaetotaxy and the presence of

extremely fine punctures on the clypeus, by the shape and chaetotaxy of the

pronotum (dorsal and sublateral rows of punctures missing, lateral puncture almost

touching lateral margin of pronotum), by the type of punctation of the elytra and the

elytral epipleura, and by both the tergite and sternite of male genital segment

emarginate apically. It has a rather isolated position within the genus and may

require an erection of a separate taxon for it (see also the discussion following the

description of Quedius).

The surface of the head and pronotum is not completely devoid of

microsculpture. Microsculpture of exceedingly fine irregular waves is present, but is

difficult to observe.

The setae at the apical margin of male sternite 8 (Fig. 236) were inferred from

the large sockets present; the acutal setae on apical portion of the stemite are almost

completely missing in the lectotype.

Revision of the Tribes Quediini and Atanygnathini

299

Qiiaest. Ent., 1988, 24 (2)

Map 26. Distribution records for: Q. rugosus {%)\ Q. intricatus ( A ); Q. gardneri ( ■ ); and Q. lineipennis ( ^ )•

300

Smetana

2. Genus Indoquedius Cameron, stat.nov.

Quedius subg. Indoquedius Cameron 1932:281, 300

Type species. — Quedius oculatus Fauvel 1895, designated by Blackwelder

1952:199, by subsequent designation.

Descriptive notes. — The genus Indocpiedius shares many characters with

Quedius but it differs in several respects. The head always bears two or three

setiferous punctures between the anterior and posterior frontal punctures, situated

directly at the median margin of the eye (Fig. 400). Eyes are very large and convex,

considerably longer than the tempora. The dorsal surface of head is without

microsculpture and is therefore highly polished. On the antenna, the large setae on

antennal segments, particularly on those past segment 3, are relatively long and

strong (Fig. 401). The penultimate segments of both the maxillary and labial palpus

bear numerous strong setae, particularly the penultimate segment of the labial palpus

on its ventromedial face. The pronotum is no more than slightly narrowed anteriorly.

The dorsal rows of pronotum are each composed of 2 punctures. The dorsal surface

of pronotum lacks microsculpture, or, rarely, the microsculpture is present only in

the narrow lateral groove. Scutellum is variably punctate. Elytra are variably, simply

punctate. Wings are fully developed. First four segments of front tarsus are strongly

dilated in both sexes, at least sometimes more so in the males. The seventh (fifth

visible) abdominal tergite bears always a whitish apical seam of palisade setae. Male

genital segment with tergite 10 and sternite 9 emarginate apically. The aedoeagus,

including the parameres, is similarly developed as in the genus Quedius; the

parameres are fused in one solid sclerite, sometimes divided anteriorly in two

branches; the microsetae are absent (Figs. 246, 250, 254, 257, 262, 266, 269). The

female genital segment as in Figs. 402 and 403, tergite 10 emarginate apically.

The genus Indocpiedius is primarily a tropical genus represented by numerous

species in the Oriental region. Some of the species reach the warm temperate zone

of the Palaeractic region. These species are e.g., I. Juno Sharp 1878 (comb.nov.) and

I. praeditus Sharp 1889 (comb.nov.). Eight species of Indocpiedius are at present

known from the Himalayan region.

Ranging and relationships. — This genus was so far considered as a subgenus of

Quedius. However, the characters discussed above leave little doubt that

Indoquedius in fact is a separate genus, to some extent intermediate between

Quedius and Bolitogyriis.

Key to species of Indoquedius

1 Lateral pronotal groove smooth and shiny, without any

microsculpture. Anterior angles of pronotum impunctate; dorsal

surface of head and pronotum without micropunctation 2

F Lateral pronotal groove with dense, almost granulose

Revision of the Tribes Quediini and Atanygnathini 301

microsculpture, opaque. Anterior angles of pronotum finely

punctate; dorsal surface of head and pronotum with

micropunctation. Length 10.0 mm

8. 1, aherrans Cameron, p. 31 1

2 (1) Large lateral puncture of pronotum situated close to lateral

pronotal groove, but distinctly not touching it. Paramere of

aedoeagus split into two branches apically (Figs. 244, 248) 3

2' Large lateral puncture of pronotum situated very close to lateral

pronotal groove, distinctly touching it. Paramere of aedoeagus

entire (Figs. 252, 256, 260, 264) 4

3 (2) Elytra bright metallic blue or greenish-blue. Median lobe of

aedoeagus slightly, arcuately widened and then rather abruptly

narrowed into apex proper (Figs. 244, 245). Length 7. 2-9.0

mm 1 . /. sikkimensis Cameron, p. 302

3' Elytra deep black. Median lobe of aedoeagus evenly, conically

narrowed anteriorly (Figs. 248, 249). Length 8.0-9. 8 mm

2. 1, haliyo spec. now, p. 304

4 (2') Pronotum and head concolourous, piceous to black 5

4’ Pronotum and head not concolourous: pronotum red and head

piceous to black. Length 6. 0-7.0 mm

6. 1.filicornis (Eppelsheim) (pars), p. 309

5 (4) Scutellum, except for narrow impunctate strip along all three

margins, with coarse and dense punctation, intervals between

punctures mostly about as large as diameters of punctures.

Punctation of elytra rather dense and coarse, intervals between

punctures, at least on area near apex of scutellum, no more than

slightly larger than diameters of punctures 6

5' Scutellum with more or less sparse and fine punctation, rather

wide areas along base and margins impunctate, scutellum

sometimes with only a few punctures. Punctation of elytra

rather sparse and less coarse, intervals between punctures

distinctly to considerably larger than diameters of punctures 7

6 (5) Antennae and legs uniformly rufo-testaceous. Aedoeagus as in

Figs. 252-254. Length 7. 8-8.7 mm

3. 1, hipunctatus Eppelsheim, p. 305

6' Antennae dark, with last 2 or 2 1/2 segments yellowish. Meso-

and metatibiae darkened, meso- and metafemora slightly

darkened. (Male unknown). Length 8. 6-8. 8 mm

4. /. oculatus Fauvel, p. 306

7 (5') Paramere of aedoeagus wide and rather short, widely rounded

at apex and with short attenuate middle portion (Fig. 256).

Length 7. 5-8. 8 mm 5. /. daai spec. now, p. 307

Quaest. Ent., 1988, 24 (2)

302

Smetana

7' Paramere of aedoeagus narrow and elongate, narrowly rounded

at apex, with long attenuate middle portion or almost

parallel-sided (Figs. 260, 264) 8

8 (V) Aedoeagus large, paramere usually slightly exceeding apex of

median lobe, with long attenuate middle portion (Fig. 260).

Length 6.8-8. 8 mm...,6. 1. filicornis (Eppelsheim) (pars), p. 309

8' Aedoeagus distinctly smaller, paramere not quite reaching apex

of median lobe, almost parallel-sided (Fig. 264). Length

7.8-8. 1 mm 7. 1, saathi spec.nov., p. 3 10

Sikkimensis Group

This species group is characterized by the following combination of characters:

lateral pronotal groove smooth and shiny, without any microsculpture; anterior

angles of pronotum impunctate; dorsal surface of head and pronotum without

micropunctations; large lateral puncture of pronotum not touching lateral pronotal

groove; paramere of aedaeagus split in two branches apically (Figs. 244, 248).

The group includes two species in the Himalayan region.

1. Indoquedius sikkimensis (Cameron) comh.nov.

Figs. 243-246; Map 27

Quedius sikkimensis Cameron 1932:294

Quedius akalita Smetana 1977:246 (syn.nov.)

Algol! cyanipennis Coiffait 19826:273 {syn.nov.)

Description. — Black, elytra dark metallic greenish-blue to dark blue, abdomen iridescent; last

segments of both maxillary and labial palpi testaceo-brunneous, antennae gradually becoming paler toward

apex, all tarsi more or less paler, rather brunneo-piceous. Head rounded, somewhat wider than long (ratio

1.17); eyes very large and convex, tempora about three times shorter than length of eyes seen from above

(ratio 0.35); two or three punctures along inner margin of eye between anterior and posterior frontal

puncture; posterior frontal puncture situated close to posteromedian margin of eye, separated from it by

distance no larger than diameter of puncture; one additional setiferous puncture between it and posterior

margin of head; area behind posterior frontal puncture with numerous fine punctures; surface of head

without any microsculpture. Antenna moderately long, hardly thickened toward apex, first three segments

with fairly long dark setae; all segments longer than wide, gradually becoming shorter, penultimate segment

about twice as long as wide at base, last segment asymmetrically acuminate, somewhat shorter than two

preceding segments combined. Pronotum rather short, feebly wider than long (ratio 1.13), broadly rounded

basally and slightly narrowed anteriorly, rather strongly transversely convex; lateral groove smooth and

shiny, without microsculpture; dorsal rows each with two punctures (occasionally with three punctures

unilaterally); sublateral rows each with two fine punctures, posterior puncture situated before or occasionally

at level of large lateral puncture; large lateral puncture situtated close to lateral pronotal groove, but

distinctly not touching it; anterior angles of pronotum impunctate; surface of pronotum without any

microsculpture. Scutellum large, punctate. Elytra moderately long, at base almost as wide as pronotum at

widest point, at suture about as long as, at sides feebly longer (ratio 1.13) than pronotum at midline;

punctation and pubescence moderately coarse and dense, surface between punctures without microsculpture.

Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae;

punctation and pubescence of abdominal tergites finer and denser than that of elytra, gradually becoming

slightly sparser toward apex. First four segments of front tarsus equally dilated in both sexes.

Revision of the Tribes Quediini and Atanygnathini

303

Male. Apical margin of sternite 8 with rather shallow and wide, arcuate emargination (Fig. 243), small

triangular area before emargination Fattened and smooth. Aedaeagus (Figs. 244-246) moderately large;

median lobe slightly, arcuately widened toward apex and then rather abruptly narrowed into apex proper.

Paramere distinctly not reaching apex of median lobe, rather wide basally but narrowed toward apex, divided

in two long branches, each branch with two minute setae apically; underside of each branch in apical portion

with numerous sensory peg setae forming an elongate group; interal sac simple, with central sclerite covered

densely with scale-like structures.

Length 7. 2-9.0 mm.

Type material. — Indoquedius sikkimensis. The original material in the collection

of the British Museum (Natural History), London, contains two male specimens

under the name Quedius sikkimensis. They are labelled as follows: Spec. No. 1

“Type” (round label with red margin)/ “Sikkim:Lachung. 9-10,000 ft.

III-IV.1920.H. Stevens.” / “H. Stevens. Brit. Mus. 1922-307.” / “Q. sikkimensis

Cam. TYPE”. Spec. No. 2 “Paratype” (round label with yellow margin)/ “Sikkim:

Lachung. 9-10,000 ft. 11I-IV.1920. H. Stevens.” / “Q. sikkimensis Cam. COTYPE”/

“M. Cameron. Bequest. B.M. 1955-147”. The specimens are conspecific. The first

male bearing the label “Type” was dissected and the sternite of pygidium and the

aedeaegus were mounted in Canada Balsam; this specimen is hereby designated as

the lectotype of I. sikkimensis', the label “LECTOTYPE Quedius sikkimensis

Cameron A. Smetana des. 1984” has been attached to it.

Indociiiedius akalita. The male holotype, deposited in the Naturhistorisches

Museum in Basel, Switzerland, is labelled as follows: “Thimphu 16.4-27.4. 2400m” /

“Nat. Hist. Museum Basel-Bhutan Expedition 1972” / “HOLOTYPE Quedius

akalita A. Smetana 1975”. The specimen cannot be specifically distinguished from

the holotype of I. sikkimensis. The name /. akalita is a junior synonym of /.

sikkimensis. My corresponding determination label has been attached to this

specimen.

Indocfuedius cyanipennis. Coiffait (l.c.) described the species from four

specimens from Bhutan. I was able to study two specimens deposited in the

Naturhistorisches Museum in Basel, Switzerland. The male holotype is labelled as

follows: “Batbalithan 10. XII. (Bumthan) 2600 m” / “Bhutan-W. Roder & L.

Caminada 77” / “TYPE” (red label)/ “Algon cyanipennis H. Coiffait 1982”. The

male paratype bears two labels identical with the two first labels of the holotype,

and a label “PARATYPE”. Neither of the specimens can be distinguished from the

lectotype of Indocjuedius sikkimensis and the name A. cyanipennis is a junior

synonym of I. sikkimensis', my determination label “Indoquedius sikkimensis Cam.

Smetana det. 1983” has been attached to each specimen.

Geographical distribution. — Indoquedius sikkimensis is known from Arunachal

Pradesh, Bhutan and Sikkim (Map 27).

Material studied. — 105 specimens.

BHUTAN. Thimphu, X. 1980, Rougemont (GDRC, ASCC) 3.

INDIA. Arunachal Pradesh. Mishmi Hills, Delai Valley, ChaChe, 22. XI. 36, M. Steele (BMNH) 1.

Sikkim. Lachung, 9000-10,000', III.IV.1920, H. Stevens (ASCC, BMNH, CNCC) 93; same, 8500-9500',

28.IL-6.III.1920, H. Stevens (BMNH) 3.

Quaest. Ent., 1988, 24 (2)

304

Smetana

Bionomics. — No details are known about the habitat requirements of this

species.

Recognition. — Indoquedius sikkiniensis can easily be recognized by the

colouration. It is the only species of the genus in the Himalayan area with metallic

greenish-blue or blue elytra.

Cameron (1932:294), oddly enough, failed to recognize this species as an

Indoquedius, assigned it to the subgenus Raphirus and compared it to Q.

himalayicus.

2. Indoquedius haliyo spec.nov.

Figs. 247-250: Map 27

Description. — Similar to /. sikkirnensis but different as follows: form in general larger and more

robust; entirely (including elytra) deep black; antennal segments 4-1 1 testaceous. Head slightly more

transverse (ratio width: length = 1.19). Prontotum more voluminous, wider and usually more distinctly

narrowed anteriorly. Scutellum larger, more densely and coarsely punctate. Elytra more coarsely and densely

punctate. Abdomen similar to that of /. sikkirnensis, punctation of tergites much finer than that of elytra.

Male. First four segments of front tarsus hardly more dilated than those of female. Apical margin of

stemite 8 with very shallow, inconspicuous arcuate emargination (Fig. 247), small triangular area before

emargination flattened and smooth. Aedoeagus (Figs. 248-250) similar to that of /. sikkirnensis but larger;

median lobe evenly, conically narrowed toward apex, paramere distinctly not reaching apex of median lobe,

strongly conically narrowed and divided into two very long, closely approximate branches, each branch with

two fine short setae apically; underside of each branch with many sensory peg setae forming a long group;

internal sac similar to that of /. sikkirnensis, quite characterisitic if evaginated.

Fength 8. 0-9. 8 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL (Prov.

Bagmati) Malemchi, 2800 m. 27. IV. 81 Lobl & Smetana”. In the collection A.

Smetana, Ottawa, Canada.

Paratypes (16): NEPAL: same data as holotype (CNCC) 4; Prov. Bagmati, Gul

Bhanjyang, 2600 m, 6.IV.81; Lobl & Smetana (CNCC, MHNG) 3; Prov. Bagmati,

Dobate Ridge NE Barahbise, 2700 m, 7.V.81, Lobl & Smetana; same, 2800 m,

2.V.81, Lobl & Smetana (CNCC, MHNG) 3; “Umg. Shermathang Helambu, Z.

Nepal Ig. H. Franz 1980 (HFCC) 1”; Bhalukop, 2800 m, IV.84, P. Morvan (ASCC,

GDRC) 2. Rasuwa Distr., Induwa Khola Valley, Forest Camp, 1950 m, 13. IV. 85, A.

Smetana (ASCC) 1. INDIA: West Bengal: Darjeeling Dist., Rimbick-Ramam,

1950-2450 m, 19.V.75, W. Wittmer (NHMB) 1; Darjiling, 7000’, 10.VIII.09, C.

Paiva (FMNH) 1.

Geographical distribution. — Indoquedius baliyo is distributed from western

Nepal to the Darjeeling area in West Bengal (Map 27).

Bionomics. — The specimens from Gul Bhanjyang were taken by sifting leaf

litter at bases of large oak trees, those from near Malemchi were taken by sifting

sap-soaked leaf litter around a freshly storm felled large oak tree; those from Dobate

Ridge were taken by sifting moist and moldy layers of leaf litter in a semideciduous

broadleaved forest.

Revision of the Tribes Quediini and Atanygnathini

305

Comparisons. — Indoquediiis haliyo can easily be distinguished from /.

sikkimensis by the colouration alone. For comparison with I. aherransi see the latter

species.

Etymology. — The specific name is the Nepali adjective baliyo (strong); it refers

to the large and rather robust form of the species.

Bipunctatus Group

This species group is characterized by the following combination of characters:

lateral pronotal groove smooth and shiny, without any microsculpture; anterior

angles of pronotum impunctate; dorsal surface of head and pronotum without

micropunctation; large lateral puncture of pronotum touching lateral pronotal

groove; paramere of aedoeagus entire (Figs. 252, 256, 260, 264).

The group includes five species in the Himalayan area.

3. Indoquedius bipunctatus (Eppelsheim) comb. nov.

Figs. 251-254; Map 27

Quedius bipunctatus E\)^Q\s>\\t\m I895b;391; Cameron 1932:301

Description. — Piceous-black to black, abdomen iridescent; palpi, antennae and legs uniformly

rufo-testaceous. Head rounded, wider than long (ratio 1.22); eyes very large and convex, tempora three times

shorter than length of eyes seen from above (ratio 0.33); two punctures along inner margin of eye between

anterior and posterior frontal puncture; posterior frontal puncture situated quite close to posteromedian

margin of eye, usually almost touching it; one additional setiferous puncture between it and posterior margin

of head; surface of head without any microsculpture. Antenna moderately long, hardly thickened toward

apex, first three segments with long dark setae; all segments longer than wide, gradually becoming shorter,

penultimate segment at least twice as long as at base wide, last segment asymmetrically acuminate, elongate,

slightly shorter than two preceding segments combined. Pronotum short, feebly wider than long (ratio 1.10),

broadly rounded basally and only feebly narrowed anteriorly, rather strongly transversely convex; lateral

groove fine, smooth and shiny, without microsculpture; dorsal rows each with two punctures; sublateral

rows each with no more than two punctures, posterior puncture, if present, at about level of large lateral

puncture; large lateral puncture situated very close to lateral groove, distinctly touching it; anterior angles of

pronotum impunctate; surface of pronotum without any microsculpture. Scutellum, except for narrow

impunctate strip along all three margins, with coarse and dense punctation, intervals between punctures

mostly about as large as diameters of punctures. Elytra rather short, at base almost as wide as pronotum at

widest point, at suture slightly shorter (ratio 0.89), at sides about as long as pronotum at midline; punctation

and pubescence rather dense and coarse, intervals between punctures, at least on area near apex of scutellum,

no more than slightly larger than diameters of punctures; surface between punctures without microsculpture.

Wings fully developed. Abdomen with tergite 7(fifth visible) bearing whitish apical seam of palisade setae;

punctation and pubescence of tergites much finer than that of elytra, slightly denser on bases of tergites and

in general gradually becoming slightly sparser toward apex of abdomen. Front tarsus equally dilated in both

sexes.

Male. Apical margin of stemite 8 with rather shallow, wide arcuate emargination (Fig. 251), small

triangular area before emargination flattened and smooth. Aedoeagus (Figs. 252-254) moderately large;

median lobe strongly narrowed into obtuse apex. Paramere is slightly exceeding apex of median lobe, rather

wide basally but strongly narrowed to form narrow and very elongate, subparallel-sided apical portion,

bearing four fine apical setae and one stronger setae at each lateral margin below apex; underside with

numerous sensory peg setae fomiing irregular elongate group. Internal sac with central sclerite covered with

scale-like structures and bearing a pair of spur-like structures in each apico-lateral comer.

Quaest. Ent., 1988, 24 (2)

306

Smetana

Length 7. 8-8. 7 mm.

Type material. — Eppelsheim (l.c.) described the species from two specimens. I

was able to study one male specimen deposited in the Eppelsheim collection in the

Naturhistorisches Museum in Wien, Austria. It is labelled as follows: pink square

label/ “Parasnath Gebg.” / “bipunctatus Epp. India orient, leg. Waagen.” / “Je ne I’ai

pas” / “c. Eppelsh. Steind. d.” / “Typus” (red label). The specimen was dissected,

the aedaeagus and 8th sternite mounted in Canada Balsam, 8th tergite and genital

segment were glued to the plate with beetle. Left antenna, except for two basal

segments, left hind tarsus and middle right tibia and tarsus are missing and the right

elytron has a big triangular hole. The specimen is hereby designated as the lectotype

of /. bipunctatus; the label “Lectotype Quedius bipunctatus Eppelsheim Smetana

des. 1983” has been attached to it.

Geographical distribution. — Indoquedius bipunctatus is distributed in the

western portion of the Himalayan range, in Himachal Pradesh and in northern Uttar

Pradesh; also known from Parashath Hills in Bihar (Map 27).

Material studied. — 8 specimens.

INDIA. No further data (FMNH) 1. (See the discussion). Himachal Pradesh. Shatingri, Mandi, 6000’,

H.G. Champion (BMNH) 1. Uttar Pradesh. Kumaon, Haldwani Distr., H.G. Champion (ASCC, BMNH) 5.

Bionomics. — No details are known about the habitat requirements of this

species.

Comparisons. — There is one female specimen of this species in the Bernhauer

collection in the Field Museum of Natural History, Chicago, Illinois, labelled as

follows: “India orient. leg. Waagen” / “bipunctatus Epp.” / “Cotypus ex Coll.

Waagen ded. Bang Haas” / “grandiceps Kr. vgl. m. Typus in Coll. Kraatz” /

“praeditus Shp. vgl. m. Typ. Lond. dieser m. etwas dunk. FUhl. u. Beinen” /

“Chicago NH Mus. M. Bernhauer Collection”. This specimen is most likely the

second specimen of the original series.

Indoquedius praeditus is another species of this group, occurring in Japan; it

differs, in addition to the differently shaped aedoeagus, by the black antennae with

gradually paler tip, and by black legs with slightly paler tarsi. As a matter of fact, I.

bipunctatus resembles much more another Japanese species, I. juno with which it

shares the same colouration. However, the latter differs, among other characters, by

the differently shaped aedoeagus with the paramere divided into two long branches.

4. Indoquedius oculatus (Fauvel) comb.nov.

Map 27

Quedius oculatus Fauve\ 1895:274; Cameron 1932:300

Description. — In all external characters extremely similar to /. bipunctatus; different only by

slightly differently coloured appendages and in general slightly narrower and almost parallel-sided

pronotum. Antennae dark, with last 2 or 2 1/2 segments yellowish. Middle and hind tibiae darkened, middle

and hind femora slightly darkened.

Male. Unknown.

Revision of the Tribes Quediini and Atanygnathini

307

Length 8. 6-8. 8 mm.

Type material. — The Fauvel eollection in the Institut Royal des Sciences

Naturelles, Bruxelles, Belgique, contains four female specimens under the name Q.

oculatus. They are labelled as follows: Spec. No. 1: “Carin Asciuii Ghecu

1400-1500 m. L. Tea. III-IV.88” / “oculatus Fvl.” / “Syntype” / “R.LSc. N.B.

17.479 Quedius Coll, et det. A. Fauvel”. Spec. No. 2: “Carin Asciuii Ghecu

1400-1500 m. L. Fea. III-IV.88” / “Syntype” / “Coll, et det. A. Fauvel Quedius

oculatus Fauv. R.I.Sc. N.B. 17.479”. Spec. No. 3: “Carin Asciuii Ghecu 1400-1500

m. L. Fea. III-IV.88” / “Carin Ghecu 900-1 100 m. Fea” / “Syntype” / “Coll, et det.

A. Fauvel Quedius oculatus Fauv. R.I.Sc. N.B. 17.479”. Spec. No. 4: “Tonkin” /

“Syntype” / “Coll, et det. A. Fauvel Quedius oculatus Fauv. R.I.Sc. N.B. 17.479”.

All specimens are not conspecific. Specimen No. 2 seems to be identical with the

holotype of /. filicornis. Specimen No. 3 is identical with specimen No. 1 and

specimen No. 4 seems to belong to yet another species.

The first specimen is hereby designated as the lectotype of I. oeulatus; the label

“Lectotype Quedius oculatus Fauv. Smetana des. 1983” has been attached to it.

Geographical distribution. — Indocpiedius oculatus is known at present only

from Burma (Map 27).

Material studied. — 2 specimens.

BURMA. See Type material.

Bionomics. — Nothing is known about the habits of this species.

Recognition. — Indocpiedius oculatus can be distinguished from /. hipunctatus at

present only by the few differences in the colouration of the appendages given in the

key and the description. The status of this species cannot be properly assessed until

the males and their aedaeagi are available for study.

Cameron (1932:300) described the emargination of the male sternite 8 of this

species as “apex broadly but not deeply emarginate, with triangular smooth

impression in front of it”. This is apparently a translation of FauveFs (l.c.)

description “segmento 1° ventrali apice late parum profunde emarginato, post

incisuram triangulariter impresso, laevi”.

I have no information about the whereabouts of the male specimen of FauveFs

original series. The specimen must come from “Carin Cheba” (see Fauvel

1895:274).

5. Indocpiedius daai spec.nov.

Figs. 255-258; Map 28

Description. — Dark brownish-piceous to piceous black, apical margins of abdominal tergites

usually paler; palpi, antennae and legs rufo-testaceous, antennae gradually becoming paler toward apex.

Head rounded, wider than long (ratio 1.27); eyes very large and convex, tempora three times shorter than

length of eyes seen from above (ratio 0.31); two punctures (occasionally three unilaterally) along inner

margin of eye between anterior and posterior frontal puncture; posterior frontal puncture situated quite close

to posteromedian margin of eye, usually touching it; one additional setiferous puncture between it and

posterior margin of head; surface of head without any microsculpture. Antenna moderately long, hardly

Quaest. Ent., 1988, 24 (2)

308

Smetana

thickened toward apex, first three segments with long setae; all segments longer than wide, gradually

becoming shorter, penultimate segment about twice as long as wide at base, last segment asymmetrically

acuminate, elongate, slightly shorter than two preceding segments combined. Pronotum short, feebly wider

than long (1.13), broadly rounded basally and slightly narrowed anteriorly, rather strongly transversely

convex; lateral groove very fine, smooth and shiny, without microsculpture; dorsal rows each with two

punctures; sublateral rows each with no more than two very fine punctures (usually with just one puncture at

anterior margin of pronotum), posterior puncture, if present, at about level of large lateral puncture; large

lateral puncture situated quite close to lateral groove, distinctly touching it; anterior angles of pronotum

impunctate; surface of pronotum without any microsculpture. Scutellum with punctation sparser and less

dense than that of/, hipunctatus, wide areas along all three margins impunctate. Elytra rather short, at base

about as wide as pronotum at widest point, at suture feebly shorter (ratio 0.93), at sides about as long as

pronotum at midline; punctation and pubescence rather sparse and only moderately coarse, intervals between

punctures distinctly larger than diameters of punctures; surface between punctures without microsculpture.

Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae;

punctation and pubescence of tergites much finer than that of elytra, slightly denser on bases of tergites and

in general gradually becoming slightly sparser toward apex of abdomen, occasionally tergites almost

impunctate in about apical fourth. Front tarsus equally dilated in both sexes.

Male. Apical margin of stemite 8 with very shallow and moderately wide, arcuate emargination (Fig.

253), small triangular area before emargination flattened and smooth. Aedoeagus (Figs. 256-258)

moderately large; median lobe relatively wide, apical portion almost parallel-sided and apex broadly arcuate.

Paramere wide and rather short, about reaching apex of median lobe, widely rounded at apex and with short

attenuate middle portion; four rather fine apical setae and two slightly finer setae at each lateral margin just

below apex; underside with numerous sensory peg setae situated as in Fig. 257. Internal sac with central

sclerite covered with sharp spine-like structures.

Length 7. 5-8. 8 mm.

Type material. — Holotype (male) and allotype (female): “INDE Kumaon (UP)

Bhim Tal env. 1500 m 1. I. Lobl 4.X.79”. In the Museum d’Histoire Naturelle de

Geneve, Geneve, Switzerland.

Paratypes (8): INDIA: Uttar Pradesh: same data as holotype (ASCC) 1;

Chakrata, Mohna, 5000', 2.V.21, M. Cameron (BMNH) 1; Mussoorie, Ami Gad,

28.V.21, M. Cameron (BMNH, CNCC) 4. Himachal Pradesh: Simla Hills, Matiana,

7900', IX.1921, M. Cameron (BMNH) 1; Dhelu, Mandi, 4500', H.G. Champion

(BMNH) 1.

Geographical distrihution. — Indoqiiedius daai is distributed in the western

portion of the Himalayan range, in Himachal Pradesh and in Uttar Pradesh (Map

28).

Bionomics. — Little is known about the habitat requirements of this species. The

three specimens from Bhim Tal were taken on an eastern slope at the edge of a small

forest by sifting fallen leaves and debris among grasses.

Comparisons. — Indoquedius daai is in all external characters very similar to /.

hipunctatus; however, it can be distinguished by the characters used in the key, and

by the distinctly differently shaped aedoeagus. Both species occur in the western

portion of the Himalayan range and seem to be largely sympatric.

Indoquedius daai is also very similar to I. filicornis; see the description, and

comparison and variation under the latter for the distinguishing characters. Unlike I.

filicornis, /. daai seems to be rather constant in colouration.

Etymology. — The specific name is the Nepali noun daai (older brother) in

apposition; it refers to the close similarity and relationship of this species to both I.

Revision of the Tribes Quediini and Atanygnathini

309

hipunctatus and I. filicornis.

6. Indoquedius filicornis (Eppelsheim) comh.nov.

Figs. 259-262, 400-403; Map 28

Quedius filicorms 1895a:55; Cameron 1932:301

C/mr/z/zi' Champion 1922:33; Cameron 1932:301

Description. — in all external characters very similar to /. daai but different as follows: colouration

similar, however, middle and hind femora often more or less darkened and rarely apical half to 2/3 of

segment 5 and apex of abdomen yellowish-red; or pronotum entirely pale red. Scutellum in general less

punctate, often with only a few punctures. Elytra in general slightly narrower, with punctation slightly

sparser and more superficial. Abdomen with apical portions of tergites more extensively and more often

impunctate.

Male. Emargination of sternite 8 similar to that of I. daai (Figs. 259). Aedoeagus (Figs. 258, 261)

slightly more elongate than that of I. daai\ median lobe with apex more arcuately rounded. Paramere usually

slightly exceeding apex of median lobe, longer and narrower than that of /. daai. narrowly rounded at apex

and with long attenuate middle portion; four rather fine apical setae and one (occasionally two) slightly finer

seta at each lateral margin below apex; underside with less numerous sensory peg setae near apex extended

backward along both lateral margins (Figs. 260-262). Internal sac similar to that of /. daai.

Length 6. 0-8. 8 mm.

Type material. — Indoquedius filicornis. The Eppelsheim collection in the

Naturhistorisches Museum, Wien, Austria, contains one female specimen (holotype

- see Eppelsheim 1895:55) under the name Q. filicornis. It is labelled as follows:

pink square label/ “Himalaja Sikkim”/ “88”/ “filicornis Epp.” / “c. Eppelsh.

Steind.d.’V “Typus” (white label)/ “Typus” (red label). The specimen was received

with head and thorax broken off and glued back; only four basal segments of each

antenna are present.

Indoquedius conicus. The collection of the British Museum (Natural History),

London, contains one male specimen (holotype) under the name Q. conicus. It is

labelled as follows: “c?” / “Type H.T.” (round label with red margin)/ “Ranikhet,

Kumaon, H.G.C.”/ “Brit. Mus. 1922-71” / “Quedius (Raphirus) conicus Champ.”

(printed label)/ “Quedius (Raphirus) conicus Champ.” (handwritten label)/ “E.M.M.

1922- det. G.C.C.”. The specimen cannot be specifically distinguished from the

holotype of I. filicornis; the name I. conicus is a junior synonym of I . filicornis.

Geographical distribution. — Indoquedius filicornis is widely distributed, from

Uttar Pradesh in the west through Nepal to Sikkim (Map 28).

Material studied. — 54 specimens.

INDIA. Sikkim. See Type material. Uttar Pradesh. Kumaon: Chaubattia nr. Ranikhet, about 1800 m,

12-13.X.79, I. Lobl (CNCC, MHNG, ASCC) 7; same 1950 m, 14.X.79, I. Fbbl (MHNG) 2; Rangarh, about

2000 m, 9.X.79, I. Fobl (ASCC, CNCC, MHNG) 1 1; Rangarh, crest, 2250 m, 9.X.79, I. Lobl (MHNG) 2.

Garhwal: au-dessus Joshimath, 2100 m, 27.X.79, 1. Lobl (MHNG) I.

NEPAL. Niiwakot Distr. Malemchi, 2800 m, 14.IV.81, Lobl & Smetana (ASCC, CNCC, MHNG) 9;

Malemchi Khola below Malemchi, 2100 m, 15. IV. 81, Lobl & Smetana (ASCC) 1. Khandhari Distr. Forest

above Ahale, 2400 m, 25. III. 82, A. & Z. Smetana (ASCC) I; Pass NE Mangmaya, 2300 m, 6. IV. 84,

Smetana & Lobl (ASCC) 1; Forest S Mansingma, 2200 m, II. IV. 84, Smetana & Lobl (ASCC) 2; Induwa

Khola Valley, 2000 m, 16.IV.84, Smetana & Lobl (ASCC, CNCC, MHNG) 4; same, 2100 m, 17.IV.84,;

same 2150 m, 18. IV. 84; all Smetana & Lobl (ASCC, CNCC, MHNG) 5. Manang Distr. Marsyandi below

Bagarchhap, 2100 m, 12. IV. 80, Martens & Ausobsky (SBMF) 1. Mustang Distr. nr. Lethe, 2460-2600 m.

Quaest. Ent., 1988, 24 (2)

310

Smetana

30.1V-1.V.80, Martens & Ausobsky (CNCC, SBMF) 3; same, 2550 m, 2.X.83, Smetana & Lobl (ASCC) 1.

Rasuwa Distr. 1.5 km NE Bhargu, 12.V.85, A. Smetana (ASCC) 1.

Bionomics. — Specimens of /. filicornis were collected at elevations between

1800 and 2600 m, mostly by sifting old vegetation, debris and fallen leaves along

old stone walls around fields, along edges and in clearings in forests by sifting fallen

leaves, occasionally also in floor litter in Finns excelsa forests.

Comparisons and variations. — Indoqnedins filicornis is in all external

characters quite similar to /. daai; the distinguishing characters given in the

description are subject to some variability and may not be always reliable. However,

both species are easily distinguished by the differences in the shape of the

aedoeagus which are quite constant.

Almost all specimens with darkened femora came from the western portion of

the distributional range (Uttar Pradesh); the specimens with yellowish-red tip of

abdomen came from central Nepal and have all uniformly rufo-testaceous legs. The

specimens with pale red pronotum are at present known only from Induwa Khola

valley in eastern Nepal; they were more abundant there than specimens with the

body entirely black.

7. Indoqnedins saatlii spec.nov.

Figs. 263-266; Map 28

Descnption. — In all external characters extremely similar to specimens of I. filicornis with entirely

black body and entirely rufo-testaceous legs, and different mainly by differently shaped aedoeagus. Elytra

slightly shorter than in average specimens of I. filicornis, at suture distinctly (ratio 0.81), at sides feebly

(ratio 0.92) shorter than pronotum at midline.

Male. Apical margin of stemite 8 with shallow arcuate emargination similar to that of I. filicornis (Fig.

263). Aedoeagus (Figs. 264-266) similar to that of I. filicornis but distinctly smaller; paramere smaller and

narrower, not quite reaching apex of median lobe, with middle portion almost parallel-sided; four rather fine

apical setae and two unequally long setae at each lateral margin slightly below apex (in one specimen one

additional, minute seta present unilaterally); underside with about 15 sensory peg setae forming a rather

solid apical group, not extended backward along lateral margins. Internal sac similar to that of /. daai and I.

filicornis (Figs. 258, 261 ).

Length 7.8-8. 1 mm.

Type material. — Holotype (male) and paratype (male): “INDIA Meghalaya

Khasi Hills 25 or 30.X.78 Shillong, 1850-1950 m Besuchet & Lobl”. Holotype

(date 25.x. 78) in the Museum d’Histoire Naturelle de Geneve, Geneve, Switzerland;

paratype in my collection (ASCC).

Geographical distribntion. — Indoqnedins saathi is at present known only from

the type locality in Khasi Hills in the state of Meghalaya of India (Map 28).

Bionomics. — The two specimens of the original series were taken on Shillong

Peak by sifting forest floor litter.

Comparisons. — Indoqnedins saathi can be distinguished positively from I.

filicornis only by the differently shaped aedoeagus. It probably is endemic to the

Khasi Hills, separated from the main Himalayan range.

Revision of the Tribes Quediini and Atanygnathini

311

Etymology. — The specific name is the Nepali noun saathi (friend). It refers to

the close similarity and relationship of this species to I. filicornis.

Ahervans Group

This species group is characterized by the following combination of characters:

lateral pronotal groove with dense, almost granulose microsculpture, opaque;

anterior angles of pronotum finely punctate; dorsal surface of head and pronotum

with micropunctation; paramere of aedoeagus entire (Fig. 268).

The group includes one species in the Himalayan region.

8. Inoquedius aherrans (Cameron) comb. now

Figs. 267-269; Map 28

Quedius aherrans Cameron 1932:299

Description . — Head and pronotum dark brunneo-piceous, elytra reddish-yellow, abdomen brown,

slightly iridescent; both maxillary and labial palpi brownish-piceous, with last segments paler, antennae

piceous, gradually becoming yellowish toward apex, legs testaceo-brunneous with slightly darkened femora

(see comparisons). Head rounded, slightly wider than long (ratio 1.11); eyes very large and convex, tempora

almost three times shorter than length of eyes seen from above (ratio 0.39); two punctures along inner

margin of eye between anterior and posterior frontal puncture; posterior frontal puncture situated close to

posteromedian margin of eye, separated from it by distance no larger than diameter of puncture; one

additional setiferous puncture between it and posterior margin of head; area behind posterior frontal

puncture with numerous fine punctures; surface of head without any micro.sculpture, but with scattered

microscopic punctures. Antenna fairly long, hardly thickened toward apex, first three segments with rather

long dark setae; all segments much longer than wide, gradually becoming shorter, penultimate .segment

almost three times as long as at base wide, last segment asymmetrically acuminate, only slighty longer than

penultimate segment. Pronotum relatively nanow, as long as wide (seemingly longer than wide), broadly

rounded basally and parallel-sided, moderately transversely convex; lateral groove with dense, almost

granulose microsculpture, opaque; dorsal rows each with two punctures; sublateral rows each with two

punctures, posterior puncture situated slightly before level of large lateral puncture; large lateral puncture

situated very close to lateral pronotal groove, distinctly touching it; surface of pronotum without any

microsculpture, but with scattered microscopic punctures becoming coarser on anterior angles, anterior

angles therefore finely punctate. Scutellum very large, entire surface densely punctate and pubescent. Elytra

at suture feebly longer (ratio 1.05), at sides somewhat longer (ratio 1.23) than pronotum at midline;

punctation and pubescence dense, interspaces between punctures no more than slightly larger than diameters

of punctures; surface between punctures without microsculpture. Wings fully developed. Abdomen with

tergite 7 (fifth visible) bearing whitish apical seam of palisade .setae; punctation and pubescence of

abdominal tergites dense but distinctly finer than that on elytra, gradually becoming finer and sparser toward

apex of abdomen.

Male. First four segments of front tarsus strongly dilated. Apical margin of stemite 8 with wide and not

deep, arcuate emargination (Fig. 267), triangular area before emargination flattened and smooth. Aedoeagus

(Figs. 268, 269) large and elongate; median lobe slightly attenuate in middle portion, nanowed into long,

obtusely acute apex. Paramere very narrow and elongate, not reaching apex of median lobe, narrowly

spatulate apically; with four small apical setae and a pair of slightly stronger setae at each lateral margin

below apex; underside with only two sensory peg setae below apex.

Length 10.0 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains one male specimen (holotype) under the name Q. aherrans. It is

labelled as follows; ‘Type” (round label with red margin)/ “Sikkim; Gopaldhara,

Quae St. Ent., 1988, 24 (2)

312

Smetana

Map 27. Distribution records for: Indoquedius sikkimensis ( • ); / haliyo ( A ); /. bipunctatus ( ■ ); and I. ocidatus ( ).

Revision of the Tribes Quediini and Atanygnathini

313

Quaest. Ent., 1988, 24 (2)

Map 28. Distribution records for: Indoquedius daai { • ); I.filicornis ( A ); / saathi { ■ ); and I. aberrans ( ^ ).

314

Smetana

Rungbong Vail. H. Stevens.” / “H. Stevens. Brit. Mus. 1922-307.” / “Quedius

aberrans Cam. TYPE”.

The specimen was dissected, the sternite 8 and the aedoeagus were mounted in

Canada Balsam, and the genital segment on plate with beetle.

Geographical distribution. — Indoquedius aherransi is at present known only

from the type locality in Sikkim (Map 28).

Material studied. — Holotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons and variations. — Indoquedius aherransi can easily be

distinguished from both I. sikkimensis and I. haliyoi, in addition to the distinctive

aedoeagus, mainly by the relatively narrow pronotum with lateral groove

microsculptured and therefore opaque, by the anterior angles finely punctate and by

the longer and densely punctate elytra.

The holotype of this species is a teneral specimen; the colouration described

above is therefore certainly atypical. I am reasonably sure that /. aberrans is in fact

black with black antennae becoming gradually yellowish toward apex, and with

piceous-black legs with paler tarsi and possibly also tibiae.

3. Genus Bolitogyrus Chevrolat

Bolitogyriis Chevrolal 1842:641; Blackwelder 1952:1 15

Cyrtothorax Kvaatz 1858:336; Fauvel, 1878:163; Cameron 1932:277; Scheerpeltz 1974:175-192.

Type species of Bolitogyrus: Bolitogyrus cribripennis Chevrolat 1842 (= B.

buphthalmus (Erichson 1840)), designated by Chevrolat (l.c.), by monotypy.

Type species of Cyrtothora.x: Quedius buphthalmus Erichson 1840, designated

by Lucas 1920:222, by subsequent designation.

Descriptive notes.— The genus Bolitogyrus shares many characters with

Quedius but it differs in several respects. The head always bears two or three

setiferous punctures between anterior and posterior frontal punctures situated

directly at median margin of eye. The eyes are always very large and convex,

considerably longer than the tempora. The frons behind the antennal insertions bears

a V-shaped impression. The dorsal surface of the head lacks microsculpture and is

therefore highly polished. On the antenna the first five segments lack the fine and

dense pubsescence and the large antennal setae, particularly those past segment 3,

are relatively long and strong. The pronotum is no more than slightly narrowed

anteriorly and the dorsal rows of pronotum are each reduced to just one puncture

situated close to the anterior margin of pronotum. Posterolateral and basal margins

of pronotum are strongly and abruptly explanate. The dorsal surface of pronotum

lacks microsculpture and is highly polished. Scutellum is punctate. Elytra are more

or less coarsely and usually to a variable extent irregularly punctate. Wings are fully

developed. The first four segments of the front tarsus are strongly dilated, more so in

the males. The middle tibia is not spinose on its lateral face. The seventh (fifth

Revision of the Tribes Quediini and Atanygnathini

315

visible) abdominal tergite bears always a whitish apical seam of palisade setae. The

female tergite 8 (sixth visible) bears usually a deep and narrow median notch. The

male genital segment has tergite 10 rounded apically (Fig. 271) and sternite 9

emarginate or subemarginate apically (Fig. 272). The aedoeagus, including the

parameres, is similarly developed as in the genus Quedius; the parameres are fused

in one solid sclerite (Fig. 274).

The genus Bolitogyrus is a tropical genus represented by numerous species in the

tropics of both the Old and New Worlds.

One species of Bolitogyrus is at present known from the Flimalayan region (see

taxonomic notes following B. vulneratus).

Comparisons and variations. — Some species of the genus Indoquedius

(particularly those from the Sikkimensis-Group) resemble in general habitus the

species of the genus Bolitogyrus, but they differ by a number of characters (see the

key and the respective descriptions).

Blackwelder (1952:1 15) considered the name Cyrtothorax as a junior synonym

of Bolitogyrus and the latter name was subsequently used for this genus by

Hammond (1984:205). The name Bolitogyrus was first published (as a nomen

nudum, since the only included species “5. crihripennis Chevrolat” was not valid)

by DeJean (1837:76) in his catalogue. Later on Chevrolat (1842:641) stated under

Bolitogyrus that the species B. crihripennis he named (no description was ever

published) was sent to Erichson, who declared it identical with his “Quedius

bupthalmus” {-Cyrtothorax huphthalmus (Erichson 1840)). Based on the above

statement of Chevrolat, Blackwelder (l.c.) attributed the genus Bolitogyrus to

Chevrolat, argued that the above Chevrolat’s statement validated the name

Bolitogyrus and consequently put the junior name Cyrtothorax in synonymy with

Bolitogyrus. This concept was later followed by Hammond (1984:205).

The genus Bolitogyrus badly needs a modern revision, based on secondary

sexual characters of the species and the shapes of their aedoeagi. The key to all

known species of Bolitogyrus presented by Scheerpeltz (1974:176-183) is based

extensively on the colouration of the body; yet the colour may distinctly vary within

one species, as even my limited material seems to indicate (see also the taxonomic

notes following B. vulneratus).

1. Bolitogyrus vulneratus (Eauvel) (comb.nov.)

Eigs. 270-274; Map 29

Cyrtothorax vulneratus Fauvel 1878:165; Cameron 1932:277

Description. — Piceous-black, abdomen slightly iridescent, namowly explanate margins of pronotum

rufo-testaceous, each elytron with obliquely transverse, rufo-testaceous spot slightly constricted toward

lateral elytral margin and connected via deflexed portion of elytron with similarly coloured, small humeral

spot; medioapical angle of elytron rufo-testaceous, apical margin narrowly, lateroapical angle of elytron

more broadly testaceous; apical margins of abdominal pleurites indistinctly paler; both maxillary and labial

palpi testaceous; antennae piceous, first segment pale testaceous with apex slightly darkened, segments 2-5

testaceous, apices of segments 2 and 3 slightly darkened; legs testaceous, hind and middle coxae and

Quaest. Ent., 1988, 24 (2)

316

Smetana

posterior base of front coxae piceous. Head wider than long (ratio 1.45) with very large and convex, bulging

eyes; tempora slightly more than 6 times shorter than length of eyes seen from above; dorsal surface with

sparsely and somewhat irregularly dispersed fine punctures, without microsculpture. Antenna short, segment

1 shorter than two following segments combined, segments 2 and 3 subequal in length, segment 2 slightly

stronger than 3, segments 4 and 5 somewhat longer than wide, segment 5 slightly shorter and wider than 4,

segments 6-10 gradually becoming shorter and wider, segments 9 and 10 slightly transverse, last segment

about as long as 2 preceding segments combined. Pronotum indistinctly wider than head (ratio 1.06) and

feebly wider than long (ratio 1.13), broadly rounded basally and indistinctly narrowed anteriorly, strongly

transversely convex; surface without microsculpture. Scutellum with several coarse punctures. Elytra short,

at suture as long as, at sides slightly longer than pronotum at midline; punctation deep and very coarse and

irregular, leaving median and lateral portion of pale transverse anterior spot and pale posterior markings on

each elytron impunctate; surface between punctures without microsculpture. Wings fully developed.

Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; bases of tergites 1^

densely and fairly coarsely punctate, discs impunctate, tergite 5 sparsely punctate, punctures elongate;

surface of tergites with excessively fine and dense microsculpture of transverse lines.

Male. Apical margin of stemite 7 distinctly, arcuately emarginate, apical margin of sternite 8 with

shallow and inconspicuous arcuate emargination (Fig. 270). Aedoeagus (Figs. 273, 274) narrow and

elongate; median lobe suddenly narrowed into minute apical portion, with apical portion slightly excavate on

surface adjacent to paramere; paramere elongate, fusiform, narrower than median lobe and slightly

exceeding apex of median lobe, with 6 long apical setae; sensory peg setae on underside of paramere

numerous, forming two rows quite close to lateral margins.

Fength 6. 7-8. 5 mm.

Type material. — Fauvel (1878:165) described the species from one single

specimen. The holotype in the Fauvel collection in the Institut Royal des Sciences

Naturelles de Belgique, Bruxelles, Belgique, is labelled as follows: “Cochinchine” /

“Tonkin” (rest illegible)/ “vulneratus Fvl.” / “TYPE” (red square label)/ “R.I.Sc.

N.B. 17.479 Cyrtothorax Coll, et det. A. Fauvel”. The specimen is a female (not a

male as stated by Fauvel, l.c.) in fair condition; only three basal segments remain

from the left antenna. The remaining 5 specimens under the name B. vulneratus do

not belong to the original series (see also taxonomic notes following B. vulneratus).

Geographical distribution. — Bolitogyrus vulneratus is distributed in eastern

portion of the Himalaya, from eastern Nepal through Darjeeling area; also in Khasi

Hills in Meghalaya (Map 29), and in Vietnam.

Material studied. — 5 specimens.

INDIA. Meghalaya. Khasi Hills, Mawsynram-Balat, 1000 m, 27.X.78, Besuchet-Fobl (MHNG) 1. West

Bengal. Darjeeling Distr., Sevoke, 200 m, 7.X.78, Besuchet-Fobl (MHNG) 1.

NEPAF. Khandbari Distr. Val. Arun ss/Num, 1050 m, 22. IV. 84, Fobl-Smetana (MHNG) 1; Arun

Valley at Num main bridge, 1050 m, 2. IV. 84, Smetana & Fobl (ASCC) 1.

VIETNAM. See Type material.

Bionomics. — The specimens from the Arun Valley were collected by sifting

various mushrooms, moss and debris around them on larger fallen trees. Those from

Khasi Hills and Darjeeling distr. were taken by sifting fallen leaves in forest ravines.

Taxonomic notes. — Bolitogyrus vulneratus is quite distinctive among the

Himalayan Quediini by its general habitus and by the distinctive colour pattern of

the very coarsely and deeply punctate elytra.

Bolitogyrus vulneratus is apparently a complex of species. But since only very

limited material is available for study and more material is essential, I have decided

not to deal with this problem at present. The female holotype (see under Type

317

Revision of the Tribes Quediini and Atanygnathini

Quaest. Ent., 1988, 24 (2)

Map 29. Distribution records for: Bolitogyrus vulneratus ( • ); and Ctenandropus nigriceps ( A ).

318

Smetana

material) has a notch in the middle of the apical margin of the abdominal tergite 8;

the female from the Arun river valley near Num in Nepal (MHNG) does not have

this notch. The additional five specimens under the name B. vulneratus in the Fauvel

collection (see Type material) come from Burma: “Carin Cheba 900-1 100 m. L.Fea

V.XII-88” (Sdd, 1$) and ‘Tennaserim. Kawkareet Fea. Maggio 1887” (Id). They

belong to at least two different species. They show intraspecific variability in

colouration; the association of any of these specimens with the female holotype is

very difficult.

The Fauvel collection contains another specimen (female) bearing the label

“Khasia Hills VII.” (apparently Khasi Hills in Meghalaya). It is similarly coloured

as the holotype of B. vulneratus, but it is at least twice as big and represents

obviously yet another species.

4. Genus Heterothops Stephens

Heterothops Stephens 1829:23; Cameron 1932:265^

Type species.— Staphy linns hinotatus Gravenhorst 1802, designated by

Westwood 1838:16, by subsequent designation.

I am not presenting here a conventional formal description of this genus; it can

be found in some recent papers, e.g., Smetana 1971:14-15.

Male. The aedoeagus of all species lacks parameres (Figs. 277, 291, 302).

Genital segment with tergite 10 and sternite 9 rounded apically, styli of tergite 9

relatively strong, each with numerous very long apical setae (Fig. 297).

Female genital segment as in Figs. 404^07, tergite 10 obtusely rounded or

subtruncate apically.

The genus is distributed worldwide. Nine species are known to occur in the

Himalayan region.

Taxonomic notes. — All species from the studied area bear two setiferous

punctures between the anterior and posterior frontal punctures; the position of the

posterior one seems to be a good distinguishing character for the species groups. It

is in correlation with the size of the eyes.

Coiffait (1978:7) erected for this genus a separate subtribe Heterothopsi, based

on the absence of the parameres on the aedoeagus in all species. He also included

the genus Atanygnatlms in this subtribe, because the aedoeagus of all species also

lacks parameres. Not only is the absence of parameres insufficient ground for

establishing a subtribe in this case, but also the two genera Heterothops and

Atanygnatlms are not even remotely related. The loss of the parameres occurred

independently. For more details see Smetana (1984) and the discussion following

the tribe Atanygnathini in this paper.

^Only references pertaining to the area treated in this revision are given. A complete synonymy with all

references up to 1970 can be found in Smetana 1971:14.

Revision of the Tribes Quediini and Atanygnathini

319

The species Heterothops flavicoUis Motschulsky 1858, described from “Indes

orientales” and treated by Cameron (1932:266), belongs in fact to the genus

Gahronthus Tottenham (1955) (comb.nov.) and is identical with Gahronthus

pulchellus (Kraatz 1859) which becomes a junior synonym of it {syn.nov.)

(Hammond, pers. comm. August 1984).

Key to species of Heterothops

1 Eyes large to very large, at least as long as tempora seen from

above. Posterior setiferous puncture between anterior and

posterior frontal punctures situated anteriad of level of posterior

margin of eye or rarely posteriad of it by about 1 1/2 diameter

of puncture 2

T Eyes small, distinctly shorter than tempora seen from above

(ratio 0.64). Posterior setiferous puncture betweeen anterior and

posterior frontal punctures situated far posteriad of level of

posterior margin of eye, distance equal to about one third length

of eye. Length 4.4 mm 8. H. saano spec.nov., p. 329

2 (1) Eyes very large, at least twice as long as tempora seen from above 3

2' Eyes large, less than twice as long as tempora, rarely about as

long as tempora seen from above 5

3 (2) Eyes more than twice as long as tempora seen from above (ratio

2.62-2.87). Aedoeagus very elongate and narrow (Eigs. 277.

278,282) 4

3' Eyes twice as long as tempora seen from above (ratio 2.0).

Aedoeagus less elongate and wider (Eig. 285), sclerites of

internal sac as in Eigs. 286, 287. Length 3. 1-4.9 mm

3. H. pusillus Coiffait, p. 323

4 (3) Apical portion of median lobe evenly narrowed toward apex

(Eigs. 277, 278), sclerites of internal sac as in Figs. 279, 280.

Eyes slightly larger (ratio length of eyes: length of tempora =

2.87). Microsculpture on head and pronotum slightly coarser

and less dense. In general slightly smaller species. Length

4. 1-4.9 mm 1. H. oculatus Fauvel, p. 320

4' Apical portion of median lobe rather suddenly attenuate toward

apex (Fig. 282), sclerites of internal sac as in Fig. 283. Eyes

slightly smaller (ratio length of eyes: length of tempora = 2.62).

Microsculpture on head and pronotum slightly finer and denser.

In general slightly larger species. Length 4. 5-5.0 mm

2. H. indicus Cameron, p. 322

5 (2') Head and pronotum with distinct microsculpture 6

5' Head and pronotum without appreciable microsculpture.

Quaest. Ent., 1988, 24 (2)

320

Smetana

Aedoeagus and internal sac as in Figs. 302, 303. Length 3.9

mm 9. H. saphaa spec.nov., p. 330

6 (5) Elytra not appreciably paler at apical margin. Punctation of

elytra and abdominal tergites very fine and very dense.

Aedoeagi and sclerites of internal sacs as in Figs. 289-292 7

6' Elytra narrowly paler at apical margin. Punctation of elytra and

abdominal tergites very fine but less dense. Aedoeagi and

sclerites of internal sacs different 8

7 (6) Median lobe of aedoeagus strongly and rather suddenly

attenuate anteriorly (Fig. 289); sclerites of internal sac as in

Fig. 290. Microsculpture on head and pronotum fine. Length

4. 3-4. 6 mm 4. H. hindustanus Cameron, p. 325

7' Median lobe of aedoeagus moderately and gradually narrowed

anteriorly (Fig. 291); sclerites of internal sac as in Fig. 292.

Microsculpture on head and pronotum coarse. Length 4.4 mm ..

5. H. khairo spec.nov., p. 326

8 (6') Internal sac of aedoeagus, when evaginated, with a pair of

highly sclerotized sclerites along lateral margins of median lobe

(Figs. 299, 300). Length 4.0-5. 1 mm

7. H.fvanzi Coiffait, p. 328

8' Internal sac of aedoeagus, when evaginated, without a pair of

highly sclerotized sclerites along lateral margins of median lobe

(Figs. 294, 295). Length 4.2-5.0 mm

6. H. persimilis Cameron, p. 326

Oculatus Group

This species group is characterized by the following combination of characters:

eyes very large, at least twice as long as tempora seen from above; head and

pronotum with distinct microsculpture; posterior puncture between anterior and

posterior frontal punctures situated anteriad of level of posterior margin of eye;

aedoeagus elongate and narrow (Figs. 277, 278, 282, 285).

The group contains three species in the Himalayan region.

1 . Heterothops oculatus Fauvel

Figs. 275-280; Map 32

Heterothops oculata Fauvel 1895:275; Cameron 1932:267 (ex parte)

Description. — Black, elytra and abdomen sometimes becoming indistinctly paler toward apex;

labial and maxillary palpi and antennae piceous-brown to piceous, basal two or three antennal segments

slightly pale; legs dark brown to piceous-brown with paler tarsi and medial face of hind tibiae more of less

darkened. Head rounded, feebly wider than long (ratio 1.10); eyes large and convex, almost three times as

long as tempora (ratio 2.87); posterior setiferous puncture between anterior and posterior frontal punctures

Revision of the Tribes Quediini and Atanygnathini

321

situated well before level of posterior margin of eye; one puncture inediad of posterior frontal puncture,

temporal puncture situated very close to posterior margin of eye, separated from it by distance no larger than

diameter of puncture; surface with moderately fine and dense microsculpture of transverse waves becoming

coarser anteriorly. Antenna moderately long, segments 2 and 3 subequal in length, segment 2 slightly

stronger than segment 3, segments 4-6 longer than wide, gradually becoming shorter and wider, segments

7-10 about as long as wide, segment 11 as long as two preceding segments combined. Pronotum feebly

wider than long (ratio 1.14), widely rounded basally and distinctly nairowed anteriorly, evenly transversely

convex; large lateral puncture separated from lateral pronotal groove by distance about equal to diameter of

puncture; surface of pronotum with microsculpture similar to that on head, but indistinctly finer and denser.

Scutellum punctate. Elytra at base about as wide as pronotum at widest point, slightly widened posteriorly, at

suture as long as, at sides slightly longer than pronotum at midline (ratio 1.20); punctation fine and rather

sparse, interspaces between punctures along longitudinal axis 2-3 times larger than diameters of punctures;

pubescence piceous-black. Wings fully developed. Abdomen with tergite 7 (fifth visible) bearing distinct

whitish apical seam of palisade setae; punctation slightly finer than that on elytra, fairly dense on bases of

tergites, gradually becoming sparser toward apical margin of each tergite; pubescence black.

Male. First four segments of front tarsus slightly dilated; apical margin of sternite 8 with small, obtusely

triangular to obtusely arcuate emargination (Figs. 275, 276). Aedoeagus (Figs. 277, 278) very elongate and

narrow; median lobe evenly narrowed toward apex; sclerites of internal sac as in Figs. 279, 280.

Fength 4. 1^.9 mm.

Type material. — The Fauvel collection in the Institut Royal des Sciences

Naturelles de Belgique, Bruxelles, contains three specimens under the name H.

oculata. They are labelled as follows: Spec. No. 1; ‘Tenasserim M. Mooleyit

1800-1900 m. Tea. Marzo 1887” / “oculatus Fvl.” / “Syntype” / “R.I.Sc.N.B. 17.479

Heterothops Coll, et det. A. Fauvel”. Spec. No. 2 and 3: exactly same first label as

Spec. No. 1/ “Syntype” / “Coll, et det. A. Fauvel Heterothops oculatus Fauv.

R.l.Sc.N.B. 17.479”.

The first two specimens are females, the third specimen is a male; it was

dissected, and the aedoeagus and sternite 8 were mounted in Canada Balsam. The

male specimen is hereby designated as the lectotype of H. oculatus', the label

“Lectotype Heterothops oculatus Fauvel Smetana des. 1983” has been attached to it.

Geographical distribution. — Heterothops oculatus is at present known from

central Nepal (Kathmandu Valley and Phulcoki) (Map 32), and from Tenasserim in

Burma.

Material studied. — 9 specimens.

BURMA. See Type material.

NEPAF. Kathmandu Distr. Gokama Forest, 1400 m, 31.111.81, Fobl & Smetana (ASCC, MHNG) 4;

same, 20.X.83, Smetana & Fdbl (MHNG) 1. Lalitpur Distr. Phulcoki, 2550 m, 21. IV. 82, A. & Z. Smetana

(ASCC) 1.

Bionomics.— The specimens from the Gokarna Forest were taken by sifting leaf

litter around bases of large trees. The specimen from Phulcoki was taken by sifting

old dead vegetation, various debris and fallen leaves at bases of large rocks on a

clearing in a forest.

Comparisons. — Heterothops oculatus has the largest eyes of all Heterothops

species known to occur in the Himalayan area. Only H. indicus comes close in this

respect; however, both species can usually be distinguished without difficulties by

external characters only (see the description of H. indicus)', the aedoeagi and the

sclerites of the internal sacs are also distinctly different. In addition, both species are

Quaest. Ent., 1988, 24 (2)

322

Smetana

(at least at present) allopatric, with H. indicus occurring only in western part of the

Himalaya (see Map 30). Cameron (1932:267) failed to distinguish H. indicus from

H. oculatus and considered it as a junior synonym of the latter species; consequently

his records (l.c.) from Dehra Dun, Mussoorie, Chakrata and Simla Hills refer to H.

indicus and not to H. oculatus.

There are some differences between the emargination of the stemite 8, the shape

of the median lobe and the development of the sclerites of the internal sac between

the lectotype and one male from the Gokarna Forest (see Figs. 277-280). However,

the differences do not seem to warrant taxonomic separation, at least not at present

when only a limited material is available for study.

2. Heterothops indicus Cameron stat.nov.

Figs. 281-283; Map 30

Heterothops indica Cameron 1926:365

Heterothops ocidatcr, Cameron 1932:267 (ex parte)

Description. — in all characters very similar to H. oculatus. but different as follows: average size

slightly larger; colouration paler: piceous, elytra narrowly paler, almost testaceous at posterior margin, apical

margins of abdominal tergites and apex of abdomen more or less paler, rufo-brunneous; both labial and

maxillary palpi testaceous, antennae piceous with three basal segments testaceous, legs uniformly pale

testaceous, except hind tibiae slightly darkened medially in some specimens. Head somewhat smaller and

indistinctly narrower, almost as long as wide; eyes slightly smaller, less than three times as long as tempora

(ratio 2.62); posterior setiferous puncture between anterior and posterior frontal punctures situated just

before level of posterior margin of eye; surface with microsculpture finer and denser than that of H.

oculatus. Antenna somewhat shorter. Surface of pronotum with microsculpture slightly finer and denser.

Punctation of elytra and abdominal tergites usually somewhat finer, pubescence paler, rather dark brown.

Male. First four segments of front tarsus dilated in a similar way as in H. oculatus', emargination in

middle of apical margin of stemite 9 (Fig. 281) similar to that of H. oculatus. Aedoeagus (Figs. 282, 283)

similar to that of H. oculatus', however, apical portion of median lobe rather suddenly attenuate toward apex;

sclerites of internal sac as in Fig. 283.

Length 4. 5-5.0 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains seven specimens under the name H. indica. They are labelled as

follows: Spec. No. 1 (male): “SYN-TYPE” (round label with blue margin)/ “Type”

(round label with red margin)/ “Mossy Falls. Mussoorie.” / “Dr. Cameron.

27.III.1921” / “Type Heterothops indica Dr. Cameron” / “M. Cameron.

Bequest.B.M. 1955-147”. Spec. No. 2 (male): “SYN-TYPE” (round label with blue

margin)/ “Co-type” (round label with yellow margin)/ “Heterothops indica Cam.” /

“Ami Gad. Mussoorie. Dr. Cameron. 16. 10. 1921”. Spec. No. 3-5 (all females):

“SYN-TYPE” (round label with blue margin)/ “Ami Gad. Mussoorie.” / “Dr.

Cameron. 1 2. IV.21.” / “M. Cameron. Bequest.B.M. 1955-147”. Spec. No. 6

(female): “SYN-TYPE” (round label with blue margin)/ “Chakrata Dist. Korawa

Khud 9100'” / “Dr. Cameron.4.V.22.” / “M. Cameron. Bequest.B.M. 1955-147”.

Spec. No. 7 (female): “Fagu 8000' Simla Hills.” / “SYN-TYPE” (round label with

blue margin)/ “Dr. Cameron. IX. 1921.” / “Heterothops indica Cam.” / “H.G.

Champion Coll. B.M.1953-156”.

Revision of the Tribes Quediini and Atanygnathini

323

Both male specimens were dissected and aedoeagi and sternite 8 of the first

specimen were mounted in Canada Balsam. The first male specimen is hereby

designated as the lectotype of H. indicus\ the label “Lectotype Heterothops indica

Cameron Smetana des. 1983” has been attached to it.

Geographical distribution. — Heterothops indicus is distributed in the western

part of the Himalaya (Himachal Pradesh and Uttar Pradesh) (Map 30).

Material studied. — 34 specimens.

INDIA. Himachal Pradesh. Simla, E.C. Ansorge (BMNH) 1. Uttar Pradesh. Chakrata Distr.: Korawa

Khud, 9100', 4.V.22, Cameron (BMNH) 3; Jadi Gad, 7000', 9.V.22, Cameron (BMNH) 2. Miissoorie Distr.

Keyarkuli, 17. IV. 22, Cameron (ASCC, BMNH) 9; Aglar River, 16. IV. 22, Cameron (BMNH) 1; Mussoorie,

1700 m, 19.x. 79, 1. LobI (MHNG) 1. Kumaon: W. Almora, H.G. Champion (BMNH) 2. Garhwal: 22 km N

Rishikesh, 450 m, 30.X.79, 1. Lobl (ASCC, MHNG) 5; 16 km de Srinagar, 550 m, 29.X.79, 1. Lbbl (MHNG)

1; below Bhatwari, 1500 m, 24.X.79, I. Lbbl (MHNG) 1; 2 km E. Dhanolti, 2250 m, 21.X.79, I. Lbbl

(MHNG) 1.

Bionomics. — The specimens from Mussoorie and Garhwal were collected by

sifting wet debris among reeds at a river, in debris on a river bank, by sifting fallen

leaves and debris in a forest ravine and by sifting fallen leaves and moss near a

seepage in a mixed Rhododendron-Ahies forest.

Recognition. — Heterothops indicus can be distinguished fairly easily from H.

oculatus by the characters given in the key and in the description. Both species

differ also in their distributional ranges (see the comparisons under H. oculatus for

details).

Cameron (1932:267) failed to distinguish H. indicus from H. oculatus; he listed

the name as a junior synonym of the latter species (see also under H. oculatus).

3. Heterothops pusillus Coiffait

Figs. 284-287; Map 31

Heterothops piisilla Coiffait 1982a:83

Description. — In all external characters similar to both H. oculatus and H. indicus, but different as

follows: piceous to piceous-black, elytra sometimes slightly paler at posterior margin, apical margins of

abdominal tergites and apex of abdomen often somewhat paler; both labial and maxillary palpi brownish to

brownish-piceous, antennae piceous with two basal segments usually at least partially paler, legs

brunneo-testaceous with posterior tibiae slightly darkened medially. Head about same as that of H. indicus,

but eyes smaller, twice as long as tempora (ratio 2.0). Microsculpture on head and pronotum same as that of

H. indicus. Punctation and pubescence of elytra and abdominal tergites not appreciably different from that of

H. indicus.

Male. Eirst four segments of front tarsus dilated in a similar way as in both compared species;

emargination in middle of apical margin of sternite 8 (Eig. 284) distinctly deeper than those of H. oculatus

and H. indicus. Aedoeagus (Figs. 285-287) less elongate and wider; median lobe almost parallel-sided in

middle portion, with apical portion rather suddenly attenuate toward subacute apex; sclerites of internal sac

as in Figs. 286, 287.

Length 3. 1-4.9 mm.

Type material. — The male holotype in the Franz collection, Modling, Austria, is

labelled as follows: “Umg. Pina 29.9-1.10.72” / “Gebiet des Rarasees Westnepal, Ig.

H. Franz” / “TYPE” / “Heterothops pusilla H. Coiffait det. 1979”. The specimen

was dissected, the aedoeagus and the 8th sternite were mounted in Canada Balsam.

Quaest. Ent., 1988, 24 (2)

324

Smetana

Geographical distribution. — Heterothops pusillus is widely distributed in the

Himalaya, from Himachal Pradesh and Uttar Pradesh through Nepal to Darjeeling

area in West Bengal (Map 31 ).

Material studied. — 41 specimens.

INDIA. Himachal Pradesh. Katrain near Kulu, H. Franz (HFCC) 1. Uttar Pradesh. Kumaon: Rangarh,

2400 m, 10.X.79, I. Lobl (MHNG) 1; Rangarh, crete, 2250 m, 9.X.79, I. Lobl (MHNG) 1; Chaubattia nr.

Ranikhet, about 1800 m, 12-13. X. 79, I. Lobl (MHNG) 1. West Bengal. Darjeeling Distr.: Algarah-Labha,

1900 m, 11.X.78, Besuchet-Lobl (MHNG) 1; 13 km N Ghoom, 1500 m, 15.X.78, Besuchet-Lobl (MHNG)

1; Ghoom-Lopchu, 2000 m, 12 or 14.X.78, Besuchet-Lobl (MHNG) 4.

NEPAL. Chaubas, 2600 m, 4 and 5.IV.81, Lobl & Smetana (ASCC, MHNG) 6; Malemchi, 2800 m, 14

and 17.IV.81, Lobl & Smetana (ASCC, CNCC) 5; Dobate Ridge NE Barahbise, 2700 m, 2. and 7.V.81, Lobl

& Smetana (ASCC) 4. Khandhari Distr. Kuwapani, 2100 m, 28. III. 82, A. & Z. Smetana (ASCC) 1; Forest

above Ahale, 2400 m, 25. III. 82, A. & Z. Smetana (ASCC) 2; Forest NE Kuwapani, 2500 m, 12. or 14. IV. 82,

A. & Z. Smetana (ASCC) 2; Induwa Khola Valley, 2100 m, 17. or 18. IV. 84, Lobl & Smetana (ASCC,

MHNG) 4. Manang Distr. Eorest W Bagarchhap, 2200 m, 2250 m, 2400 m, 21.22. and 23. IX. 83, Smetana &

Lobl (ASCC, MHNG) 4; Latha Manang W Bagarchhap, 2400 m, 23.IX.83, Smetana & Lobl (MHNG) 1.

Parhat Distr. Ghoropani Pass, N slope, 2800 m, 5.X.83, Smetana & Lobl (ASCC) 1. Weg von Pokhara z.

Goropani, H. Franz (HFCC) 1.

Bionomics. — Heterothops pusillus seems to occur mainly in forested habitats at

elevations between 1500 m and 2800 m. Most specimens were taken by sifting leaf

litter, various debris and moss on the floor of semideciduous broad leaved forests,

sometimes in rather wet habitats, such as seepages, etc.

Comparisons. — The size of the eyes and the emargination of the male sternite 8

are the best external characters for distinguishing H. pusillus from both H. oculatus

and H. indicus (see the key and the respective descriptions). Heterothops pusillus

also resembles H. franzi; see the comparisons under the latter species for

distinguishing characters of these two species.

There is a chance that two species are in fact combined here under this name.

The specimens from the Darjeeling district seem to have a slightly different

aedoeagus; however, since the available aedoeagi are not preserved well enough, I

did not treat the Darjeeling specimens as taxonomically different.

Hindustanus Group

This species group is characterized by the following combination of characters:

eyes large, but less than twice as long as tempora seen from above (ratio 1.40-1.66);

head and pronotum with distinct microsculpture; posterior puncture between

anterior and posterior frontal punctures situated at level of posterior margin of eye;

punctation of elytra and abdominal tergites very fine and very dense; elytra not paler

at apical margin.

The group contains two species in the Himalayan region.

Revision of the Tribes Quediini and Atanygnathini

325

4. Heterothops hindustamis Cameron

Figs. 288-290; Map 31

Heterothops hindiistana Cameron 1932:267

Description. — Piceous-black, elytra brunneo-piceous, apical margins of abdominal tergites and

apex of abdomen indistinctly paler; both labial and maxillary palpi brunneous, antennae piceous with three

basal segments feebly paler, legs brunneotestaceous, middle and hind tibiae slightly darkened medially.

Head rounded, vaguely wider than long (ratio 1.07); eyes large and convex, longer than tempora (ratio 1.4);

posterior setiferous puncture between anterior and posterior frontal punctures situated at level of posterior

margin of eye; one puncture mediad of posterior frontal puncture; temporal puncture separated from

posterior margin of eye by distance slightly longer than two diameters of puncture; surface with fine and

superficial, moderately dense microsculpture of transverse and oblique waves. Antenna moderately long,

segments 2 and 3 subequal in length, segment 2 slightly stronger than segment 3, segments 4 and 5 longer

than wide, segment 6 slightly longer than wide, segments 7-10 about as long as wide, gradually becoming

slightly wider, last segment as long as two preceding segments combined. Pronotum feebly wider than long

(ratio 1.10), widely rounded basally and distinctly narrowed anteriorly, evenly transversely convex; large

lateral puncture touching lateral pronotal groove; surface of pronotum with microsculpture similar to that on

head. Scutellum rather elongate, densely punctate. Elytra at base about as wide as pronotum at widest point,

slightly widened posteriorly and relatively long, at suture slightly longer (ratio 1.16), at sides distinctly

longer (ratio 1.34) than pronotum at midline; punctation very fine and very dense, punctures often almost

touching each other along transverse axis; pubescence dark, brownish-piceous. Wings fully developed.

Abdomen with tergite 7 (fifth visible) bearing distinct whitish apical seam of palisade setae; punctation

about equally fine and dense as that on elytra, gradually becoming sparser toward apical margin of each

tergite; pubescence dark, brownish-piceous.

Mate. First four segments of front tarsus slightly dilated; apical margin of stemite 8 with small, obtusely

triangular emargination (Fig. 288). Aedoeagus (Figs. 289, 290) with median lobe parallel-sided in middle

part and then strongly and rather suddenly attenuate toward apex, sclerites of internal sac as in Fig. 290.

Length 4.3— 4.6 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains one female specimen under the name H. hindustanus. It is labelled

as follows: “SYN-TYPE” (round label with blue margin)/ “TYPE” (round label with

red margin)/ “Dr. Cameron 4.V.22.” / “M. Cameron. Bequest. B.M. 1955-147” /

“Chakrata Dist. Korawa Khud 91007 “H. hindustanus Cam. TYPE”. The specimen

is hereby designated as the lectotype of H. hindustanus', the label “Lectotype

Heterothops hindustanus Cameron Smetana des. 1983” has been attached to it.

Geographical distribution. — Heterothops hindustanus is at present known only

from Uttar Pradesh and western Nepal (Map 31 ).

Material studied. — 2 specimens.

INDIA. Uttar Pradesh. See Type material.

NEPAL. NordI Dhaulagiri, Gompa/Tarakot, 3300-3400 m, 2-6. VI. 73, J. Martens (SBMF) 1.

Bionomics. — Nothing is known about the habitat requirements of this species.

Recognition. — Heterothops hindustanus is well characterized among the species

with microsculptured head and pronotum and with the eyes less than twice as long

as tempora, by the rather long, unicoloured and very finely and densely punctate

elytra, by the temporal puncture on the head situated rather far from posterior

margin of the eye (see the description) and by the characters on the aedoeagus and

internal sac (Figs. 289, 290).

Quaest. Ent., 1988, 24 (2)

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Smetana

The aedoeagus and sternite 8 were drawn from the male specimen from Nepal,

which was compared with female lectotype.

5. Heterothops khairo spec.nov.

Figs. 291,292; Map 31

Description. — in all external characters very similar to H. hindustanus but different as follows:

head with eyes slightly longer, ratio of length of eyes: length of tempora = 1.66; microsculpture on head and

pronotum coarse, however, that of pronotum slightly less coarse than that on head.

Male. First four segments of front tarsus slightly less dilated than those of H. hindustanus', sternite 8

with apical emargination similar to that of H. hindustanus. Aedoeagus (Figs. 291, 292) with median lobe

distinctly narrower, moderately and gradually narrowed toward apex, sclerites of internal sac as in Fig. 292.

Length 4.4 mm.

Type material. — Holotype (male); INDIA; “Dehra Dun Dr. M. Cameron 4/3

1921.” / “M. Cameron. Bequest.B.M. 1955-147”. In the British Museum (Natural

History), London.

Geographical distribution. — Heterothops khairo is at present known only from

the type locality in Uttar Pradesh (Map 31 ).

Bionomics. — The holotype was taken in a termite nest.

Comparisons. — Heterothops khairo differs from the two following species H.

persimilis and H.franzi with similar ratio of length of eyes to the length of tempora,

by the larger and wider head, by the larger pronotum, by the distinctly coarser

microsculpture on the head and pronotum, by the longer and distinctly more densely

punctate elytra, etc.

Etymology. — The specific name is the Nepali adjective khairo (brown); it refers

to the general colour of the species.

Persimilis Group

This species group is characterized by the following combination of characters;

eyes large, but less than twice as long as or rarely about as long as tempora seen

from above; head and pronotum with distinct microsculpture; posterior puncture

between anterior and posterior frontal punctures situated at level of posterior margin

of eye or posteriad of it by about 1 1/2 diameter of puncture; punctation of elytra

and abdominal tergites very fine but moderately dense; elytra narrowly paler at

apical margin.

The group contains two species in the Himalayan region.

6. Heterothops persimilis Cameron

Figs. 293-295, 404^07; Map 32

Heterothops persimilis Cameron 1932:267

Description. — Brunneo-piceous, piceous to piceous-black, elytra paler, rather dark brunneous and

paler at apical margin, apical margins of abdominal tergites and apex of abdomen paler, usually

Revision of the Tribes Quediini and Atanygnathini

327

rufo-brunneous; both labial and maxillary palpi brunneous, antennae testaceo-brunneous to dark brown, legs

testaceo-brunneous, hind tibiae slightly darkened medially. Head rounded, vaguely wider than long (ratio

1.08); eyes large and convex, longer than tempora (ratio 1.53); posterior setiferous puncture between anterior

and posterior frontal punctures situated at level of posterior margin of eye; temporal puncture separated from

posterior margin of eye by distance slightly larger than diameter of puncture; one puncture mediad of

posterior frontal puncture; surface with fine and dense microsculpture of transverse and oblique waves.

Antenna moderately long, segments 2 and 3 subequal in length, segment 2 slightly stronger than .segment 3,

segments 4 and 5 slightly longer than wide, segment 6 hardly longer than wide, segments 7-10 about as long

as wide to slightly transverse, last segment slightly shorter than two preceding segments combined.

Pronotum feebly wider than long (ratio 1.11), widely rounded basally and moderately naiTowed anteriorly,

evenly transversely convex; large lateral puncture separated from lateral pronotal groove by distance about

equal to diameter of puncture; surface of pronotum with microsculpture similar to that on head, but slightly

finer. Scutellum punctate. Elytra moderately long, at base about as wide as pronotum at widest point, slightly

widened posteriorly, at suture about as long as, at sides longer (ratio 1.26) than pronotum at midline;

punctation fine but not very dense, interspaces between punctures along longitudinal axis 2-3 times larger

than diameters of punctures; pubescence dark brown. Wings fully developed. Abdomen with tergite 7 (fifth

visible) bearing distinct whitish apical seam of palisade setae; punctation somewhat finer than that on elytra,

dense on bases of tergites but becoming much sparser toward apical margin of each tergite; pubescence

brownish piceous.

Male. First four segments of front tarsus slightly dilated; apical margin of stemite 8 with small, obtusely

triangular emargination (Fig. 293). Aedoeagus (Figs. 294, 295) with median lobe almost parallel-sided in

middle portion, with apical portion rather suddenly narrowed toward subacute apex; sclerites of internal sac

as in Fig. 295, no pair of highly sclerotized sclerites along lateral margins of median lobe when internal sac

evaginated.

Length 4. 2-5.0 mm.

Type material. — The original series in the British Museum (Natural History),

London, contains two male specimens under the name H. persimilis. They are

labelled as follows: Spec. No. 1: “SYN-TYPE” (round label with blue margin)/

“Type” (round label with red margin)/ “Kotgarh 7000' Simla Hills” / “Dr. Cameron

IX.1922” / “H. persimilis Cam. TYPE” / “M. Cameron. Bequest. B.M. 1955-147”.

Spec. No. 2; “SYN-TYPE” (round label with blue margine)/ “Kotgarh 7000' Simla

Hills” / “Dr. Cameron IX.1922” / “M. Cameron. Bequest. B.M. 1955-147” /

“persimilis” (handwritten label). Both specimens were dissected, and aedoeagi and

8th sternites were mounted in Canada Balsam. Specimen No. 1 is hereby designated

as the lectotype of H. persimilis', the label “Lectotype Heterothops persimilis

Cameron Smetana des. 1983” has been attached to it.

Geographical distribution. — Heterothops persimilis is distributed from

Himachal Pradesh and Uttar Pradesh to eastern Nepal (Map 32).

Material studied. — 19 specimens.

INDIA. Himachal Pradesh. Simla Hills, Narkanda, 9239', 15.IX.21, Cameron (BMNH) 1. Uttar

Pradesh. Chakrata Distr.: Bodyar, 8200’, 8-12.V.22, Cameron (BMNH) 1; Khedar Khud, 7500’, 11.V.22,

Cameron (BMNH) 1. Mussoorie; Dhobi Ghat, 14. IV. 22, Cameron (BMNH) 1. Kumaon; W. Almora, H.G.

Champion (BMNH) 2.

NEPAL. Khandhari Distr. Forest NE Kuwapani, 2450 m, 13. IV. 82, A. & Z. Smetana (ASCC) 1; same,

2250 m, 24. IV. 84, Lbbl & Smetana (MNHG) 1. Lalitpur Distr. Phulcoki, 20. and 21. IV. 82, 2550 or 2600 m,

A. & Z. Smetana (ASCC) 3; same, 28., 29. and 30.IV.84, 2500-2550 m, Smetana & Lobl (ASCC, MHNG)

6.

Bionomics. — In Nepal, specimens of this species were taken by sifting leaf litter

and other debris on forest floor on moist places, particularly in the oak forest on

Phulcoki, also by sifting moist moss on large rocks (Eorest NE Kuwapani) or by

Quaest. Ent., 1988, 24 (2)

328

Smetana

sifting dead vegetation, moss and debris at bases of large rock blocks in forest

clearings (Phulcoki).

Comparisons. — Heterothops persimilis resembles in general habitus and

colouration H. indicus. Both species are sympatric in Himachal Pradesh and Uttar

Pradesh, but H. persimilis differs, in addition to the differences on the aedoeagus, by

the smaller eyes and by the different position of the posterior setiferous puncture

between the anterior and posterior frontal punctures (see the respective

descriptions).

7. Heterothops franzi Coiffait

Figs. 296-300; Map 30

Heterothops franzi Coiffait 1982a:82

Description. — In all characters extremely similar to H. persimilis but different as follows: in

general darker, piceous to piceous-black, abdominal tergites uniformly dark or with only feebly paler apical

margins; antennae piceous with basal 1 to 3 segments only vaguely paler, legs pale brunneous to brunneous,

hind tibiae distinctly darkened medially. Head narrower, about as long as wide, eyes smaller, ratio of length

of eyes to length of tempora = 1.25, in some specimens tempora almost as long as length of eyes seen from

above; posterior setiferous puncture between anterior and posterior frontal punctures situated at about level

of posterior margin of eye or posteriad of it by about 1 1/2 diameter of puncture.

Male. First four segments of front tarsus dilated in similar way as in H. persimilis, apical margin of

stemite 8 (Fig. 296) with emargination similar to that of H. persimilis. Aedoeagus (Figs. 298-300) very

similar to that of H. persimilis but with attenuate apical portion longer; sclerites of internal sac as in Figs.

299, 300, with a pair of highly sclerotized sclerites along lateral margins of median lobe when internal sac

evaginated.

Length 4.0-5. 1 mm.

Type material. — The male holotype in the Franz collection, Modling, Austria, is

labelled as follows: “Umg. Pina 29.9.-1.10.72” / “Gebiet des Rarasees Westnepal,

Ig.H. Franz” / “TYPE” / “Heterothops franzi H. Coiffait det. 1979”. The specimen

was dissected and the aedoeagus and stemite 8 were mounted in Canada Balsam.

I have not seen the female paratype deposited in the Coiffait collection (Museum

National d’Histoire Naturelle, Paris).

Geographical distribution. — Heterothops franzi is distributed in western part of

the Himalaya, from Uttar Pradesh to about central Nepal (Map 30).

Material studied. — 12 specimens.

INDIA. Uttar Pradesh. Garhwal, 10 km E Dhanolti, 2450 m, 21.X.79, 1. Lobl (MHNG) 2.

NEPAL. Gurjakhani, 83°14'E, 28°37'N, 8500', 4-7.VII.54, K.H. Hyatt (BMNH) 1; Chaubas, 2500 m,

4.IV.81, Lobl & Smetana (ASCC) 1; Malemchi, 2800 m, 17.IV.81, Lobl & Smetana (ASCC) 1. Lalitpur

Distr. Phulcoki, 2600-2650 m, 21-22. III. 80, Martens & Ausobsky (SBME) 2; same, 2600 m, 20. and

22.IV.82, A. & Z. Smetana (ASCC, MHNG) 3; same, 2550 m, 29.IV.84, Smetana & Lobl (ASCC) 1.

Bionomics. — Specimens of H. franzi were taken by sifting forest floor litter, leaf

litter and other debris in depressions of forest floor, around bases of large trees, etc.

Some were also taken by sifting wet fermenting wood cuttings.

Comparisons. — Heterothops franzi is rather difficult to distinguish from H.

persimilis; the best external distinguishing character seems to be the smaller size of

the eyes in H. franzi (see the respective descriptions). Both species occur together in

Revision of the Tribes Quediini and Atanygnathini

329

some localities (Phulcoki).

Saano Group

The single species of this group shares most characters of the Persimilis Group

but differs as follows: eyes small, distinctly shorter than tempora seen from above;

posterior puncture between anterior and posterior frontal punctures situated far

posteriad of level of posterior margin of eye.

The species group may prove to be superfluous when the species of Heterothops

from the studied area will become better known.

8. Heterothops saano spec.nov.

Map 30

Description. — - Piceous, pronotum dark brown, elytra brown with suture and apical margin

testaceous, apical margins of abdominal tergites and apex of abdomen distinctly paler, testaceo-rufous; both

labial and maxillary palpi testaceous, antennae testaceo-brunneous with first three segments testaceous, legs

testaceous with bases of femora indistinctly darkened. Head relatively narrow, as long as wide, almost

parallel-sided; eyes small and hardly convex, not protruding from lateral contours of head, distinctly shorter

than tempora seen from above (ratio 0.64); posterior setiferous puncture between anterior and posterior

frontal punctures situated far posteriad of level of posterior margin of eye, distance equal to about one third

of length of eye; one puncture mediad of posterior frontal puncture; temporal puncture situated closer to

posterior margin of head than to posterior margin of eye; surface of head with dense and very fine

microsculptue of transverse and oblique waves. Antenna rather short, segments 2 and 3 subequal in length

but segment 2 somewhat stronger, segment 4 slightly longer than wide, segment 5 about as long as wide,

segments 6-10 gradually becoming shorter, segments 8-10 transverse, segment 1 1 about as long as two

preceding segments combined. Pronotum wider than long (ratio 1.16), widely rounded basally, distinctly

narrowed anteriorly, evenly transversely convex; large lateral puncture separated from lateral pronotal

groove by distance about equal to diameter of puncture; surface of pronotum with microsculpture similar to

that on head. Scutellum sparingly punctate. Elytra moderately long, at base slightly narrower than pronotum

at widest point, slightly widened posteriorly, at suture as long as, at sides slightly longer (ratio 1.18) than

pronotum at midline; punctation moderately dense, interspaces between punctures along longitudinal axis

about twice as large as diameters of punctures; pubescence dark brown. Wings fully developed. Abdomen

with tergite 7 (fifth visible) bearing distinct whitish apical seam of palisade setae; punctation of tergites

somewhat finer than that on elytra, dense on tergal bases and becoming slightly sparser toward apical

margins; pubescence dark brown.

Male. Unknown.

Length 4.4 mm.

Type material. — Holotype (female): INDIA: “Kashmir Aru Ig. H. Franz, Okt.

1977”. In the Franz collection, Modling, Austria.

Geographical distribution. — Heterothops saano is known only from the type

locality in Kashmir (Map 30).

Bionomics. — No details are known about the habitat requirements of this

species.

Recognition. — Heterothops saano is unique among the Himalayan species due

to the small size of the eyes and the position of both the posterior setiferous

puncture between anterior and posterior frontal punctures (situated far posteriad of

the posterior margin of eye) and the temporal puncture (situated very far from the

Quaest. Ent., 1988, 24 (2)

330

Smetana

posterior margin of eye - see the description).

Etymology. — The specific name is the Nepali adjective saano (small). It refers to

the small size of the eyes.

Saphaa Group

The single species of this species group is characterized by the lack of any

microsculpture on head and pronotum.

9. Heterothops saphaa spec. now

Figs. 301-303; Map 32

Description. — Black, elytra piceous-black with feebly paler apical margin, apical margins of

abdominal tergites and apex of abdomen vaguely paler; both labial and maxillary palpi brunneopiceous,

antennae piceous, first three segments vaguely paler, legs brunneous with paler tarsi, middle and posterior

tibiae darkened medially. Head relatively narrow, as long as wide, rounded; eyes moderately large, larger

than tempora seen from above (ratio 1.62); posterior setiferous puncture between anterior and posterior

frontal punctures situated just anteriad of posterior margin of eye, one puncture mediad of posterior frontal

puncture; temporal puncture separated from posterior margin of eye by distance slightly larger than diameter

of puncture; surface of head without any microsculpture. Antenna rather short, segments 2 and 3 subequal in

length but segment 2 somewhat stronger, segment 4 slightly longer than wide, segments 3 and 6 as long as

wide, segments 7-10 indistinctly wider than long, last segment as long as two preceding segments

combined. Pronotum as long as wide, widely arcuate basally and only moderately narrowed anteriorly,

evenly transversely convex; large lateral puncture almost touching pronotal lateral groove; surface of

pronotum without any microsculpture. Scutellum densely punctate. Elytra long, at base as wide as pronotum

at widest point, slightly widened posteriorly, at suture slightly (ratio 1.20), at sides distinctly longer (ratio

1.40) than pronotum at midline; punctation fine and not dense; interspaces between punctures along both

transverse and longitudinal axes about twice as large as diameters of punctures; pubescence brown. Wings

fully developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae;

punctation of tergites finer than on elytra, dense on tergal bases but becoming distinctly sparser toward

apical margins of tergites and toward apex of abdomen in general; pubescence piceous.

Male. First four segments of front tarsus dilated; apical margin of stemite 8 with small, obtusely

triangular emargination (Fig. 301). Aedoeagus (Figs. 302, 303) fairly short; median lobe parallel-sided in

middle portion, gradually and evenly narrowed toward relatively acute apex; sclerites of internal sac as in

Fig. 303.

Fength 3.9 mm.

Type material. — Holotype (male): Nepal: “148 Manang Dist. Marayandi

3800^100 m oberhalb Manang Richtung Thorung, Gebiish Martens & Ausobsky

19 Apr. 80”. In the Senckenberg Museum, Frankfurt am M., Federal Republic of

Germany.

Geographical distribution. — Heterothops saphaa is known only from the type

locality near Manang (Map 32).

Bionomics. — The holotype was taken from under a rock in rather dry, open

growths of Juniperus, Berheris and Rosa (Martens, in litt.).

Recognition. — Heterothops saphaa is unique among the Himalayan species due

to the absence of microsculpture dorsally on the head and pronotum, in combination

with the rather long elytra.

331

Revision of the Tribes Quediini and Atanygnathini

Quaest. Ent., 1988, 24 (2)

Map 30. Distribution records for: Heterothops indicus ( • ); H. franzi ( ▲ ); and H. saano ( ^ ).

332

Smetana

Map 31. Distribution records for; Heterothops piisillus { % Y, H. hindustaniis ( A ); and H. khairo ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

333

Quaest. Ent., 1988, 24 (2)

Map 32. Distribution records for: Heterothops persimilis (•);//. ociilatiis ( A ); and H. saphaa ( ^ ).

334

Smetana

Etymology. — The specific name is the Nepali adjective saphaa (clean). It refers

to the shiny surface of the head and pronotum due to the lack of microsculpture.

5. Genus Ctenandropus Cameron

Ctenandropiis Cameron 1926:348; 1932:268

Type species. — Ctenandropus nigriceps Cameron 1926, designated by Cameron

(1926:348) by original designation and monotypy.

Cameron (1926:348 and 1932:268) gave a detailed description of this genus and

I therefore present here only some additions to his description.

Descriptive notes. — Body rather flat. Head without infraorbital ridge. Male

genital segment with both tergite 10 and sternite 9 weakly sclerotized, sternite 9

obtusely rounded apically; tergite 10 with membranous, rounded apical lobe; styli of

tergite 9 strong and wide, each with numerous, very long setae (Fig. 306);

aedoeagus without paramere (Fig. 307).

At present, only one species, C. nigriceps, is included in Ctenandropus; it is

widely distributed in the Oriental region and reaches its northern distributional

limits in the Siwalik Range. However, according to A.F. Newton, Jr. (personal

communication), there is at least one additional species in this genus, C. magniceps

(Bernhauer 1920) (comb.nov.), distributed in Australia, Philippines, Sumatra and

Fiji, and originally described by Bernhauer in the genus Heterothops.

The genus Ctenandropus seems to belong to the group of the south temperate

Quediini. The ventral comb of the first segment of the middle tarsus is of the same

type found in a number of Australian species of Quediiis {sensu lato), as mentioned

by Lea (1925:240) (A.F. Newton, Jr., personal communication). The extension of

this group into mainland Asia is quite surprising.

1. Ctenandropus nigriceps Cameron

Figs. 304-307; Map 29

Ctenandropus nigriceps Cameron 1926:348; 1932::268

Description. — Head piceous-black to black, pronotum, elytra and abdomen testaceous, abdominal

segments 4—6 or 5 and 6 slightly darkened, elytra occasionally indistinctly darkened; mouthparts, antennae

and legs testaceous. Head of obtusely rectangular shape, parallel-sided and without narrowed neck,

indistinctly wider than long (ratio 1.09); eyes small, tempora about 1.5 times longer than diameter of eyes

seen from above; no additional punctures between anterior frontal punctures, posterior frontal puncture close

to line of neck, separated from it by distance slightly larger than diameter of puncture; one puncture

posteromedially of posterior frontal puncture; temporal puncture separated from posterior margin of eye by

distance of about 2 1/2 times diameter of puncture; surface of head with dense and extremely fine

microsculpture of irregular transverse waves. Antenna with segment 3 somewhat narrower and indistinctly

shorter than segment 2, segments 4 and 5 slightly longer than wide, segment 6 and 7 about as long as wide,

outer segments slightly transverse, last segment almost as long as two preceding segments combined.

Pronotum fairly flat, about as long as wide, broadly rounded basally, almost parallel-sided except narrowed

in posterior third; dorsal rows each with 2 punctures; no sublateral rows of punctures; large lateral puncture

separated from lateral margin by distance about equal to diameter of puncture; microsculpture of pronotum

still somewhat finer and denser than that on head. Scutellum finely punctate. Elytra moderately long, at

Revision of the Tribes Quediini and Atanygnathini

335

suture about as long as, at sides somewhat longer than pronotum at midline (ratio 1.14); surface finely and

densely punctate and pubescent; surface between punctures with extremely fine, irregular microsculpture.

Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; punctation and

pubescence similar to that on elytra, gradually becoming sparser toward apex of abdomen; surface between

punctures with extremely fine and dense microsculpture of transverse lines.

Male. First segment of middle tarsus with comb of dense, stiff and black setae ventrally. Apical margin

of stemite 8 with deep and rather wide obtusely rounded emargination (Fig. 305). Apical margin of tergite 8

with shallow emargination (Fig. 304). Aedoeagus (Fig. 307) small and narrow, with apical portion strongly

narrowed into quite sharp apex.

Length 2. 5-2. 8 mm.

Type material. — The collection of the British Museum (Natural History),

London, contains seven specimens belonging to the original material of C.

nigriceps.. They are labelled as follows: Spec. No. 1 (male): “Type H.T.” (round

label with red margin)/ “SYN-TYPE” (round label with blue margin)/ “Under Bark”

/ “Asarori Siwaliks. Dr. Cameron.29.X.1922.” / “TYPE Ctenandropus nigriceps Dr.

Cameron” / “M. Cameron. Bequest. B.M. 1955-147”. Spec. Nos. 2-5 (all females):

“Asarori Siwaliks.” / “Dr. Cameron.29.X.1922.” / “M. Cameron.

Bequest.B.M. 1955- 147.” / “SYN-TYPE” (round label with blue margin). Spec. No.

6 (male): “Timli, Siwaliks.” / “Dr. Cameron.27.XI.21.” / “M. Cameron.

Bequest.B.M. 1955- 147.” / “SYN-TYPE” (round label with blue margin). Spec. No.

7 (female): “Under Bark” / “Port Blair, Andaman Islds. Dr. Cameron” / “M.

Cameron. Bequest.B.M. 1955-147.” / “SYN-TYPE” (round label with blue margin).

The first male is hereby designated as the lectotype of C. nigriceps. The label

“Lectotype Ctenandropus nigriceps Cameron Smetana des. 1984” has been attached

to it.

Geographical distribution. — Ctenandropus nigriceps is widely distributed in the

Oriental region, particularly in the Malay Peninsula, Sumatra and the Philippine

Islands. I have also seen specimens from Marianas Islands (Tinian IsL, Saipal IsL,

Guam IsL), Caroline Islands (Ponape IsL, Moy Mog IsL) and from Fiji (Taveuni).

From the Himalayan region known only from the Siwalik Range in Uttar Pradesh

(Map 29).

Material studied. — 20 specimens.

INDIA. Uttar Pradesh. See Type material.

Bionomics. — Ctenandropus nigriceps lives under the bark of trees and is

apparently slow in its motions (Cameron 1932:269).

Recognition. — Ctenandropus nigriceps is quite conspicuous due to its general

habitus, particularly the rather flat body, and the wide head that is not constricted

posteriorly. It cannot be confused with any other quediine occurring in the

Himalayan region.

6. Genus Paratolmerus Cameron

Paratolmerus Cameron 1932; 169

Type species. — Paratolmerus pilosiventris Cameron 1932, designated by

Cameron (1932:169), by monotypy.

Quaest. Ent., 1988, 24 (2)

336

Smetana

The formal description of the genus was given by Cameron (l.c.). I therefore

present here only some additional characters not mentioned by Cameron (l.c.).

First segment of antenna with fine and dense pubescence. Mandibles large,

sickle-shaped as those of Anchocerus but symmetrical, each with two large and

rather sharp teeth on medial margin. Gula very short, gular sutures fused.

Infraorbital ridge fine but complete. Basal portion of head in front of neck not

abruptly and strongly depressed. Prothoracic hypomera considerably less inflexed

than in Acylophorus, except basally, and therefore distinctly visible in lateral view.

Prosternum relatively large, with short and sharp intercoxal process slightly curved

ventrally. All tibiae without spines on lateral face, except middle tibia with one or

two fine and short spines there. Dorsal side of last four segments of middle and hind

tarsus smooth and without setae except for long bristles at distal margin of each

segment. Front claws dinstinctly longer than middle and posterior claws.

Penultimate segment of middle and hind tarsus with a pair of very long apical setae

exceeding apex of last tarsal segment. Empodial setae of middle and hind tarsus

short, hardly visible between claws and considerably shorter than claws. No male

secondary sexual characters on abdominal sternites or tergites. Male genital segment

with tergite 10 and sternite 9 rounded apically, styli of tergite 9 strong and wide,

each with relatively fine apical spine (Fig. 308).

Paratolmerus is a monobasic genus from Sikkim and eastern Nepal.

Taxonomic notes. — Cameron (1932:169) assigned the genus Paratolmerus to

the tribe Staphylinini, subtribe Staphylini and compared it to the genus Tolmerinus

Bemhauer 1923. However, the presence of an infraorbital ridge on the head, the

configuration of the antennae, the mandibles and the prostemal area, and all the

characters on the legs and on the abdomen, including the male genital segment

confirm the assignment of Paratolmerus in the tribe Quediini and in the

''Acylophorus- lineage”.

1. Paratolmerus pilosiventris Cameron

Figs. 308-311; Map 33

Paratolmerus pilosiventris Cameron 1932: 169

Description. — Dark reddish-piceous to piceous, pronotum slighty paler, rather dark reddish-brown;

mouthparts testaceous; antennae and legs reddish-brown, tarsi indistinctly paler toward apex. Head orbicular,

feebly longer than wide (ratio 1.08); eyes small and flat, tempora about twice as long as length of eyes seen

from above (ratio 2.08); entire surface of head covered with fine and very dense punctation, except for

elongate-oval area on vertex; each puncture bearing fine seta; impunctate area on vertex without any

microsculpture. Antenna long, geniculate; first segment extremely long, slightly longer than four following

segments combined; segments 2 and 3 very long, subequal in length, each almost twice as long as segment

four, segments 5-8 longer than segment four, more than twice as long as at apex wide, segment 8 about 1.5

times as long as at apex wide, segment 10 yet somewhat shorter, last segment about as long as segment 10.

Pronotum feebly longer than wide (ratio 1.10), rather strongly rounded anteriorly, slightly narrowed

posteriorly, with lateral margins feebly concave behind middle and with basal margin shallowly emarginate

in middle portion; surface without any microsculpture. Scutellum large, densely and fairly coarsely punctate

and pubescent. Elytra moderately long, at base about as wide as pronotum at widest point, slightly dilated

Revision of the Tribes Quediini and Atanygnathini

337

posteriorly, at suture about as long as, at sides somewhat longer (ratio 1.17) than pronotum at midline;

punctation moderately coarse, very dense and slightly asperate, pubescence stiff, long and very dense. Wings

fully developed. Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of palisade setae;

punctation of abdominal tergites very fine, very dense on basal portion of first visible tergite but becoming

distinctly sparser toward apical margin of the tergite and toward apex of abdomen in general, fifth visible

tergite therefore only sparsely punctate; apical margins of all tergites with long and stiff, closely set setae;

pubescence long and semi-erect, particularly conspicuous on paratergites.

Male. Aedoeagus (Figs. 309-311) small; apical margin of median lobe obtusely angulate; paramere

relatively large and wide, divided into two fairly narrow branches widely separated from each other; sensory

peg setae forming a very close group on ventromedian portion of each branch; internal sac simple and

inconspicuous (Figs. 309-31 1 ).

Length 7.0-8. 0 mm.

Type material. — The Cameron collection in the British Museum (Natural

History), London, contains one female specimen under the name Paratolmerus

pilosiventris. It is labelled as follows: “Type” (round label with red margins)/

“Sikkim: Gopaldhara Rungbong Vail. H. Stevens” / “H. Stevens. Brit.

Mus. 1922-307.” / “Paratolmerus pilosiventris Cam. TYPE”. Left front tibia and

tarsus are missing. The genital segment was dissected and glued to plate with beetle.

The specimen is hereby designated as the lectotype of P. pilosiventris; the label

“Lectotype Paratolmerus pilosiventris Cameron Smetana des. 1985” has been

attached to it.

Geographical distribution. — Paratolmerus pilosiventris is known from the

eastern portion of the Himalaya, from eastern Nepal and Sikkim (Map 33).

Material studied. — 3 specimens.

INDIA. Sikkim. See the Type material.

NEPAL. Khandhan Distr. Below Sheduwa, 2100-2550 m, 9.IV.82, A. & Z. Smetand (ASCC) 2.

Bionomics. — The specimens from below Sheduwa were taken in a small creek

on a steep slope from very wet debris composed mainly of rotting twigs.

Recognition. — Paratolmerus pilosiventris is quite conspicuous due to its general

habitus, resembling the genus Acylophorus, the long and strongly geniculate

antennae, and the conspicuously pubescent abdomen.

7. Genus Acylophorus Nordman^

Acylophorus tSordmann 1837: 127; Cameron 1932:302.

Type species. — Acylophorus ahrensii Nordmann 1837 (=Staphylinus

glaherrimus Herbst 1784), designated by Blackwelder 1943:466, by subsequent

designation.

I am not presenting here a conventional formal description of this genus; it can

be found in Cameron (1932:302) and in some recent papers, e.g., Smetana

1971:246. I only mention some additional characters not given in previous

descriptions, or important for distinguishing the genera of the area studied.

Only references pertaining to the area treated in this revision are given. A complete synonymy with all

references up to 1970 can be found in Smetana 1971:246.

Quaest. Ent., 1988, 24 (2)

338

Smetana

Descriptive notes. — First segment of antenna with fine and dense pubescence

(Fig. 409). Mandible moderate in size, each wide basally and then strongly

narrowed into curved and narrow, usually needle-sharp apical portion, before

narrow apical portion with bicuspid or tricuspid tooth on medial margin. Maxillary

palpus with last segment either about as wide as penultimate segment and more or

less gradually and symmetrically acuminate apically, or penultimate segment

swollen, wider than penultimate segment and variably, asymmetrically acuminate

apically. Basal portion of head in front of neck abruptly and strongly depressed.

Dorsal side of head and pronotum impunctate, except for large setiferous punctures.

Prothoracic hypomera strongly inflexed and therefore not visible in lateral view. All

tibiae with spines on lateral face, spines most numerous and distinct on middle tibia.

Dorsal side of last four segments of middle and posterior tarsus smooth and without

setae except for long bristles at distal margin of each segment (Fig. 412). Front

claws distinctly longer than middle and hind claws (Figs. 413, 414). Penultimate

segment of middle and hind tarsus without a pair of very long apical setae (Fig.

413). Empodial setae of middle and hind tarsus long, distinctly protruding between

claws and at least as long as claws, but usually longer (Figs. 412, 413). No male

secondary sexual characters on abdominal sternites or tergites. Male genital segment

with tergite 10 and stemite 9 rounded apically, styli of tergite 9 long and strong,

each with strong apical spine (Fig. 324).

Female genital segment as in Fig. 416, tergite 10 acuminate apically and with

long apical setae.

Acylophorus is a large, predominantly tropical genus, represented by many

species in all zoogeographical regions. Fourteen species are at present known from

the Himalayan region; one species {A. bipunctatus), included here for practical

reasons, occurs in southern India (see under A. bipunctatus).

Classification of species. — Acylophorus requires a thorough revision. The

characters on the aedoeagus, including the internal sac, are essential for the

recognition of the species; unfortunately, these structures are described and

illustrated only for a small portion of the known species. The genus was split into

several subgenera (Bierig 1938; Smetana 1971); however, the status of these

subgenera, which were established for limited faunas, may have to be reevaluated

once the study of the genus is undertaken on a world-wide basis.

For similar reasons I refrained from establishing definite species-groups in this

treatment. Although there seem to be assemblages of species linked together by

some characters, I found it very difficult to meaningfully characterize them as

species-groups. This only can be done based on larger faunas, in this case at least on

the entire Oriental fauna.

Acylophorus puncticeps is quite isolated by the character of the densely punctate

head, a character shared with the only species of the genus Paratolmerus.

Acylophorus bipunctatus is unique by the narrow elongate head with the anterior

frontal punctures situated medially and posteriad of hind margins of eyes. The

Revision of the Tribes Quediini and Atanygnathini

339

remaining species can be divided into two large groups based on the shape of the

last segment of the maxillary palpus. In one group this segment is not enlarged

(about as wide as penultimate segment) and more or less gradually and

symmetrically acuminate apically. Acylophorus heesoni, A. ch.illo, A. daai, A. siyo,

A. furcatus and A. khairo belong here; the aedoeagi of all these species are

characterized by the presence of a strongly sclerotized paramere with numerous peg

setae on the underside; however, a similar type of aedoeagus appears also in A.

puncticeps, which also has the same type of the last segment of the maxillary palpus.

In the second group of species, which includes A. ruficollis, A. halchhi, A.

micTocephalus, A. raato, A. charaa and A. tibialis, the last segment of the maxillary

palpus is swollen, wider than penultimate segment and variably, asymmetrically

acuminate apically. In four species of this group the aedoeagus has a weakly

sclerotized paramere with long and narrow branches without any peg setae on the

underside; however, the aedoeagus of A. tibialis has a strongly sclerotized paramere

with very numerous peg setae on underside.

Acylophorus nepalicus Coiffait (1981:328), described from a single female from

Nepal “Kluihakani, Halambu, Nepal central”), is a species of the genus Erichsonius

Fauvel 1874 (comb.nov.), most likely identical with E. basalis Motschulsky 1858

(type seen).

I do not treat here Acylophorus microcerus Fauvel 1895, described from Burma

(Bhamo) and included by Cameron (1932). I was unable to locate the original

material of the species and found it impossible to interpret the species using only the

description. It should be considerably smaller than any other species of the genus of

the Himalayan region.

Key to species of Acylophorus

1 Pronotum bright red, contrasting in colour with dark head.

Aedoeagus as in Figs. 328-330. Length 6. 8-7.5 mm

7. A. ruficollis Motschulsky, p. 349

F Pronotum not bright red, of about same colour as head.

Aedoeagi different 2

2 (F) Eyes small, in dorsal view considerably shorter than tempora;

anterior frontal punctures located medially and slightly

posteriad of hind margins of eyes. Length 8.0 mm

14. A. bipunctatus Cameron, p. 358

2' Eyes moderately large to large, in dorsal view at least as long as

tempora; anterior frontal punctures located between medial

margins of eyes well in front of hind margins of eyes 3

3 (2') Last segment of maxillary palpus swollen, wider than

penultimate segment and variably, asymmetrically acuminate

apically 4

Quaest. Ent., 1988, 24 (2)

340

Smetana

3' Last segment of maxillary palpus not swollen, about as wide as

penultimate segment and more or less gradually and

symmetrically acuminate apically 9

4 (3) Head wide and rounded, slightly wider than long (ratio 1.10);

anterior frontal puncture situated away from medial margin of

eye, separated from it by distance almost twice as large as

diameter of puncture. Aedoeagus as in Figs. 342, 343. Length

7.0 mm 13. A. tibialis Cameron, p. 357

4' Head narrower, feebly longer than wide or as long as wide,

narrowed anteriorly; anterior frontal puncture situated close to

medial margin of eye, separated from it by distance no larger

than diameter of puncture 5

5 (4') Distinctly bicoloured; head, pronotum and elytra

rufo-testaceous, abdomen dark piceous with apical margins of

tergites and apex paler. Aedoeagus and internal sac as in Figs.

334, 335. Length 5. 0-6.0 mm 9. A. raato spec.nov., p. 351

5' Not bicoloured, forebody not rufo-testaceous and not

contrasting in colour with abdomen. Aedoeagi and internal sacs

different (Figs. 331-333, 336-341) 6

6 (5') Large lateral puncture on pronotum situated close to lateral

pronotal groove, but not touching it. Aedoeagus with median

lobe hook-like apically (lateral view. Fig. 333). Length 6. 1-6.9

mm 8. A. halchhi spec.nov., p. 350

6' Large lateral puncture on pronotum touching lateral pronotal

groove. Aedoeagus with median lobe straight apically, not

hook- like 7

7 (6') Legs uniformly rufo-brunneous. Aedoeagus with apical portion

of median lobe narrow, internal sac inconspicuous (Fig. 337, 341) 8

7' Legs brunneotestaceous with outer faces of middle and hind

femora distinctly darkened. Aedoeagus with apical portion of

median lobe wide, apex broadly arcuate; internal sac

conspicuous (Fig. 338, 339). Length 6. 5-7. 5 mm

11. A. charaa spec.nov., p. 356

8 (7) Median lobe of aedoeagus gradually and evenly narrowed

toward narrowly obtuse apex (Fig. 336); internal sac as in Fig.

337. Length 5. 9-6.8 mm

10. A. microcephalus Cameron, p. 352

8' Median lobe of aedoeagus rather abruptly narrowed toward

truncate, minutely emarginate apex (Fig. 340); internal sac as in

Fig. 341. Length 6.2 mm 12. A.flavipes Motschulsky, p. 356

9 (3') Head densely and finely punctate, except for small impunctate

median area anteriorly. Aedoeagus as in Figs. 344-346. Length

Revision of the Tribes Quediini and Atanygnathini

341

7. 6-8.0 mm 15. A. puncticeps Fauvel, p. 359

9' Head, except for punctate tempora, with only a few large

setiferous punctures (Fig. 408) 10

10 (9') Antenna with segments 2-7 elongate, much longer than wide,

segment 4 subequal in length to segment 5; segment 8

appreciably longer than wide. Paramere of aedoeagus divided

in two branches apically (Fig. 312, 314, 316) 1 1

10' Antenna with segments 2-7 less elongate, distinctly longer than

wide, segment 4 shorter and usually also smaller than segment

5; segment 8 no more than feebly longer than wide. Paramere

of aedoeagus solid (Figs. 319, 323, 327). Size smaller: 5. 2-6.7 mm ..... 13

11 (10) Branches of paramere long and divergent, apical portion of

median lobe wide, broadly arcuate or obtusely truncate apically

(Figs. 313, 315) ...12

IF Branches of paramere short and not appreciably divergent,

apical portion of median lobe nanow, obtusely acuminate

apically (Figs. 316, 317). Length 8.4 mm

...3. A. khairo spec.nov., p. 344

12 (11) Branches of paramere about half as long as length of entire

paramere; apex of median lobe broadly arcuate (Fig. 313).

Length 8. 6-9.4 mm LA. heesoni Cameron, p. 341

12' Branches of paramere slightly more than 1/3 as long as length

of entire paramere; apex of median lobe obtusely truncate (Fig.

315). Length 7.9 mm 2. A. chillo spec.nov., p. 343

13 (10') Paramere of aedoeagus narrowed into very long, narrow, acute

apical portion (Fig. 319). Length 5. 5-6. 7 mm

4. A. siyo spec.nov., p. 344

13' Paramere of aedoeagus moderately narrowed anteriorly, apex

more or less obtuse (Figs. 323, 327) 14

14 (13') Aedoeagus large, apical half of paramere lancet-shaped;

scale-like structures of internal sac coarse and very distinct

(Figs. 325-327). Length 5.2-6. 3 mm

6. A. daai spec.nov., p. 348

14' Aedoeagus smaller, apical half of paramere fusiform; scale-like

structures of internal sac fine and inconspicuous (Figs.

320-323). Length 5. 3-6.4 mm

5. A.furcatus Motschulsky, p. 345

1 . Acylophorus heesoni Cameron

Figs. 312, 313; Map 33

Acylophorus beesoni Cameron 1926:371; 1932:304.

Quaest. Ent., 1988, 24 (2)

342

Smetana

Acylophorus ventralis Coiffait 1983a: 168 (syn.nov.).

Description. — Piceous-black, apical margins of abdominal tergites and 6th visible segment

reddish-brunneous; abdomen distinctly iridescent; mouthparts testaceous; antennae testaceous, apical portion

of segment 1 and segments 2 and 3 piceous, segments 4 and 5 slightly darker than remaining segments; legs

rufo-brunneous with paler tarsi. Head feebly longer than wide (ratio 1.10); eyes moderately large, tempora

about as long as length of eyes seen from above; anterior frontal puncture situated at about level of posterior

third of length of eye and separated from medial margin of eye by distance slightly larger than diameter of

puncture; one puncture posteromediad of posterior frontal puncture; tempora finely punctate and pubescent;

surface of head without any microsculpture. Last segment of maxillary palpus not swollen, about as wide as

penultimate segment and more or less gradually and symmetrically naiTowed anteriorly. Antenna long, first

segment slightly longer than five following segments combined, segments 2-7 elongate, much longer than

wide but gradually becoming shorter, segment 4 subequal in length to segment 5, segment 8 appreciably

longer than wide, segment 9 feebly longer than wide, segment 10 as long as wide, segment 1 1 short, shorter

than preceding segment. Pronotum slightly wider than long (ratio 1.15), broadly rounded basally, strongly

arcuately narrowed anteriorly; large lateral puncture separated from lateral pronotal groove by distance about

equal to diameter of puncture; surface without microsculpture. Scutellum densely punctate. Elytra at base

slightly narrower than pronotum at widest point, slightly dilated posteriorly, at suture shorter (ratio 0.85), at

sides as long as pronotum at midline; punctation dense and moderately coarse, slightly asperate, interspaces

between punctures along transverse axis smaller than diameters of punctures, punctation becoming finer and

denser near elytral base. Pubescence stiff, dense. Wings fully developed. Abdomen with tergite 7 (fifth

visible) with distinct whitish apical seam of palisade setae; punctation of abdominal tergites dense at tergal

bases with punctures more or less elongate, gradually becoming finer, sparser and simple toward apical

margins of tergites; pubescence long and stiff.

Male. Aedoeagus (Figs. 312, 313) small; apical portion of median lobe with apex broadly arcuate;

paramere elongate, divided in two long, divergent branches, each with numerous sensory peg setae along

median margin; internal sac without sclerotized structures.

Length 8. 6-9.4 mm.

Type material. — A. heesoui. The original series in the British Museum (Natural

History), London, contains two specimens under the name A. heesoni. They are

labelled as follows: Spec. No. 1 (female): “Type” (round label with red margin)/

“SYN-TYPE” (round label with blue margin)/ “Kaligad, Dehra Dun Dr. Cameron.

26.VI.21.” / “TYPE Acylophorus beesoni Cam.” / “M. Cameron. Bequest. B.M.

1955-147.”. Spec. No. 2 (male): “SYN-TYPE” (round label with blue margin)/

“Kaligad, Dehra Dun. 10.VII.21.” / “M. Cameron. Bequest. 1955-147.”

The second (male) specimen was dissected and the genital segment and

aedoeagus were mounted on plate with beetle. The specimen is hereby disignated as

the lectotype of A. heesoni\ the label “Lectotype Acylophorus beesoni Cameron

Smetana des. 1984” has been attached to it.

Acylophorus ventralis. Coiffait (1983:168) described the species from a single

specimen from Nepal. The holotype in the collection Coiffait, Museum National

d’Histoire Naturelle, Paris, France, is labelled as follows: “Nepal VIII. 82 Val. de

Kathmandu Bandha T.D.” / “HOLOTYPE” / “Acylophorus ventralis Coiffait det.

19.” The specimen is a male (not a female as stated in the original description); it

was dissected and the aedoeagus and genital segment were mounted on plate with

beetle. The specimen cannot be specifically distinguished from the lectotype of A.

heesoni; the name A. ventralis is a junior synonym of A. beesoni. My determination

label “Acylophorus beesoni Cam. Smetana det. 1984” has been attached to the

specimen.

Revision of the Tribes Quediini and Atanygnathini

343

Geographical distribution. — Acylophorus heesoni is distributed in the western

portion of the Himalayan range, from Uttar Pradesh eastward to central Nepal (Map

33).

Material studied. — 6 specimens.

INDIA. Uttar Pradesh. See Type material of A. heesoni.

NEPAL. Kathmandu Distr. Gokarna Forest 1300 m, 10. IX. 83, Smetana and Ldbl (CNCC, MHNG) 3.

Bionomics. — The specimens from Gokama Forest were taken among soaking

wet fallen leaves on a forest seepage.

Comparisons. — Acylophorus heesoni is characterized by the large size, in

combination with the long antenna (see description), the not swollen last segment of

the maxillary palpus, and the shape of the aedoeagus (Figs. 312, 313). It shares the

first three characters with the two following species; however, each of the two

following species can be easily distinguished by the different aedoeagus (Figs. 314,

316).

2. Acylophorus chillo spec, now

Figs. 314, 315; Map 33

Description. — In all characters very similar to A. heesoni. but different as follows: size smaller and

form less robust, colouration slightly paler; head rufo-brunneous, pronotum rufopiceous, first antennal

segment uniformly testaceous, segments 2-5 not appreciably darkened. Head slightly narrower with

somewhat smaller eyes, tempora slightly longer than length of eyes seen from above (ratio 1.21). Antenna

with middle segments not quite as elongate as in A. heesoni. Pronotum slightly narrower and more narrowed

anteriorly.

Maie. Aedoeagus very similar to that of A. heesoni but somewhat smaller and naiTower; apical portion

of median lobe not dilated anteriorly and with apex obtu.sely truncate; paramere narrower with branches

shorter, slightly more than 1/3 as long as length of entire paramere; internal sac without sclerotized

structures (Figs. 314, 315).

Length 7.9 mm.

Type material. — Holotype (male): “INDIA Assam Manas 200 m 22.X.78

Besuchet-Ldbl”. In the Museum d’Histoire Naturelle de Geneve, Geneve,

Switzerland.

Geographical distribution. — Acylophorus chillo is known only from the type

locality in Assam (Map 33).

Bionomics. — The holotype was taken by sifting leaf litter and floor debris in a

forest.

Comparisons. — Acylophorus chillo is also similar to A. khairo, but differs from

it by the shorter antenna and by the quite different aedoeagus (Figs. 314, 316).

Etymology. — The specific name is the Nepali adjective chillo (smooth). It refers

to the smoothness of the dorsal surface of the head and pronotum.

Quaest. Ent., 1988, 24 (2)

344

Smetana

3. Acylophorus khairo spec, now

Figs. 316, 317: Map 33

Description. — in all characters very similar to A. heesoni, but different as follows: form less robust,

colouration slightly paler, same as in A. chillo. Head slightly narrower with somewhat smaller eyes, tempora

slightly longer than length of eyes seen from above (ratio 1.27). Pronotum slightly narrower and more

narrowed anteriorly.

Male. Aedoeagus (Figs. 316, 317) with median lobe narrow, gradually naiTowed anteriorly, apical

portion narrow with apex obtusely acuminate; paramere relatively wide, with branches short and wide,

narrowly separated medially and not appreciably divergent; sensory peg setae on underside of each branch

numerous, forming elongate group; internal sac without sclerotized structures.

Length 8.4 mm.

Holotype (male): “INDIA Meghalaya Garo Hills 2.XI.78 Songsak 400 m

Besuchet-Lobl”. In the Museum d’Histoire Naturelle de Geneve, Geneve,

Switzerland.

Geographical distribution. — Acylophorus khairo is known only from the type

locality in Garo Hills in Meghalaya (Map 33).

Bionomics. — The holotype was taken in a forest by sifting forest floor debris,

particularly under bamboo growths.

Comparisons. — Acylophorus khairo shares most of the external characters with

A. chillo, but differs from it by the longer antenna (same as described for A. heesoni)

and by the quite different aedoeagus (Figs. 314, 316).

Etymology. — The specific name is the Nepali adjective khairo (brown). It refers

to the colour of this species.

4. Acylophorus siyo spec, now

Figs. 318, 319; Map 34

Description. — Piceous-black, pronotum occasionally slightly paler, humeral angles and apical

margin of elytra, and apical margins of abdominal tergites often feebly paler; abdomen distinctly iridescent;

mouthparts pale testaceous; antennae brunneo-piceous, base of first segment and last segment vaguely paler;

front legs testaceous with paler coxae, middle and hind legs brunneo-piceous to piceous with paler tarsi.

Head feebly wider than long (ratio 1.11), distinctly dilated posteriorly behind eyes and then strongly

narrowed toward neck; eyes moderately large, tempora feebly longer than length of eyes seen from above

(ratio 1.11); anterior frontal puncture situated at about level of posterior third of length of eye and separated

from medial margin of eye by distance equal to diameter of puncture; one puncture posteromediad of

posterior frontal puncture; tempora finely punctate and pubescent; surface of head without microsculpture.

Last segment of maxillary palpus not swollen, as wide as penultimate segment, strongly attenuate apically.

Antenna moderately long, first segment as long as four following segments combined, second segment as

long as segments 3 and 4 combined, segment 4 slightly smaller and shorter than segment 5, segments 6 and 7

slightly longer than wide, segment 8 about as long as wide, segments 9 and 10 slightly transverse, last

segment short, shorter than two preceding segments combined. Pronotum wider than long (ratio 1.20),

moderately narrowed anteriorly; base broadly rounded, but flattened in middle portion; large lateral puncture

separated from lateral pronotal groove by distance about equal to diameter of puncture; surface without

microsculpture. Scutellum densely punctate. Elytra at base slightly narrower than pronotum at widest point,

slightly dilated posteriorly, at suture about as long as pronotum at midline, at sides slightly longer than

pronotum at midline (ratio 1.15); punctation fine and dense, slightly asperate, interspaces between punctures

along transverse axis about equal to diameters of punctures; punctation becoming denser and finer toward

elytral base; pubescence dense. Wings fully developed. Abdomen with tergite 7 (fifth visible) with distinct

Revision of the Tribes Quediini and Atanygnathini

345

whitish apical seam of palisade setae; punctation of abdominal tergites fine and dense at tergal bases with

punctures feebly elongate, gradually becoming finer and sparser toward apical margins of tergites;

pubescence long, dense and stiff.

Male. Aedoeagus (Figs. 318, 319) rather small; middle portion of median lobe slightly dilated, apex

broadly rounded, entirely covered by paramere; paramere elongate, narrowed into very long and narrow,

acute apical portion; sensory peg setae on underside of paramere very numerous, fomiing a large group on

middle portion of paramere; internal sac without sclerotized structures.

Length 5. 3-6. 7 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Distr. 2 km E Mansingma 1900 m 19. IV. 84 Smetana & Ldbl. In the Smetana

Collection, Ottawa, Canada.

Paratypes (8): Nepal; Kosi, 2 km E Mansingma, 1900 m, 19. IV. 84,

Lobl-Smetana (CNCC, MHNG) 7; Khandbari Distr., Induwa Khola Valley, 2050 m,

17.IV.84, Smetana & Lobl (ASCC) 1.

Geographical distribution. — Acylophoriis siyo is known from two localities in

eastern Nepal just east of Arun river (Map 34).

Bionomics. — The specimens were collected at a small creek by sifting wet moss

and debris along its edges and by sifting small flood-debris piles accumulated during

two recent storms. One specimen was also found on a wet talus slope in a

semideciduous broad-leaved forest by sifting wet debris and humus among rocks on

a seepage area.

Comparisons. — Acylophoriis siyo resembles externally A. kailo and A. daai;

however, it differs from both of them, in addition to the distinctive aedoeagus, by

the darker colouration of body, by the darker antennae and middle and hind legs,

and by the more densely punctate elytra and abdominal tergites.

Etymology. — The specific^name is the Nepali noun siyo (needle). It refers to the

shape of the paramere of the aedoeagus of this species.

5. Acylophoriis furcatus Motschulsky

Eigs. 320-323, 408-f 15; Map 34

Acylophorus furcatus Motschulsky 1858a:657.

Description. — in all external characters very similar to A. siyo but different as follows: colouration

paler; rufo-brunneous, elytra rarely feebly darkened, all appendages uniformly testaceous; abdomen strongly

iridescent. Head with eyes longer, tempora slightly shorter than length of eyes seen from above (ratio 0.77)

(Fig. 408). Pronotum more distinctly narrowed anteriorly, large lateral puncture situated somewhat farther

from lateral pronotal groove, separated from it by distance equal to about two diameters of the puncture (Fig.

415). Punctation of both elytra and abdominal tergites sparser.

Male. Aedoeagus (Figs. 32(3-323) fairly small, elongate; apex of median lobe truncate; paramere

elongate and rather narrow, considerably exceeding apex of median lobe, its apical half fusiform, sensory

peg setae on underside of paramere numerous, scattered over apical portion; internal sac with fine and

inconspicuous scale-like structures.

Length 5. 3-6.4 mm.

Type material. — The Motschulsky collection at the Zoological Museum,

Moscow Lomonosov State University, contains one female specimen under the

name A. furcatus. It is labelled as follows: round yellow disc/ “Ind. or.” /

Quaest. Ent., 1988,24 (2)

346

Smetana

Map 33. Distribution records for: Paratolmerus pilosivenths ( ^ ); Acylophoriis beesoni (•);/!. chillo ( ■ ); and/t. khairo ( A ).

Revision of the Tribes Quediini and Atanygnathini

347

Quaest. Ent., 1988, 24 (2)

Map 34. Distribution records for; Acylophorus siyo ( • ); A.furcatus ( A ); /I. charaa ( ■ ); and Anchoceriis hirmanus ( ^ ).

348

Smetana

“Ancylophorus furcatus Motch. Ind. or.”. The specimen was cleaned and remounted

on a larger plate (original plate attached to pin); the genital segment is mounted

separately on plate. The specimen is in relatively good condition: left hind tibia and

tarsus, and middle and hind right tibiae and tarsi are missing. The specimen is

hereby designated as the lectotype of A. furcatus; the label “Lectotype Acylophorus

furcatus Motschulsky, A. Smetana des. 1986” has been attached to it.

Geographical distribution. — Acylophorus furcatus is distributed over eastern

portion of the Himalaya, in West Bengal (Darjeeling area) and Sikkim; also in

Assam and Meghalaya (Garo and Khasi Hills) (Map 34).

Material studied. — 14 specimens.

INDIA. Assam. Manas, 200 m, 22.X.78, Besuchet-Lobl (ASCC, MHNG) 4. Meghalaya. Garo Hills:

Songsak, 400 m, 2. XI. 78, Besuchet-Lobl (MHNG) 1; Garo Hills: Rongrengiri, 400 m, 3.X.78,

Besuchet-Lobl (MHNG) 1; Khasi Hills: Dawki, 500-800 m, 29.X.78, Besuchet-Lobl (MHNG) 2. Sikkim.

Singhik, 4480', 5. III. 32, T. Clay (BMNH) 1. West Bengal. Darjeeling distr.: Sukna, 200 m, 7.X.78,

Besuchet-Lobl (ASCC, MHNG) 2; Sevoke, 200 m, 7.X.78, Besuchet-Lobl (ASCC, MHNG) 2;

Teesta-Rangpo 350 m, 12.X.78, Besuchet-Lobl (MHNG) 1.

Bionomics. — Acylophorus furcatus apparently prefers lower elevations, up to

1360 m. Specimens were collected in an evergreen forest from “moss, leaves,

liverworths” (at Singhik, Sikkim), on a marshy land by sifting dead “Elephant grass”

and by sifting forest floor litter, moss and fallen leaves.

Comparisons. — Acylophorus furcatus can easily be distinguished from A. siyo

by the characters given in the key and in the description. It also closely resembles A.

daai, even in the shape of the aedoeagus. See under the latter for notes about the

distinguishing characters, and Figs. 320, 325.

6. Acylophorus daai spec.nov.

Figs. 324-327; Map 35

Description. — in all external characters very similar to A. siyo but different as follows: colouration

paler, same as in A. furcatus, except apical portion of first antennal segment and antennal segments 2-5

slightly darkened. Head nan'ower, feebly longer than wide (ratio 1.12), only slightly dilated posteriorly

behind eyes, tempora evenly, arcuately narrowed toward neck; eyes moderately large, tempora about as long

as length of eyes seen from above. Pronotum narrower, only slightly wider than long (ratio 1.14), more

distinctly naiTOwed anteriorly. Punctation of both elytra and abdominal tergites sparser, about same as that of

A. furcatus.

Male. Aedoeagus (Figs. 325-327) quite similar to that of A. furcatus but longer; apical half of paramere

lancet-shaped, sensory peg setae on underside of paramere distributed in a similar way to those of A.

furcatus-, scale-like structures of internal sac coarse and very distinct.

Length 5. 2-6. 3 mm.

Type material. — Holotype (male): “NEPAL Khandbari District” / “below

Sheduwa 2100-2500 m 9. IV. 1982 A. & Z. Smetana ”. Allotype (female): “NEPAL

(Prov. Bagmati) Dobate Ridge NE Barahbise, 2800 m, 2.V.81 Lobl & Smetana.” In

the collection of A. Smetana, Ottawa.

Paratypes (11): INDIA: Uttar Pradesh: Garhwal, Mussoorie, 1700 m, 19.X.79, 1.

Lobl (MHNG) 1; Garhwal, 4 km S Bhatwari, 1400 m, 23.X.79, I. Lobl (MHNG) 1.

NEPAL:: Khandbari Distr. Induwa Khola Valley, 2000 m, 14.IV. 84, Lobl-Smetana

Revision of the Tribes Quediini and Atanygnathini

349

(MHNG) 1; Arun Khola Valley at Num main bridge, 1050 m, 21. IV. 84, Smetana &

Lbbl (ASCC) 1; Val. Arun ss/Num, 1050 m, 20. IV. 84, Lobl-Smetana (MHNG) 1.

Lalitpur Distr.: Godawari, 23.V.76, W. Wittmer and C. Baroni-Urbani (NHMB) 1.

Parbat Distr.: Umg. Goropani w. Bokhara, Sept.-Okt, 1971, H. Franz (HGCC) 1.

Sindhupalchok Distr.: Dobate Ridge NE Barahbise, 2800 m, 2.V.81, Lobl &

Smetana (ASCC, MHNG) 3; Barahbise, Ting-Sang-La, H. Franz (HFCC) 1.

Geographical distribution. — Acylophoriis daai is widely distributed; from Uttar

Pradesh through central Nepal to the Arun river valley in eastern Nepal (Map 35).

Bionomics. — Specimens were collected by sifting moist to wet leaf litter in a

depression in a semideciduous broad-leaved forest, in wet detritus at edges of a

small, fast running creek, by sifting thin layer of soaking wet leaves and other debris

in a sandy bank of a creek, by sifting moist leaf litter in a forest ravine and by sifting

of moss and ferns at the edge of a degraded forest.

Comparisons. — Acylophorus daai shares most of the external characters with A.

furcatus; however, it can be distinguished, in addition to the differences on the

aedoeagus (Figs. 320, 325), by the different shape of the head, the slightly shorter

eyes, and the slightly darkened antennal segments 2-5.

Etymology. — The specific name is the Nepali noun daai (older brother). It refers

to the similarity of this species to A. furcatus.

7. Acylophorus ruficoUis Motschulsky

Figs. 328-330; Map 35

Acylophorus ruficoUis Motschulsky 1858a:657; Cameron 1932:303.

Acylophorus ruficoUis Kraatz 1859:65 (syu. now).

Description. — Black, apical margins of first four visible abdominal tergites narrowly, apical portion

of tergite 5 and base of tergite 6 paler, rufo-brunneous to rufo-testaceous; pronotum bright red, abdomen

strongly iridescent; mouthparts and legs testaceous to rufo-testaceous; antennae piceous, gradually becoming

paler toward apex, basal portion of first segment testaceous. Head relatively narrow, about as long as wide,

behind eyes gradually arcuately narrowed toward neck; eyes large, tempora distinctly shorter than length of

eyes seen from above (ratio 0.65); anterior frontal puncture situated at about level of middle of length of eye

and separated from medial margin of eye by distance about equal to diameter of puncture; no puncture

posteromediad of posterior frontal puncture; tempora extensively, finely and densely punctate and

pubescent; surface of head without any microsculpture. Last segment of maxillary palpus slightly swollen,

wider than penultimate segment and asymmetrically acuminate apically. Antenna long, first segment as long

as five following segments combined, segment 2 elongate, longer than segment 3, segment 4 shorter and

slightly smaller than segment 5, segments 6-8 longer than wide, gradually becoming shorter and wider

toward apex, segment 9 as long as wide, segment 10 slightly transverse, last segment short, shorter than two

preceding segments combined. Pronotum only moderately transversely convex and relatively narrow, only

slightly wider than long (ratio 1.10), distinctly nan'owed anteriorly; base broadly rounded but flattened in

middle portion; large lateral puncture separated from lateral pronotal groove by distance somewhat larger

than diameter of puncture; surface of pronotum without microsculpture. Scutellum densely punctate. Elytra

at base slightly narrower than pronotum at widest point, slightly dilated posteriorly, at suture about as long

as, at sides longer than pronotum at midline (ratio 1.17); punctation dense and moderately coarse, asperate,

interspaces between punctures along transverse axis smaller than diameters of punctures; pubescence dense

and stiff. Wings fully developed. Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of

palisade setae; punctation of abdominal tergites finer than that on elytra, dense at tergal bases with punctures

slightly elongate, gradually becoming sparser and more or less finer toward apical margins of tergites;

Quaest. Ent., 1988, 24 (2)

350

Smetana

pubescence long and stiff.

Mate. Aedoeagus (Figs. 328-330) short, stout and wide; median lobe narrowed anteriorly with apex

obtusely arcuate; paramere short, solid, slightly dilated anteriorly, apical margin deeply, obtusely

triangularly emarginate, with three pairs of fine apical setae; sensory peg setae on underside of paramere

very numerous, distributed as in Fig. 330; paramere either covering entire apical portion of median lobe, or

apex of median lobe exposed in emargination of apical margin of paramere; internal sac without large

sclerotized structures, developed as in Fig. 329.

Length 6. 8-7. 5 mm.

Type material. — The Motschulsky collection at the Zoological Museum,

Moscow Lomonosov State University, Moscow, contains two conspecific female

specimens glued on one plate under the name A. nificolUs. They are labelled as

follows: round yellow disc with a part cut off and illegible symbols/ “Ancylophorus

ruficollis Motch. Ind. or.” The specimens were cleaned and remounted on a larger

plate in exactly the same postion as they were glued on the original plate (original

plate attached to pin). The specimens are in relatively good condition: the front

specimen is missing the left elytron, the medioapical portion of the right elytron and

right hind tarsus; the posterior specimen is missing left hind leg. The posterior

specimen is hereby designated as the lectotype of A. ruficollis; the label “Lectotype

(posterior specimen) Acylophorus ruficollis Motschulsky, A. Smetana des. 1986”

has been attached to it.

Geographical distribution. — Acylophorus ruficollis is widely distributed, from

Sri Lanka northwards through Indian subcontinent to Nepal and Assam (Map 35).

Also known from the Malay Peninsula and from Ishigakijima IsL, Okinawa, Japan

(Shibata 1984:139).

Material studied. — 4 specimens.

INDIA. Assam. Gauhati, 200 m, 24.X.78, Besuchet-Lobl (MHNG) 1.

NEPAL. Kathmandu Distr. Kathmandu, H. Franz (HFCC) 1.

Bionomics. — No details are known about the habitat requirements of this

species. It seems to occur at low elevations, up to about 1400 m (Kathmandu).

Recognition. — Acylophorus ruficollis can easily be recognized by the

colouration alone (see the description).

Kraatz (1859:65) actually described A. ruficollis as a new species, using by

chance the same name as Motschulsky one year earlier.

8. Acylophorus halchhi spec. nov.

Figs. 331-333; Map 35

Description. — Piceous-black, apical portions of visible tergites 5 and 6 of abdomen in some

specimens slightly paler; abdomen strongly iridescent; mouthparts testaceous; antennae piceous, indistinctly

paler at apex, first segment testaceous; legs rufo-testaceous with somewhat paler tarsi. Head rather small,

about as long as wide, behind eyes strongly narrowed toward neck; eyes large, tempora about as long as

length of eyes seen from above; anterior frontal puncture situated at about level of middle of length of eye

and very close to medial margin of eye, almost touching it; one puncture posteromediad of posterior frontal

puncture; tempora finely and densely punctate and pubescent; surface of head without any microsculpture.

Last segment of maxillary palpus swollen, wider than penultimate segment and asymmetrically acuminate

apically. Antenna rather short, first segment almost as long as seven following segments combined, segment

Revision of the Tribes Quediini and Atanygnathini

351

2 only moderately elongate, somewhat longer than segment 3, segments 4-7 longer than wide, gradually

becoming shorter and wider, segment 4 slightly shorter and smaller than segment 5, segments 8-10 as long

as wide to slightly wider than long, last segment short, shorter than two preceding segments combined.

Pronotum wider than long (ratio 1.25), broadly rounded basally, strongly narrowed anteriorly; large lateral

puncture situated close to pronotal lateral groove, separated from it by distance slightly less to about equal to

diameter of puncture; surface of pronotum without microsculpture. Scutellum punctate, except smooth

basally. Elytra at base about as wide as pronotum at widest point, slightly dilated posteriorly, at suture about

as long as, at sides slightly longer than pronotum at midline (ratio 1.16); punctation moderately coarse and

dense, asperate, interspaces between punctures along transverse axis slightly larger than diameters of

punctures; pubescence moderately dense, stiff. Wings fully developed. Abdomen with tergite 7 (fifth visible)

with distinct whitish seam of palisade setae; punctation of abdominal tergites finer than that of elytra,

moderately dense at tergal bases, gradually becoming sparse and finer toward apical margins of tergites;

pubescence moderately dense, stiff.

Male. Aedoeagus (Figs. 331-333) nanow and elongate; median lobe suddenly naiTowed into short,

acute and hook-like curved apical portion; paramere weakly sclerotized, divided in two long, narrow and

acute branches, broadly arcuately separated from each other basally; no sensory peg setae on underside of

paramere; internal sac with numerous scale-like structures (Fig. 332).

Fength 6. 1-6.9 mm.

Type material. — Holotype (male) and allotype (female): “INDIA Meghalaya

Khasi Hills 1000 m Mawsynram-Balat 27.X.78 Besuehet-Lbbl. In the Museum

d’Histoire Naturelle de Geneve, Switzerland.

Paratypes (2): same data as holotype (ASCC, MHNG) 2.

Geographical distrihution. — Acylophorus halchhi is at present known only from

Khasi Hills in Meghalaya (Map 35).

Bionomics. — The specimens were collected by sifting leaf litter and other debris

in a forest ravine.

Comparisons. — Acylophorus halchhi can be best distinguished from the similar

species (mainly A. furcatus, A. raato, A. charaa) by the following combination of

characters: last segment of maxillary palpus swollen, large lateral puncture on

pronotum not touching lateral pronotal groove, aedoeagus of characteristic shape

(see the description and Fig. 331).

Etymology. — The specific name is the Nepali noun balchhi (hook). It refers to

the shape of apex of median lobe of aedoeagus of this species.

9. Acylophorus raato spec. nov.

Figs. 334, 335, 416; Map 36

Description. — In all external characters similar to A. halchhi but different as follows: size smaller

and form slightly less stout; colouration paler; bicoloured, head, pronotum and elytra rufo-testaceous,

abdomen dark piceous with apical margins of tergites and apex paler, antennae almost uniformly

brownish-testaceous, occasionally vaguely darkened in middle portion. Head slightly smaller and narrower.

Pronotum somewhat more narrowed anteriorly, large lateral puncture situated close to lateral pronotal

groove, separated from it by distance somewhat smaller than diameter of puncture, in a few specimens

almost touching it.

Male. Aedoeagus (Figs. 334, 335) rather small, elongate; apex of median lobe subtruncate to truncate

apically; paramere short and slightly sclerotized, divided in two long and naiTow, subacute branches, broadly

arcuately separated from each other basally; no sensory peg setae on underside of paramere; internal sac

densely covered with spine-like structures, without larger sclerotized sclerites (Fig. 335).

Quaest. Ent., 1988, 24 (2)

352

Smetana

Length 5. 0-6.0 mm.

Type material. — Holotype (male): “NEPAL, Khandbari District” / “Arun River

at Num 1500-1600 m 10.IV.1982 A. & Z. Smetana”. Allotype (female): “NEPAL

Khandbari Dis. Arun Valley at Num main bridge 1050 m 21. IV. 1984 Smetana &

Lobl”. In the collection A. Smetana, Ottawa, Canada.

Paratypes (18): INDIA: Meghalaya: Khasi Hills, Weloi, 1700 m, 27.X.78,

Besuchet-Lobl (MHNG) 1. West Bengal: Darjeeling Distr., Mahanadi, 1200 m, 6. or

19.X.78, Besuchet-Lobl (ASCC, MHNG) 7. NEPAL: Khandbari Distr.: same data

as holotype (ASCC) 2; same data as allotype (ASCC, CNCC) 5; Val. Arun ss/Num,

1050 m, 21.IV.84, Lobl-Smetana (MHNG) 3.

Geographical distribution. — Acylophoriis raato is distributed in the eastern

portion of the Himalaya, in eastern Nepal and in the Darjeeling area; also in Khasi

Hills in Meghalaya (Map 36).

Bionomics. — Specimens were collected in the Arun River valley by sifting moist

to wet layers of fallen leaves in depressions of the forest floor, and by sifting thin

layer of soaking wet leaves and other debris on a sandy bank of a creek. The

specimens from the Darjeeling district were taken by sifting forest floor litter in a

degraded forest, those from Khasi Hills by sifting forest floor litter.

Recognition. — Acylophorus raato can easily be distinguished by its bicoloured

body with the head, pronotum and elytra rufo-testaceous, and the abdomen dark

piceous (see the description for details), in combination with the swollen last

segment of maxillary palpus, and the shape of the aedoeagus (Pig. 334).

Etymology. — The specific name is the Nepali adjective raato (red). It refers to

the colour of this species.

10. Acylophorus microcephalus Cameron

Pigs. 336, 337; Map 37

Acylophorus microcephalus Cameron 1932:305.

Acylophorus furcatus', Cameron 1932:304 (nec Motschulsky 1858).

Description. — Dark rufopiceous to piceous, pronotum and/or elytra sometimes slightly paler, apical

margins of abdominal tergites and apical portion of sixth visible tergite in some specimens somewhat paler,

rather rufo-brunneous; abdomen strongly iridescent; mouthparts pale testaceous; antennae testaceo-

brunneous, becoming slightly paler toward apex, basal portion of first segment indefinitely paler; legs

uniformly rufo-brunneous. Head relatively small and narrow, feebly longer than wide (ratio 1.12), behind

eyes gradually arcuately narrowed toward neck; eyes moderately large, tempora slightly longer than length

of eyes seen from above (ratio 1.12); anterior frontal puncture situated at about level of middle of length of

eye and separated from medial margin of eye by distance slightly smaller than diameter of puncture; one

puncture posteromediad of posterior frontal puncture; tempora finely and densely punctate and pubescent;

surface of head without any microsculpture. Last segment of maxillary palpus swollen, much wider than

penultimate segment and slightly asymmetrically acuminate apically. Antenna rather short, first segment as

long as six following segments combined, segment 2 only moderately elongate, somewhat longer than

segment 3, segments 4—7 longer than wide, gradually becoming shorter and wider, segment 8 as long as

wide, segments 9 and 10 slightly transverse, last segment short, shorter than two preceding segments

combined. Pronotum broad and voluminous, wider than long (ratio 1.22), moderately, arcuately narrowed

anteriorly; base broadly rounded but flattened in middle portion; large lateral puncture touching pronotal

353

Revision of the Tribes Quediini and Atanygnathini

Qiiaest. Ent., 1988, 24 (2)

Map 35. Distribution records for: Acylophoriis daai (•);/!. nificollis ( A ); and A. balchhi ( ■ ).

354

Smetana

Map 36. Distribution records for: Acylophorus raato ( • ); A. puncticeps ( A ); and Anchocerus monticola ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

355

lateral groove; surface without microsculpture. Scutellum densely punctate. Elytra at base narrower than

pronotum at widest point, slightly widened posteriorly and rather short, at suture shorter (ratio 0.84), at sides

as long as pronotum at midline; punctation dense and moderately coarse, asperate, interspaces between

punctures along transverse axis about equal to diameters of punctures; punctation gradually becoming finer

and denser toward elytral base; pubescence dense and stiff. Wings fully developed. Abdomen with tergite 7

(fifth visible) with distinct whitish apical seam of palisade setae; punctation of abdominal tergites more or

less finer than that on elytra, dense at tergal bases with punctures elongate, becoming much sparser toward

apical margins of tergites and toward apex of abdomen in general; pubescence long and stiff.

Male. Aedoeagus (Figs. 336, 337) very long and narrow; median lobe with apical portion narrow and

with apex narrowly obtuse; paramere weakly sclerotized, divided in two long and narrow, almost hair-like

branches, broadly arcuately separated from each other basally; no sensory peg setae on underside of

paramere; internal sac inconspicuous, without large sclerotized structures (Fig 337).

Fength 5. 9-6. 8 mm.

Type material. — The original series in the British Museum (Natural History),

London, contains six specimens under the name A. mierocephalus. They are

labelled: Spec. No. 1 (male): “Type (round label with red margin)” “SYN-TYPE”

(round label with blue margin) “Arni Gad. Mussoorie.2”/ “Dr. Cameron, 12. VI. 21.”/

“A. mierocephalus Cam. TYPE”/ “M. Cameron. Bequest. B.M. 1955-147.” Spec.

Nos. 2-6: “Arni Gad. Mussoorie.” / “Dr. Cameron, 12. VI. 21 [28.V.21, 16.X.21]” /

“M. Cameron. Bequest. B.M. 1955-147.”

The male specimen No. 1 was dissected and the genital segment and aedoeagus

were mounted on plate with beetle. The specimen is hereby designated as the

lectotype of A. microcephalus\ the label “Lectotype Acylophorus mierocephalus

Cameron, Smetana des. 1984” has been attached to it.

Geographical distribution. — Acylophorus mierocephalus is distributed mainly

in the western portion of the Himalayan range, in Uttar Pradesh; however, one

record is also known from the Darjeeling area in the eastern part of the Himalayan

range (Map 37).

Material studied. — 137 specimens.

INDIA. Uttar Pradesh. Kaligad, Dehra Dun, 10. VII. 21, Cameron (BMNH) 2. Mussoorie Distr.: Dhobi

Ghat, 14.IV.22, Cameron (BMNH) I. Kumaon: Haldwani distr., H.G. Champion (ASCC, CNCC, BMNH)

125; Haldwani distr., banks of Deoba Nadi, 27.V.23, H.G. Champion (BMNH) 1; Garhwal: 20 km S

Chamba, 1150 m, 20.X.79, I. LobI (MNHG) 1. West Bengal. Darjeeling Distr., Rangpo, 400 m, 10.X.78,

Besuchet and Fobl (MHNG) 1.

Bionomics. — Little is known about the habitat requirements of this species. The

specimens from Garhwal were taken in a ravine with a spring by sifting

accumulated twigs, pieces of wood and grasses on gravel, those from Darjeeling

district by sifting debris along the edges of a creek.

Comparisons. — Acylophorus mierocephalus differs from all medium sized and

more or less dark coloured species of the genus, except for A. charaa, by the

position of the large lateral puncture on pronotum which touches the pronotal lateral

groove; the aedoegus is also characteristic (Pig. 336).

Rare specimens of A. raato with the large lateral punctute on pronotum almost

touching lateral pronotal groove can be distinguished from those of A.

mierocephalus by the different colouration (see the description) and by the different

aedoeagus (Pigs. 334, 336).

Quae St. Ent., 1988, 24 (2)

356

Smetana

1 1. Acylophorus charaa spec. nov.

Figs. 338. 339; Map 34

Description. — In all external characters similar to A. microcephalus, but different as follows: size

larger; colouration darker: uniformly black, mouthparts brunneotestaceous, last segment of maxillary palpus

more or less paler; antennae piceous, first segment becoming paler toward base, last segment indistinctly

paler; legs brunneotestaceous with outer faces of middle and hind femora distinctly darkened. Head with

frons between eyes slightly elevated. Antenna stronger. Pronotum narrower, less voluminous and only feebly

wider than long (ratio 1.1), more strongly narrowed anteriorly. Punctation of elytra in general denser and

becoming more distinctly denser toward base. Scutellum more densely punctate. Punctation of abdominal

tergites denser, particularly on bases of tergites.

Male. Aedoeagus (Figs. 338, 339) with median lobe slightly dilated toward broadly arcuate apex;

paramere weakly sclerotised, divided in two long and narrow, acuminate branches, arcuately separated from

each other basally; no sensory peg setae on underside of paramere; internal sac conspicuous, with apical

portion resembling head of an eared owl (Fig. 339).

Length 6. 5-7. 5 mm.

Type material.— Holotype (male) and allotype (female): “NEPAL Lalitpur

Distr. 2 km S Godavari 1700 m 12. IX. 83 Smetana & Lobl”. In the collection A.

Smetana, Ottawa, Canada.

Paratypes (9): same data as holotype (ASCC, BMNH, CNCC, MHNG) 9.

Geographical distrihution. — Acylophorus charaa is at present known only from

the type locality at the foot of the mountain Phulcoki near Kathmandu (Map 34).

Bionomics. — All specimens of the original series were taken by sifting a pile of

fresh fermenting wood shavings in a forest clearing.

Comparisons. — Acylophorus charaa is well characterized, in addition to the

characteristic aedoeagus, by the uniformly black body and the colouration of the

appendages (see the description), the swollen last segment of maxillary palpus and

the position of the large lateral puncture on the pronotum, which is touching the

lateral pronotal groove. On the other hand, it is quite similar to A. flavipes and can

be positively distinguished from it only by the shape of the aedoeagus (Fig. 338,

340).

Etymology. — The specific name is the Nepali noun charaa (bird). It refers to the

shape of the apical portion of the internal sac of the aedoeagus which resembles the

head of an eared owl.

12. Acylophorus flavipes Motschulsky

Figs. 340, 341

Acylophorus flavipes Motschulsky 1858a:657; Cameron 1932:304.

Description. — in all external characters extremely similar to A. charaa and different only by

uniformly rufo-brunneous legs and by characters on aedoeagus. Aedoeagus (Figs. 340, 341) narrow and

elongate; middle portion of median lobe parallel-sided, rather abruptly narrowed toward truncate, minutely

emarginate apex; internal sac inconspicuous. (Fig. 341).

Length 6.2 mm.

Type material. — The Motschulsky collection at the Zoological Museum,

Moscow Lomonosov State University, Moscow, contains one male specimen under

Revision of the Tribes Quediini and Atanygnathini

357

the name A. flavipes. It is labelled as follows: ‘Tnd.or.’V “Ancylophorus flavipes

Motch. Ind.or.”. The specimen was cleaned and remounted on larger plate (original

plate attached to pin); it was dissected, the aedoeagus was mounted on a separate

transparent plate in Canada Balsam, and the genital segment was glued to plate with

beetle. The specimen is in relatively good condition: segments 4-1 1 of left antenna,

segments 1 and 2 of right antenna and left front tibia and tarsus are missing. The

specimen is hereby designated as the lectotype of A. flavipes\ the label “Lectotype

Acylophorus flavipes Motschulsky, A. Smetana des. 1986” has been attached to it.

Geographical distribution. — Acylophorus flavipes was described from “East

India”. Its distributional range is not known at present.

Material studied. — See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons. — Acylophorus flavipes can be positively distinguished from A.

charaa only by the differently shaped aedoeagus (Figs. 338, 340).

Acylophorus flavipes may not occur in the Himalayan area; however, since it

shows relationships with species from the area, it was included in this revision.

13. Acylophorus tibialis Cameron

Figs. 342, 343; Map 37

Acylophorus tibialis Cameron 1932:305.

Description. — Piceous-black, apical margins of abdominal tergites and 6th visible segment vaguely

paler; abdomen distinctly iridescent; mouthparts testaceous; antennae piceous, becoming vaguely paler

toward apex; coxae and tibiae of all legs piceous, all femora and tarsi testaceous. Head wide and rounded,

indistinctly wider than long (ratio 1.08); eyes moderately large, tempora indistinctly shorter than length of

eyes seen from above (ratio 0.88); anterior frontal puncture situated at about middle of length of eye and

away from medial margin of eye, separated from it by distance almost twice as large as diameter of

puncture; one puncture posteromediad of posterior frontal puncture; tempora finely and densely punctate and

pubescent; surface of head without any microsculpture. Last segment of maxillary palpus swollen, wider

than penultimate segment and asymmetrically acuminate apically. Antenna rather short, first segment about

as long as five following segments combined, segment 2 not very elongate, only slightly longer than

segment 3, segments 4-7 longer than wide, gradually becoming shorter and wider, segment 4 somewhat

smaller and shorter than segment 5, segment 8 as long as wide, segments 9 and 10 slightly wider than long,

last segment short, shorter than two preceding segments combined. Pronotum wider than long (ratio 1.20),

slightly arcuately narrowed anteriorly; base broadly rounded but flattened in middle portion; large lateral

puncture separated from lateral pronotal groove by distance about equal to diameter of puncture, several fine

punctures around it; surface without microsculpture. Scutellum densely punctate. Elytra at base about

equally wide as pronotum at widest point, slightly dilated posteriorly and relatively long, at suture about as

long as, at sides longer than pronotum at midline (ratio 1.26); punctation dense and moderately coarse,

slightly asperate, interspaces between punctures along transverse axis about equal to diameters of punctures ;

punctation gradually becoming denser and finer toward elytral base; pubescence dense, stiff. Wings fully

developed. Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of palisade setae;

punctation of abdominal tergites distinctly finer than that on elytra, moderately dense at tergal bases,

gradually becoming much sparser toward apical margins of tergites and toward apex of abdomen in general;

pubescence long and stiff.

Male. Aedoeagus (Figs. 342, 343) rather short and wide; median lobe short, arcuate apically; paramere

short and wide, covering most of median lobe, slightly narrowed and arcuate apically, with fine setae at each

lateral margin below apex and additional shorter setae below them; sensory peg setae on underside of

paramere very numerous, covering most of apical half of paramere; internal sac simple, without large

Quaest. Ent., 1988, 24 (2)

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Smetana

scierotised structures.

Length 7.0 mm.

Type material. — The original series in the British Museum (Natural History),

London, contains one male specimen under the name A. tibialis. It is labelled as

follows: “64520’V “Type”(round label with red margin)/“Birmah Ruby Mes” /

“Acylophorus tibialis Cam. TYPE”. The specimen was dissected, the genital

segment mounted on plate with beetle and the aedoeagus mounted in Canada

Balsam.

The specimen is hereby designated as the lectotype of A. tibialis; the label

“Lectotype Acylophorus tibialis Cameron A. Smetana des. 1984” has been attached

to it.

Geographical distribution. — Acylophorus tibialis is at present known only from

the type locality in Burma (Map 37).

Material studied. — The holotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Recognition. — Acylophorus tibialis can easily be recognized by the wide and

rounded head, in combination with the swollen last segment of maxillary palpus,

and the relatively long elytra.

14. Acylophorus bipunctatus Cameron

Acylophorus hipunctatus C'dmeron 1920; 219.

Anchocerus bipunctatus-, Cameron 1932: 308.

Description. — Brunneo-piceous, apical margins of abdominal tergites and 6th visible segment

rufo-brunneous; abdomen iridescent; mouthparts and antennae rufo-testaceous, antennal segments 3-6

slightly darkened, last four segments paler, pale yellowish; legs rufo-brunneous with paler tarsi. Head

narrow, longer than wide (ratio 1.18); eyes small, tempora about twice as long as length of eyes seen from

above; anterior frontal punctures situated medially and slightly posteriad of hind margins of eyes; one

puncture posteromediad of postertior frontal puncture; temporal puncture situated distinctly closer to

posterior margin of head than to posterior margin of eye, tempora with several fine punctures; surface of

head without any microsculpture. Last segment of maxillary palpus as wide as penultimate segment and

gradually narrowed anteriorly. Antenna moderately long, first segment slightly longer than three following

segments combined, second segment slightly shorter than segments 3 and 4 combined, segment 3 distinctly

longer than wide, segments 4—7 longer than wide, gradually becoming shorter, segment 8 slightly longer

than wide, segments 9 and 10 as long as wide, last segment short, much shorter than two preceding segments

combined. Pronotum feebly wider than long (ratio 1.09), broadly rounded basally and strongly narrowed

anteriorly; large lateral puncture situated away from lateral groove of pronotum, separated from it by

distance equal to about three diameters of puncture; surface without microsculpture. Scutellum densely

punctate, except smooth basally. Elytra at base about as wide as pronotum at widest point, slightly dilated

posteriorly, at suture shorter (ratio 0.85), at sides as long as pronotum at midline; punctation fine and

moderately dense, slightly asperate, interspaces between punctures along transverse axis somewhat larger

than diameters of punctures; punctation becoming slightly denser near elytral base; pubescence moderately

dense. Wings fully developed. Abdomen with tergite 7 (fifth visible) with distinct whitish apical seam of

palisade setae; punctation of abdominal tergites dense at tergal bases with punctures elongate, gradually

becoming sparser and simple toward apical margins of tergites; pubescence long and stiff.

Male. Unknown.

Length 8.0 mm (abdomen extended).

Revision of the Tribes Quediini and Atanygnathini

359

Type material. — Cameron (1920:219) described the species from a single

specimen from Nilgiri Hills. The female holotype, deposited in the British Museum

(Natural History), London, is labelled as follows: “Type H.T. (round label with red

margin)/ “368”/ “H.L. Andrews Nilgiri Hills”/ “A. bipunctatus Cam.”/ “M.

Cameron. Bequest. 1955-147.”

Geographical distribution. — Acylophorus bipunctatus is known only from the

type locality in Nilgiri Hills in southern India. The species does not occur in the

Himalayan region; it was included only for practical reasons (to treat all the species

of Acylophorus included in Cameron (1932)).

Material studied. — The holotype.

Bionomics. — Nothing is known about the habitat requirements of this species.

Recognition. — Acylophorus bipunctatus can easily be recognized by the narrow

head with the anterior frontal punctures situated medially and slightly posteriad of

hind margins of eyes. It cannot be confused with any species of Acylophorus known

to occur in the Himalaya.

The species was originally described by Cameron (1920:219) as an Acylophorus^

however, Cameron (1932:308) assigned it later to the genus Anchocerus, without

giving any reasons for this new combination. I believe that the chaetotaxy of the

pronotum (large lateral puncture not doubled as in all species of Anchocerus) and

the shape of the labrum warrant the reassignment of this species to Acylophorus.

Cameron (1920, 1932) gives the length of this species as “9 mm.”. However, the

holotype, even with the abdomen extended, is only 8.0 mm long.

15. Acylophorus puncticeps Fauvel

Figs. 344-346; Map 36

Acylophorus puncticeps Fauvel 1895:275; Cameron 1932:303.

Description. — Piceous-black with black head, apical margins of abdominal tergites and 6th visible

segment paler, rufo-brunneous; abdomen iridescent; mouthparts testaceous, last two segments of antennae

rufo-testaceous; legs rufo-brunneous with paler tarsi. Head wide, about as long as wide; eyes large, tempora

slightly shorter than length of eyes seen from above; surface of head densely and finely punctate, except for

small impunctate median area anteriorly. Last segment of maxillary palpus about as wide as penultimate

segment and symmetrically narrowed anteriorly. Pronotum slightly wider than long, broadly rounded basally

and strongly arcuately naiTowed anteriorly; surface without microsculpture. Scutellum finely and densely

punctate. Elytra at base slightly narrower than pronotum at widest point and slightly widened posteriorly;

punctation coarse and rather dense, interspaces between punctures along transverse axis smaller than

diameters of punctures, punctation becoming distinctly finer and denser near elytral base. Abdomen with

tergite 7 (fifth visible) with whitish apical seam of palisade setae, punctation of abdominal tergites dense and

rather fine, gradually becoming sparser medioapically, apical portions almost smooth, impunctate.

Male. Aedoeagus (Figs. 344-346) moderately large; median lobe slightly dilated toward broadly arcuate

apical margin; paramere with two long, slightly divergent branches, each with Hattened medio-apical area

densely covered with sensory peg setae, and with one apical seta; internal sac as in Fig. 345.

Length 7. 6-8.0 mm.

Type material. — Fauvel (1895:275) described the species from specimen(s?)

from Burma: “Carin Ghecu, alt. 1300-1400 meters, Feb.-Mar. (L.Fea).” I was not

able to study the original material which is not deposited in the collection Fauvel,

Quaest. Ent., 1988, 24 (2)

360

Smetana

Map 37. Distribution records for: Acylophorus microcephalus { • ); /t. tibialis ( A ); and Anchocenis pimctatissimus ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

361

Bruxelles, Belgique. My concept of the species is based on two specimens from the

collection Fauvel I studied. They are labelled as follows: Spec. No. 1 (female):

“Irawaddi Birmanie” / “Coll, et det. A. Fauvel Acylophorus puncticeps Fauv. R.I.

Sc. N.B. 17.479.” Spec. No. 2 (male): “Birmanie Flelfer” / “puncticeps Fvl.” /

“R.I.Sc.N.B. 17.479 Acylophorus Coll, et det. A. Fauvel.”

The male specimen was dissected, the aedoeagus was mounted in Canada

Balsam, and the genital segment glued to plate with beetle.

Geographical distribution. — Acylophorus puncticeps is known from two

localities (one of them not located) in Burma (Map 36).

Material studied. — 2 specimens.

BURMA. See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparison. — Acylophorus puncticeps is unique in having the head densely and

finely punctate. In this respect, it resembles Paratolmerus pilosiventris; however,

the latter differs abundantly in other characters.

Although my concept of A. puncticeps is not based on the study of the original

material, it is very likely that the described specimens really represent the species

Fauvel described.

8. Gtnws Anchocerus Fauvel

A}Kiiocerus¥^u\t\ 1905: 141; Cameron 1932:306

Type species. — Anchocerus hirmanus Fauvel 1905, designated by Fauvel

(1905:141), by original designation and monotypy.

In addition to the characters given by Fauvel (l.c.) and Cameron (l.c.), I note the

following features.

Descriptive notes. — First segment of antenna lacking fine and dense

pubescence. Mandibles large, sickle-shaped and with asymmetrically developed

teeth on medial margin; right mandible with large and rounded, plate-like tooth in

basal third and with another, much smaller and narrower, subacute tooth just in front

of it; left mandible only with rounded plate-like basal tooth. Gula very short, gular

sutures fused. Infraorbital ridge complete but becoming very fine and inconspicuous

anteriorly. Basal portion of head in front of neck abruptly and strongly depressed, as

in Acylophorus. Dorsal surface of head and pronotum with double punctation (fine

punctures intermixed with coarser ones) in some species. Prothoracic hypomera

strongly inflexed, not visible in lateral view. Prosternum with central, fairly sharp

spine-like protuberance. Front and middle tibia with numerous and strong spines on

lateral face, hind tibia without spines on lateral face. Dorsal side of all tarsal

segments pubescent. Front claws only indistinctly longer and larger than middle and

hind claws. Penultimate segment of middle and hind tarsus without a pair of very

long apical setae. Empodial setae of middle and hind tarsus short, hardly visible

between claws and considerably shorter than claws. No male secondary sexual

Quaest. Ent., 1988, 24 (2)

362

Smetana

characters on abdominal stemites or tergites. Male genital segment with tergite 10

and sternite 9 rounded apically, styli of tergite 9 long, strong and wide, each with

three strong apical spiniform setae (Fig. 351).

A?ichocerus is an Oriental genus with eight species known at present. Four

species are known to occur in the Himalayan region.

Comparisons. — Anchocevus can be distinguished from Acylophorus, in addition

to the characters in the key, by the in general larger and more parallel-sided form,

and by the distinct, double punctation of the dorsal side of the head and pronotum in

some species.

Key to species of Anchocerus

1 Head wide, wider than long, tempora each with narrow, finely

punctate and pubescent area posteriorly. Surface of head and

pronotum very finely but distinctly punctate. Paramere of

aedoeagus with two thin, hair-like branches (Figs. 349, 352,

male of one species not known) 2

1’ Head narrow, about as long as wide, tempora without narrow,

finely punctate and pubescent area posteriorly. Surface of head

and pronotum almost impunctate. Paramere of aedoeagus solid,

without hair-like branches (Fig. 347). Length 7.2 mm

\. A. monticola Cameron, p. 363

2 (1) Surface of head and pronotum with very fine, simple and fairly

sparse punctation. Aedoeagus with median lobe moderately

long and moderately narrowed anteriorly (Figs. 349, 350).

Length 9.5-1 1.2mm 2. A. hirmanus Fauvel, p. 364

2' Surface of head and pronotum with extremely fine, dense

punctures intermixed with sparser and coarser punctures.

Aedoeagus (known only for one species) with median lobe long

and strongly narrrowed anteriorly (Figs. 352, 353) 3

3 (2') No setiferous puncture near posterior margin of eye. Elytra

longer, at sides about as long as pronotum at midline;

punctation of elytra dense, intervals between punctures about as

large as diameters of punctures. Average size larger. Length

8. 5- 9. 5 mm 3. A. punctatissimus Smetana, p. 365

3' Fine setiferous puncture near posterior margin of eye. Elytra

shorter, at sides shorter than pronotum at midline (ratio 0.80);

punctation of elytra very dense, intervals between punctures

along transverse axis smaller than diameters of punctures.

Average size smaller. Aedoeagus as in Eigs. 352, 353. Length

7. 5- 8. 5 mm 4. A. nepalicus spec.nov., p. 366

Revision of the Tribes Quediini and Atanygnathini

363

1 . Anchocerus monficola Cameron

Figs. 347, 348; Map 36

Anchocerus monticola Cameron 1926:371; 1932:307

Description. — Piceous-black, pronotum, elytra and apical margins of abdominal tergites

indistinctly paler, apex of abdomen appreciably paler; both maxillary and labial palpi testaceous, antennae

and legs riifo-testaceoiis, tarsi slightly paler. Head relatively narrow, about as wide as long, almost

parallel-sided behind eyes and then strongly nanowed toward neck; eyes small and only slightly convex,

tempora considerably longer than length of eyes seen from above (ratio 2.1), with narrow, finely punctate

and pubescent area posteriorly in front of neck; anterior frontal punctures situated clo,se to each other on

median portion of frons, distance between them distinctly smaller than distance separating each puncture

from median margin of eye; posterior frontal puncture situated close to posterior margin of head, one

puncture posteromediad of it; minute setiferous puncture at posterior margin of eye; temporal puncture

situated much closer to posterior margin of head than to posterior margin of eye; surface of head without

microsculpture and with only very few scattered, extremely fine punctures. Antenna with segment 1 slightly

dilated anteriorly, segment 2 about 1/3 longer than segment 3, segment 4 slightly shorter than segment 3,

segments 5-8 longer than wide, gradually becoming shorter and wider, segments 9 and 10 about as long as

wide, last segment very short, transverse. Pronotum slightly wider than long (ratio 1.19), broadly rounded

basally and very inconspicuously narrowed anteriorly; dorsal rows each with only one puncture situated just

before middle of pronotum; sublateral rows missing; large lateral puncture doubled; surface of pronotum

without microsculpture and with only very few scattered, extremely fine punctures. Scutellum punctate on

apical portion, without microsculpture. Elytra at base only slightly narrower than pronotum and rather short,

at suture about as long as and at sides feebly longer (ratio 1.09) than pronotum at midline; punctation rather

dense, intervals between punctures about as large as diameters of punctures; pubescence fine, dark; surface

between punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible)

bearing fine whitish apical seam of palisade setae; punctation dense, coarse and rather asperate on basal

portions of tergites, gradually becoming sparser, finer and hardly asperate toward apical margin of each

tergite, tergite 6 only sparsely and finely punctate; pubescence dense, dark and long; surface without

microsculpture.

Male. Aedoeagus (Figs. 347, 348) with median lobe elongate and nanow, gradually narrowed and

subacuate apically; paramere in form of solid, slightly transverse plate with anterior margin subemarginate in

middle; internal sac, when evaginated, with two strongly sclerotised apical spines (Fig. 348).

Fength 7.2 mm.

Type material. — The original series in the British Museum (Natural History),

Lonidon, contains four specimens under the name A. monticola. They are labelled as

follows: Spec. No. 1 (male): “Type” (round label with red margin)/ “SYN-TYPE”

(round label with blue margin)/ “Kaligad, Dehra Dun” / “Dr. Cameron. 19. VI. 21.” /

“Anchocerus monticola Cam.” / “M. Cameron. Bequest B.M. 1955-147.” Spec. Nos.

2-A (Nos. 2 and 4 females. No. 3 male): “Kaligad, Dehra Dun” / “Dr. Cameron.

26.VI.21.” / “M. Cameron. Bequest B.M. 1955-147.” / “SYN-TYPE” (round label

with blue margin).

The first specimen was dissected and the aedoeagus mounted in Canada Balsam.

The specimen is hereby designated as the lectotype of A. monticola; the label

“Lectotype Anchocerus monticola Cameron Smetana des. 1984” has been attached

to it. The remaining three specimens are not conspecific with the lectotype and

belong to A. hirmanus.

Geographical distribution. — Anchocerus monticola is at present known only

from the type locality in Uttar Pradesh (Map 36).

Quaest. Ent., 1988, 24 (2)

364

Smetana

Material studied. — The lectotype .

Bionomics. — Nothing is known about the habitat requirements of this species.

Recognition. — Anchocerus monticoia can easily be recognized by the relatively

narrow head without nan'ow, finely punctate and pubescent area on each side in

front of neck, and by paramere of the aedoeagus lacking any branches.

2. Anchocerus hirmanus Fauvel

Figs. 349, 350; Map 34

Anchocerus hirmanus F'duve\ 1905:141; Cameron 1932:306; Smetana 1977a:249.

Description. — Dark reddish-piceous to piceous, pronotum and elytra often paler, apical margins of

abdominal tergites and apex of abdomen slightly paler; both maxillary and labial palpi testaceous, antennae

darker testaceous, legs testaceo-rufous with paler tarsi. Head rounded, wider than long (ratio 1.24), slightly

widened behind eyes and then strongly narrowed toward neck, widest at about posterior fourth; eyes small

and flat, tempora considerably longer than length of eyes seen from above (ratio 2.33), each with narrow,

finely punctate and pubescent area posteriorly in front of neck; anterior frontal punctures situated close to

each other on median portion of frons, distance between them distinctly smaller than distance separating

each puncture from median margin of eye; posterior frontal puncture situated close to posterior margin of

head, one puncture posteromediad of it; temporal puncture situated in densely punctate and pubescent area in

front of neck; setiferous puncture at posterior margin of eye missing; surface of head without

microsculpture, with very fine and not dense punctation gradually becoming denser laterally. Antenna with

segment 2 almost twice as long as segment 3, segment 4 slightly shorter than segment 3, segments 5-8

longer than wide, gradually becoming shorter and wider, segments 9 and 10 about as long as wide, last

segment short, much shorter than two preceding segments combined. Pronotum wider than long (ratio 1.18),

broadly rounded basally and slightly narrowed anteriorly; dorsal rows each with only one puncture situtated

just before middle of pronotum; sublateral rows each reduced to Just one fine puncture distant from anterior

margin and situated rather laterally (often missing, even bilaterally); larger lateral puncture doubled; surface

without microsculpture, punctation similar to that on head but even finer and sparser. Scutellum punctate on

apical portion, without microsculpture. Elytra at base narrower than pronotum and rather short, at suture

shorter (ratio 0.81), at sides about as long as pronotum at midline; punctation rather dense and moderately

coarse, intervals between punctures about as large as diameters of punctures; pubescence fine, dark; surface

between punctures without microsculpture. Wings fully developed. Abdomen with tergite 7 (fifth visible)

bearing fine whitish apical seam of palisade setae; punctation dense, rather coarse and more or less asperate

on basal portions of tergites, gradually becoming distinctly sparser, finer and hardly asperate toward apical

margin of each tergite, sixth visible tergite only sparsely and finely punctate; pubescence dense, dark and

long; surface slightly iridescent, with extremely fine and dense microsculpture of incomplete transverse

waves.

Male. Aedoeagus (Figs. 349, 350) with median lobe slightly asymmetrical, moderately long and obtuse

apically; paramere with two hairdike branches; internal sac as in Fig. 350.

Fength 9.5-11.2 mm.

Type material.—- The original series of A. hirmanus was studied by Smetana

(1977:249) who also designated the female lectotype. See Smetana (l.c.) for details.

Geographical distribution. — Anchocerus hirmanus is at present known from

Burma (not mapped) and from northern Uttar Pradesh in India (Map 34).

Material studied. — 8 specimens.

BURMA. Tenasserim. (Fectotype - see Smetana 1977:249).

INDIA. Uttar Pradesh. Dehra Dun, Kaligad, 26.VI.21, Cameron (ASCC, BMNH) 3; Dehra Dun, Nun

Nadi, 17.VII.21, Cameron (BMNH) 1.

Bionomics. — Nothing is known about the habitat requirements of this species.

Revision of the Tribes Quediini and Atanygnathini

365

Comparisons and taxonomic notes. — Anchocerus hirmanus differs from the two

similar Himalayan species, A. pimctatissimus and A. nepalicus, by the simple, fine

punctation of the head and pronotum lacking intermixed coarser punctures, and by

the shape of the aedoeagus (male of A. pimctatissimus not known).

The large gap between the localities in Burma and in India may be due to the

inadequate collecting in the interlying areas. It is also possible that the two

populations actually represent two very similar separate species. Males of the

Burmese population are needed for the clarification of this question.

The three specimens from Kaligad (see above) belong to the original series of A.

monticola; they differ significantly from the lectotype of A. monticola (see “Type

material” under A. monticola).

Cameron (1932:307) described the 6th visible sternite of the male as “very finely

crenulate”. I am not able to detect any crenulation of the apical margin of this

sternite in the single male available (from Kaligad); the sternite is identically

developed in both sexes.

3. Anchocerus pimctatissimus Smetana

Map 37

Anchocerus punctatissimHS Smetana 1977a:248.

Description. — Very similar to A. hirmanus but different as follows; average size smaller; slightly

darker, piceous to piceous-black, elytral humeri, apical margins of abdominal tergites and apex of abdomen

somewhat paler, rather piceo-rufous to rufo-brunneous. Surface of head densely covered with extremely fine

and superficial punctation with intermixed coarser punctures becoming gradually more frequent toward

lateral portions of head. Surface of pronotum with dual punctation similar to that on head; however, both

fine and coarser punctures somewhat finer, and coarser punctures more difficult to see, especially laterally.

Male. Unknown.

Length 8. 6-9. 2 mm.

Type material. — The species was described from one female specimen

(holotype) from Bhutan, labelled as follows: “Changra 18 km S. Tongsa, 1900 m,

22/6”/ “Nat. -Hist. Museum Basel-Bhutan Expedition 1972” / “HOLOTYPE Ancho-

cerus punctatissimus A. Smetana 1975”. The hololtype is deposited in the

Naturhistorisches Museum in Basel, Switzerland.

Geographical distribution. — Anchocerus punctatissimus is at present known

from Bhutan and from eastern and central Nepal (Map 37).

Material studied. — 5 specimens.

BHUTAN. See Type material.

NEPAL. Kathmandu Distr. Gokama Forest nr. Kathmandu, 1300 m, 10. IX. 83, Smetana and Lobl

(ASCC) 1. Khandbari Distr. Arun Valley at Num main bridge, 1050 m, 21. IV. 84, Smetana and Lobl

(ASCC) 3.

Bionomics. — All specimens from Nepal were taken at relatively low elevations

(below 1500 m). The specimen from Gokama Eorest was taken by sifting very wet

fallen leaves and other debris on a forest seepage area. The specimens from the

Arun valley were taken together with Q. nilo, by sifting a thin layer of soaking wet

leaves and other debris on a sandy bank of a creek.

Quaest. Ent., 1988, 24 (2)

366

Smetana

Comparisons. — Anchocerus punctatissimiis differs from A. hinnanus mainly by

the different punctation of the head and pronotum. Males are needed to differentiate

it further from A. hirmaniis.

For a comparison with A. nepalicus, see the comparisons under the latter species.

4. Anchocerus nepaliciis spec, now

Figs. 351-353; Map 38

Description. — Very similar to A. hinnanus but different as follows: size smaller; antenna shorter,

middle segments less elongate, segments 9 and 10 distinctly transverse, last segment very short, transverse.

Surface of head and pronotum with dual punctation similar to that of A. punctatissinius, but punctation

denser and identically developed on both head and pronotum. Head with fine setiferous puncture near

posterior margin of eye, at least unilaterally. Pronotum distinctly narrowed anteriorly, almost subtruncate

basally and therefore of almost rhomboid shape. Elytra distinctly shorter, at suture much shorter (ratio 0.66),

at sides somewhat shorter (ratio 0.83) than pronotum at midline; punctation and pubescence very dense,

intervals between punctures along transverse axis smaller than diameters of punctures. Brachypterous, wings

in form of short, apically slightly folded, nonfunctional stumps. Punctation and pubescence of abdominal

tergites in general somewhat denser; whitish apical seam of palisade setae on tergite 7 (fifth visible) very

delicate.

Male. Aedoeagus (Figs. 352, 353) with median lobe long, fairly symmetrical, strongly narrowed and

with apex slightly curved; paramere with two hair-like branches; internal sac as in Fig. 353.

Fength 7. 5-8. 5 mm.

Type material. — Holotype (male): “NEPAL Khandbari District” / “For. above

Ahale 2300 m 26.IIL82 A. & Z. Smetana”. Allotype (female): “NEPAL Khandbari

District” / “For. NE Kuwapani 2450 m 13. IV. 82 A. & Z. Smetana”. In the Smetana

collection, Ottawa, Canada.

Paratypes (3); NEPAL: Khandbari Distr., Forest S Mansingma, 2250 m,

12.IV.84, Smetana & Lobl (ASCC) 1; Kosi: Foret S. Mansingma, 2300 m, 13.IV.84,

Lobl-Smetana (MHNG) 1. INDIA: West Bengal: Darjeeling Distr., Ghoom-Lopchu,

2000 m, 14.X.78, Besuchet & Lobl (MHNG) 1.

Geographical distribution. — Anchocerus nepalicus is at present known from a

few localities in eastern Nepal and in the Darjeeling district in West Bengal (Map

38).

Bionomics. — The specimens from Nepal were collected by sifting wet moss on

rocks or fallen trees in shady and moist forest habitats.

Comparison. — Anchocerus nepalicus can easily be distinguished from both A.

hinnanus and A. punctatissimus by the presence (at least unilaterally) of a fine

setiferous puncture near posterior margin of the eye, by the different shape of the

pronotum (see above), by the shorter and more densely punctate and pubescent

elytra, and by the reduced, nonfunctional wings. It also differs from A.

punctatissimus by the punctation of the pronotum which is identical with that on the

head (finer than that on the head with coarser punctures more difficult to see in A.

punctatissimus).

Etymology. — The specific name is an adjective derived from the name of the

country in which most specimens of the original series were collected.

Revision of the Tribes Quediini and Atanygnathini

367

Quaest. Ent., 1988, 24 (2)

Map 38. Distribution records for: Anchocerus nepalicus ( • ); Atanygnathus pictus ( A ); /t. sasuraa ( ■ ); and A. bindii ( ^ ).

368

Smetana

2. Tribe Atanygnathini

Tanygnathinini Reitter 1909:105; Szujecki 1980:152.

Tanygnathini Kuhnt 1913:247; Casey 1915:424; Portevin 1929:333.

Atanygnathini Lohse 1964:220.

The tribe is characterized by the following: tarsal formula 5,4,4; head under eyes

with mandibular ridge (for definition see Smetana 1977b: 180) which begins,

because of strong elongation of head, on posterior genal region (not at level of

insertion of mandible) and gradually disappears behind eyes; mouthparts very

elongate, stipes of maxilla and palpiger partially fused, palpiger with long setae on

lateral margin and along fusion line between stipes and palpiger (see Fig. 2 in

Smetana 1984:280); sternite 9 of male genital segment obsolete (Fig. 357);

aedoeagus without paramere; female genital segment as in Figs. 417^20, tergite 10

subacute and with long and strong apical setae. General habitus resembling

tachyporine genus Tachyporus Gravenhorsted 1802.

The aedoeagus rests in the abdomen with the ventral side (where the proximal

opening is) facing ventrally.

The tribe contains only the predominantly tropical genus Atanygnathus. The

genus, and the tribe along with it, was alternately put either in the subfamily

Tachyporinae or Staphylininae (in the latter case either as a separate tribe, or, as

most modern authors did, within the tribe Quediini). The main reason for the

inclusion in the Quediini was apparently the presence of a ridge on the head below

the eye, which was believed to be the infraorbital ridge of the Quediini. However,

this ridge in fact is not homologous with the infraorbital ridge of the Quediini; I

suggested the name “mandibular ridge” for it (see Smetana 1977b: 180 and 1984:279

for details). Since Atanygnathus does not show any other characters of the tribe

Quediini, I recently suggested (Smetana 1984:281) that a separate tribe within the

subfamily Staphylininae should be kept for it, at least until its phylogenetic

relationships are clearly demonstrated, using also the characters of the larva which is

unfortunately still undescribed.

1. Genus Atanygnathus Jakobson*

Tanygnathus Erichson 1839:417 (nec Wagler 1832).

Atanygnathus iakohson 1909:520 {nom.nov.y, Cameron 1921:355,379; 1932:308.

Tanygnathinus Reitter 1909:105 (nom. now).

Type species. — Tanygnathus terminalis Erichson 1839, designated by Jakobson

(1909:520).

I am not presenting here a conventional formal description of this genus; it can

be found in Cameron (1932:308) and in some recent papers, e.g., Smetana

1971:269.

g

Only references pertaining to the area treated in this revision are given. A complete account of references

up to 1970 can be found in Smetana 1971:269.

Revision of the Tribes Quediini and Atanygnathini

369

Some additional characters, not mentioned in previous descriptions can be found

above under the tribe Atanygnathini.

The genus Atanygnathus is distributed worldwide, particularly in tropical and

wami temperate areas. Eight species are at present known from the Himalayan

region.

Taxonomic notes. — Atanygnathus is a fair size genus in need of thorough

modern revision. The Himalayan species of Atanygnathus are very similar to each

other and are, with the exception of A. pictus and A. sasuraa, rather dark coloured.

The shape of the aedoeagus and particularly the arrangement of the sclerites of the

internal sac are quite important for the positive determination of the species. The

aedoeagi of most species are in general of similar shape, with apical portion of

median lobe wide and with apex obtuse, arcuate or rounded (Figs. 361, 364); A.

hindu is unique in this respect with its median lobe of aedoeagus strongly narrowed

anteriorly into a needle-sharp apex (Fig. 370). Males of all Himalayan species lack

the brush of thick black bristles at lower median margin of front femora present in

all Nearctic species, but have the first three segments of front tarsus provided with

long and dense pale hairs ventromedially.

Cameron (1932:309) recorded the Palaearctic species A. terminalis Erichson

1839 with the varieties v. ruficoUis and v. fuscus as occurring in India. But both

varieties proved to be separate species and the specimens coloured in a similar way

as the typical A. terminalis from Europe also proved to be specifically different,

belonging to A. sasuraa. To the best of my knowledge A. terminalis does not occur

in the Himalayan region and most likely not on the Indian subcontinent.

Key to species of Atanygnathus

1 Pronotum relatively narrow, with ratio length/width above 0.8.

Aedoeagus and sclerites of internal sac in situ as in Figs. 354,

355. Length 3.9-4. 1 mm 1. A. pictus Motschulsky, p. 370

T Pronotum wider, with ratio of length/width no more than 0.78.

Aedoeagi and sclerites of internal sacs in situ different 2

2 (!') Median lobe of aedoeagus with apical portion more or less

wide and with apex obtuse, broadly arcuate or rounded (Figs.

358, 361,364, 367,374, 377) 3

2' Median lobe of aedoeagus strongly narrowed into an extremely

narrow, needle-sharp apical portion (Fig 370). Length 4. 0-5.0

mm 6. A. hindu spec. now, p. 376

3 (2) Internal sac of aedoeagus in situ without a strongly sclerotised,

tweezer-like structure proximally (Fig. 359). Length 3. 3^. 7

mm 2. A. sasuraa spec. now, p. 372

3' Internal sac of aedoeagus in situ with a strongly sclerotised,

tweezer-like structure proximally (Figs. 362, 365, 368, 375, 378) 4

Quaest. Ent., 1988, 24 (2)

370

Smetana

4 (3') Internal sac of aedoeagus in situ with distal sclerites simple,

composed of only one pair of narrow, slightly lyre-shaped

structures; apical portion of median lobe short (Figs. 362, 365) 5

4' Internal sac of aedoeagus in situ with distal sclerites complex;

apical portion of median lobe long (Figs. 368, 375, 378) 6

5 (4) Proximal tweezer-like structure of internal sac of aedoeagus

short, with branches widely separated basally; apical portion of

median lobe fairly narrow, with apex obtusely arcuate (Figs.

364, 365). Length 3.9 mm .... 4. A. piceiis (Motschulsky), p. 374

5' Proximal tweezer-like structure of internal sac of aedoeagus

long, with branches narrowly separated basally; apical portion

of median lobe wide, with apex broadly rounded (Figs. 361,

362). Length 4.2^.4 mm 3. A. hrevicollis Fauvel, p. 373

6 (4') Internal sac of aedoeagus in situ with a pair of acute, triangular

sclerites in distal group of sclerites (Fig. 378). Length 3. 7-3. 9

mm 8. A. purha spec. now, p. 379

6' Internal sac of aedoeagus in situ without acute, triangular

sclerites in distal group of sclerites (Figs. 368, 375) 7

7 (6’) Median lobe of aedoeagus slightly dilated anteriorly, apex

broadly rounded (Fig. 374); sclerites of internal sac of

aedoeagus as in Fig. 375. Length 3.1— 4.0 mm

7. A. chiso spec. now, p. 'ill

7 Median lobe of aedoeagus parallel-sided with apex obtusely

arcuate (Fig. 367); sclerites of internal sac of aedoeagus as in

Fig. 368. Length 4.3-5. 1 mm 5. A. paani spec. now, p. 375

\. Atanygnathus pictus (Motschulsky)

Figs. 354, 355; Map 38

Tanygiiathus pictus Motschulsky 1858b:213.

Tanygnathus ruficollis Kraatz 1859:64 {syn. now)

Atanygnathus terminalis var. ruficollis', Cameron 1932:310.

Description. — Head brownish-piceous; pronotum pale testaceous, vaguely darkened anteriorly;

elytra brownish-piceous, posterolateral angles extensively and posterior margin narrowly pale testaceous;

abdomen testaceous, basal portions of first three tergites brownish-piceous; mouthparts and legs uniformly

yellowish, antennae brownish, gradually becoming pale yellowish both toward base and apex. Head slightly

wider than long (ratio 1.16), surface with excessively fine, rudimentary microsculpture appreciable on

anterior portion and gradually obliterating toward posterior margin. Antenna with segments 2 and 3 equal in

length, segment 2 somewhat stronger than segment 3, segments 4-8 distinctly longer than wide and

gradually becoming shorter and wider, segments 9 and 10 slightly longer than wide, segment 1 1 as long as 2

preceding segments combined. Pronotum relatively narrow, with ratio length/width 0.84, widely rounded

basally and strongly narrowed anteriorly, evenly transversely convex; surface with excessively fine,

rudimentary microsculpture. Scutellum finely and densely punctate. Elytra moderately long, slightly

widened posteriorly and at base slightly narrower than pronotum at widest point, at suture about as long as,

at sides slightly longer than pronotum at midline (ratio 1.14); punctation very fine and dense, interspaces

between punctures along transverse axis about equal to diameters of punctures; pubescence brownish. Wings

Revision of the Tribes Quediini and Atanygnathini

371

fully developed. Abdomen with tergite 7 (fifth visible) bearing distinct whitish apical seam of palisade setae;

punctation extremely fine and dense on basal portions of tergites, gradually becoming sparser toward apical

margin of each tergite and in general toward apex of abdomen; pubescence brownish.

Male. First three segments of front tarsus with long and dense pale hairs ventromedially; apical margin

of stemite 8 with moderately wide and shallow, slightly angulate emargination. Aedoeagus (Figs. 354, 355)

rather small and of characteristic shape, strongly narrowed from about middle; median lobe broadly arcuate

apically; sclerites of internal sac in situ as in Fig. 355, without large and strongly sclerotised structures

distally.

Female. Apical margin of sternite 8 broadly rounded.

Length 3.9^. 1 mm.

Type material. — Atanygnafhus pictiis. The Motschulsky collection at the

Zoological Museum, Lomonosov State University, Moscow, contains one female

specimen under the name T. pictus. It is labelled as follows: round yellow disc with

a part cut off /“type”/ “Tanygnathus pictus Motch. ruficollis Kraatz Ind. or”. The

specimen is in fair condition; both antennae except for basal segments and both

middle legs are missing. The left front leg and the genital segment are glued

separately on plate with beetle. The specimen is hereby designated as the lectotype

of A. pietus; the label “Lectotype Tanygnathus pictus Motschulsky, A Smetana des.

1986” has been attached to it.

Atanygnathus ruficollis. The collection Kraatz in DEI, Eberswalde, German

Deomocratic Republic, contains 26 specimens under the name T. ruficollis, but only

four belong to the original series. They are labelled as follows: Spec. No. 1 (9):

“138”/ “Ceylon”/ “Syntypus”/ “Tanygnat. ruficollis Kr.”/“Coll. Kraatz” / “Col. DEI

Eberswalde”. The remaining three specimens (2d'd', 19) bear the following identical

labels: “Ceylon”/ “Syntypus”/ “Coll. Kraatz”/ “Col. DEI Eberswalde”. Spec. No. 4

is missing the label “Coll. Kraatz”. The two males (Spec. Nos. 2 and 4) were

dissected, both tergite and sternite 8 and the genital segment were glued to plates

with beetles, and the aedoeagi were mounted in Canada Balsam. Specimen No. 4 is

hereby designated as the lectotype of A. ruficollis', the label “Lectotype Tanygnathus

ruficollis Kraatz A. Smetana des. 1985” has been attached to it.

The remaining 22 specimens come from Kambodja, Lagos, Sumatra (Padang)

and “Aegypt. Sudan” (Bahr el Ghazal). They were not closely studied; they most

likely belong to different species.

One of the specimens from Assam in MHNG was compared with the lectotype

and accordingly labelled.

Geographical distribution. — Atanygnathus pictus is at present known from Sri

Lanka (no other data) and from northeastern India (Map 38).

Material studied. — 8 specimens.

INDIA. Assam. Manas, 200 m, 23.X.78, Besuchet-Lobl (MHNG) 1; Gauhati, 200 m, 24.X.78,

Besuchet-LobI (ASCC, MHNG) 2. Meghalaya. Garo Hills, Rongrengiri, 400 m, 3. XI. 78, Besuchet-LbbI

(MHNG) 1.

SRI LANKA. See Type material of A. ruficollis.

Bionomics. — The specimens from Assam were taken by sifting fallen leaves and

other forest floor debris; and by sifting deep layer of leaves accumulated at base of a

steep slope; those from Garo Hills by sifting forest floor debris, particularly under

Quaest. Ent., 1988, 24 (2)

372

Smetana

bamboo growths.

Recognition and variation. — Atanygnathus pictus can best be recognized, in

addition to the distinctive shape of the aedoeagus, by the relatively narrow pronotum

and by the extremely fine and dense punctation on the basal portions of the first

three abdominal tergites.

All specimens seen were either slightly teneral or faded (original series). The

actual colouration of the species may therefore be darker than indicated in the

description.

2. Atanygnathus sasuvaa spec. nov.

Figs. 356-359, 417^20; Map 38

Description. — in all characters very similar to A. pictus, but different as follows: colour very

variable: head piceous to piceous black; pronotum rufo-testaceous, occasionally with indistinct cloud of

slightly darker colour in about apical third, or uniformly dark brownish to piceous-brown; elytra piceous

with apical protions to various extent paler, rufotestacous, sometimes piceous with only apical margin

narrowly paler; abdomen piceous to piceous-black, apical portions of tergites and apex of abdomen

rufo-testaceous, paler apical portions of tergites gradually becoming wider toward apex of abdomen;

mouthparts yellowish, antennae yellowish with middle segments (usually 2-8) variably darkened; legs pale

testaceous. Pronotum somewhat wider, with ratio length/width 0.78. Punctation of elytra somewhat coarser

and less dense; punctation of bases of abdominal tergites slightly less dense.

Male. First three segments of front tarsus as in /\. ruficollis', apical margin of sternite 8 with wide and

moderately deep, obtusely angulate emargination (Fig. 356). Aedoeagus (Figs. 358, 359) larger, median lobe

evenly and gradually narrowed anteriorly, apex arcuate; sclerites of internal sac in situ as in Fig. 359,

without a strongly sclerotised tweezer-like structure proximally.

Female. Apical margin of sternite 8 subtruncate.

Length 3. 3-4. 7 mm.

Type material. — Holotype (male) and allotype(female): “NEPAL (Prov.

Bagmati) Tarang Marang 1000 m, 27. IV. 81 Lobl & Smetana”. Both holotype and

allotype in the collection A. Smetana, Ottawa, Canada.

Paratypes (73): INDIA: Assam: Manas, 200 m, 23.X.78, Besuchet-Lobl (ASCC,

MHNG) 10. Meghalaya: Garo Hills, Songsak, 400 m, 2.XI.78, Besuchet-Lobl

(MHNG) 1. Uttar Pradesh: Kumaon: Haldwani Distr., H.G. H.G. Champion (ASCC,

CNCC, BMNH) 22; Haldwani Distr., Nandhaur R., H.G. Champion (BMNH) 1;

Haldwani Distr., banks of Deoba Nadi, 27.V.23, H.G. Champion (BMNH) 2;

Ranikhet, H.G. Champion (BMNH) 2; Sarju Valley, 4000', H.G. Champion

(BMNH) 1; W. Almora, H.G. Champion (BMNH) 1; W. Almora Dvn., IV.1917,

G.H. Champion (BMNH) 1; C. Almor Dvn., I. 1920, H.G. Champion (BMNH) 1;

Garjia, 10 km off Ramagar, 450 m, 15.X.79, I. Lobl (MHNG) 2. Mussoorie Distr.:

Kolhu Khet Gad, LXI.21, Cameron (BMNH) 1; Keyarkull, 12.XI.21, Cameron

(BMNH) 1; Dehra Dun Distr.: Lachiwala, 21.11.32, H.G. Champion (ASCC,

BMNH) 7; Dehra, Narkaunda, 2000', 23.X.17, H.G. Champion (BMNH) 8; Dehra

Dun, Raipur, 30.X.27, H.G. Champion (BMNH) 1; Dehra Dun, Phandowala, Suewa

R., 1.IV.28, H.G. Champion (BMNH) 2; Dehra Dun, R. Song, 2.IV.22, Cameron

(BMNH) 2. Siwaliks, Kheri Ran, 30.X.21 Cameron (BMNH) 1; Siwaliks, Mohan

Revision of the Tribes Quediini and Atanygnathini

373

Ran, 17.X.21, Cameron (BMNH) 1. West Bengal: Darjeeling Distr., Sukna, 200 m,

7.x. 78, Besuchet-Ldbl (MHNG) 1. NEPAL: “Wald unterhalb Fulung”, Franz

(HFCC) 1. Khandbari Distr.: Pangma, 1700 m, 4. IV. 84, Smetana & Lobl (ASCC) 3.

Geographical distribution. — Atanygnathus sasiiraa is widely distributed

throughout the Himalayan region, from Uttar Pradesh through Nepal to Assam and

Meghalaya (Map 38).

Bionomics. — Atanygnathus sasuraa seems to occur mainly at lower elevations

(below 2000 m). Specimens were collected by sifting wet fallen leaves and various

debris around seepages, among grass on wet meadows along irrigation canals and by

sifting forest floor debris, particularly under bamboo growths.

Recognition and variations. — Atanygnathus sasuraa varies considerably in

colour (see the description). Specimens with pale pronotum, particularly those with

indefinite darker anterior spot, resemble specimens of A. pictus particularly closely,

but they can be distinguished by the characters given in the description, especially

by the differences on the aedoeagus. Darker coloured specimens resemble those of

the following six species and in most cases the aedoeagus is needed for a positive

identification.

Etymology. — The specific name is the Nepali noun sasuraa (father in law). It

refers to the similarity of this species with A. pictus.

3. Atanygnathus hrevicollis (Fauvel)

Figs. 360-362; Map 39

Tanygnathus hrevicollis Fauvel 1895:276.

Description. — Head piceous; pronotum brownish to brownish-piceous; elytra piceous with suture

and apical margin slightly and narrowly paler; abdomen piceous, slightly iridescent, ‘apical portions of

tergites and apex of abdomen paler, rufo-brunneous; mouthparts pale testaceous; antennae with first segment

testaceous, following segments brunneous, gradually becoming paler toward antennal apex, so that last 2-3

segments are pale testaceous; legs rufo-testaceous. Head slightly wider than long (ratio 1.20), surface with

fine and dense microsculpture of transverse meshes. Antenna with segments 2 and 3 equal in length,

segment 2 somewhat stronger than segment 3, following segments much longer than wide; gradually

becoming shorter and wider, segment 10 distinctly longer than width at apex, last segment slightly shorter

than two preceding segments combined. Pronotum wide, with ratio length/width 0.71, widely rounded

basally and moderately narrowed anteriorly, evenly transversely convex; surface with extremely fine

microsculpture of mostly incomplete transverse and/or oblique meshes. Scutellum finely punctate. Elytra

moderately long, slightly widened posteriorly, at suture feebly (ratio 1.06), at sides slightly longer than

pronotum at midline (ratio 1.22); punctation very fine and very dense, interspaces between punctures along

transverse axis about equal to diameters of punctures; pubescence decumbent, brownish. Wings fully

developed. Abdomen with tergite 7 (fifth visible) bearing distinct whitish apical seam of palisade setae;

punctation very fine and dense on basal portions of tergites, gradually becoming distinctly sparser toward

apical margin of each tergite and in general toward apex of abdomen; pubescence brownish.

Male. First three segments of front tarsus with long and dense pale hairs ventromedially; apical margin

of stemite 8 with moderately wide and rather shallow, obtusely angulate emargination (Fig. 360). Aedoeagus

(Figs. 361, 362) relatively wide; apical portion of median lobe short and wide, with apex broadly rounded

(Fig. 361); internal sac in situ with distal sclerites simple, composed only of one pair of narrow, slightly

lyre-shaped structures, and with proximal tweezer-like structure long and with branches narrowly separated

basally (Fig. 362).

Quaest. Ent., 1988, 24 (2)

374

Smetana

Female. Apical margin of sternite 8 broadly arcuate.

Length 4. 2-4.4 mm.

Type material. — The Fauvel collection in the Institute Royal des Sciences

naturelles de Belgique, Bruxelles, contains three specimens under the name A.

hrevieollis. They are labelled as follows: Spec. No. 1 (female): “Carin Ghecu

1300-1400 m. L. Tea 11-111.88”/ “brevicollis Fvl.”/ “R.I.Sc.N.B. 17.479

Atanygnathus Coll, et det. A. Fauvel”/ “Syntype” Spec. No. 2 (female) and Spec.

No. 3 (male): “Carin Asciui Ghecu 1400-1500 m. L. Fea.” “HI-IV.88”/ “Coll, et det.

A. Fauvel Atanygnathus brevicollis Fauv. R.I. Sc. N.B. 17.479”/ “Syntype”.

The male specimen was dissected and the sternite 8 and the aedoeagus were

mounted in Canada Balsam. The specimen is hereby designated as the lectotype of

A. hrevieollis', the label “Lectotype Atanygnathus brevicollis Fauv. Smetana des.

1983” has been attached to it.

Geographieal distribution. — Atanygnathus hrevieollis is at present known only

from the type locality in Burma (Map 39).

Material studied. — 3 specimens.

BURMA. See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species.

Comparisons and taxonomic notes. — Atanygnathus hrevieollis is in all external

characters extremely similar to the two following species; the shape of the

aedoeagus and the sclerites of the internal sac are necessary for positive

identification.

Cameron (1932:311) records A. hrevieollis from Assam (Naga Hills, in

Nagaland) and Uttar Pradesh (Mussoorie Distr. and from the Siwaliks). However,

his records apply in fact to different species, apparently to A. paani and/or to dark

specimens of A. sasuraa.

4. Atanygnathus piceus (Motschulsky)

Figs. 363-365

Tany^nathus piceus Motschulsky 1858b:213

Tanygnathus fuscus Kraatz 1859:65 [syn. nov.)

Atanygnathus fuscus', Cameron 1932:310

Description. — in all characters extremely similar to A. hrevieollis and different mainly in some

characters on aedoeagus. Size smaller; pronotum slightly narrower, with ratio length/width 0.73.

Male. First three segments of front tarsus dilated in similar way to those of A. hrevieollis-, sternite 8 with

apical emargination much deeper (Fig. 363). Aedoeagus very similar to that of A. hrevieollis but smaller;

apical portion of median lobe fairly narrow, with apex obtusely arcuate (Fig. 364); internal sac in situ with

distal sclerites similar to those of A. hrevieollis, but proximal tweezerdike structure short and with branches

widely separated basally (Fig. 365).

Female. Apical margin of sternite 8 broadly arcuate.

Length 3.9 mm.

Type material. — Atanygnathus piceus. The Motschulsky collection at the

Zoological Museum, Lomonosov State University, Moscow, contains three

conspecific specimens {2dd and 1 ?) glued on one plate under the name T. piceus.

Revision of the Tribes Quediini and Atanygnathini

375

They are labelled as follows: round yellow disc with a part cut off/“type”/

“Tanygnathus piceus Motch. fuscus Kraatz Ind. or.” The specimens were cleaned

and remounted on a larger plate in exactly the same position as they were glued on

the original plate (original plate attached to pin). Both males were dissected; the

aedoeagus and tergite and sternite 8 of the first male were mounted in Canada

Balsam on a separate transparent plate, which was attached to pin; the aedoeagus

and the genital segment of the second male were glued to plate with beetles. The

specimens are in poor condition. The first male (with the aedoeagus and segment 8

mounted in Canada Balsam) is hereby designated as the lectotype of A. piceus; the

label “Lectotype (first male) Tanygnathus piceus Motschulsky A. Smetana des.

1986” has been attached to it; the specimen is missing right antenna, five apical

segments of left antenna and the tarsus of left middle leg (right front leg and the

genital segment are glued separately on plate with beetles).

Atanygnathus fuscus. The collection Kraatz in DEI, Eberswalde, German

Democratic Republic, contains one male specimen under the name T. fuscus. It is

labelled as follows: “43”/ “Ost-Indien”/ “Holotype”/ “Tanygnath. fuscus Kr.”/

“Coll. DEI Eberswalde” / “A. fuscus Kr.”. The specimen was dissected; the genital

segment and the tergite and sternite 8 were glued to plate with beetle, and the

aedoeagus was mounted in Canada Balsam. The specimen is hereby designated as

the lectotype of A. fuscus; the label “Lectotype Tanygnathus fuscus Kraatz A.

Smetana des. 1986” has been attached to it.

The specimen is in poor shape: right maxillary palpus, four outer segments of the

left and all but 2 basal segments of right antenna and all legs except for left hind leg

are missing. The specimen cannot be distinguished from the lectotype of A. piceus;

the name is a junior synonym of A. piceus.

Geographical distribution. — Atanygnathus piceus was described from “East

India”. Its distributional range is not known at present.

Material studied. — See Type material.

Bionomics. — Nothing is known about the habitat requirements of this species.

Notes. — Atanygnathus piceus may not occur in the Himalayan area; however,

since it shows relationships to several species from the area, it was included in this

revision.

5. Atanygnathus paani spec. nov.

Eigs. 366-368; Map 39

Atanygnathus hrevicolUs\ Cameron 1932:31 1 (ex parte)

Description. — in all characters extremely similar to A. hrevicollis and different only in characters

on aedoeagus and slightly different emargination of male sternite 8.

Male. First three segments of front tarsus as in A. hrevicollis, apical margin of stemite 8 with wider and

deeper emargination (Fig. 366). Aedoeagus (Figs. 367, 368) longer than that of A. hrevicollis: median lobe

parallel-sided and elongate, with apex obtusely arcuate; intemal sac in situ with distal sclerites more

complex, developed as in Fig. 368, proximal tweezer-like structure with branches widely separated basally.

Quaest. Ent., 1988, 24 (2)

376

Smetana

Length 4.3-5. 1 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Lamjung

Distr. Besisahar 900 m 17. IX. 83 Smetana & Lobl”. In the collection A. Smetana,

Ottawa, Canada.

Paratypes (23): INDIA: Meghalaya: Khasi Hills, Mawsynram-Balat, 1000 m,

27.X.78, Besuchet-Lobl (MHNG, ASCC) 4. Assam: Manas, 200 m., 23.X.78,

Besuchet-Lobl (MHNG) 2. Uttar Pradesh: Haldwani Distr., Kumaon, H.G.

Champion (ASCC, CNCC, BMNH) 14; Mussoorie Distr., Arni Ghad, 28.V.21, Dr.

Cameron (BMNH, FMNH) 2; Garhwal, 20 km S Chamba, 1 150 m, 20.IX.79, 1. Lobl

(MHNG) 1.

Geographical distribution. — Atanygnathus paani is widely distributed

throughout the Himalaya, from Uttar Pradesh through Nepal to Assam and

Meghalaya (Map 39).

Bionomics. — Atanygnathus paani seems to occur at lower elevations, up to

about 1200 m. Specimens were taken in Nepal (holotype and allotype) in soaking

wet vegetation and moss around a small waterfall; other specimens were taken by

sifting leaf litter and other debris in a forest ravine, and in a ravine with a spring by

sifting accumulated twigs, pieces of wood and grasses on gravel.

Recognition. — Atanygnathus paani can be positively identified only by the

characters of the aedoeagus and internal sac.

Etymology. — The specific name is the Nepali noun paani (water). It refers to the

habitat of the specimens taken in Nepal (see under “Bionomics”).

6. Atanygnathus hindu spec. nov.

Figs. 369-372; Map 38

Description. — in all characters extremely similar to A. hrevicollis and different in drastically

different aedoeagus and different emargination of male stemite 8.

Male. First three segments of front tarsus as in /t. hrevicollis; apical margin of stemite 8 with wide and

very deep obtusely triangular emargination (Fig. 369). Aedoeagus (Figs. 370-372) of unique shape, elongate

and very namow; median lobe strongly narrowed into extremely narrow, needle-sharp apical portion; internal

sac in situ (Fig. 371) composed of two very narrow and elongate, slightly sclerotised structures bearing

dense setae and one distal, heavily sclerotised semicircular structure; however, in most specimens internal

sac is evaginated (Fig. 372).

Length 4. 0-5.0 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Dis. Arun Valley at Num main bridge 1050 m, 20. IV. 1984 Smetana & Lobl. In the

collection A. Smetana, Ottawa, Canada.

Paratypes (34): INDIA: Meghalaya. Khasi Hills, between Mewsynram and Balat,

1000 m, 27.x. 78, Besuchet-Lobl (MHNG) 2; Khasi Hills, Cherapunjee, 1200 m,

26.X-78, Besuchet-Lobl (MHNG) 3; Khasi Hills, Dawki, 500-800 m, 29.X.78,

Besuchet-Lobl (BMNH, MHNG) 3. NEPAL: Khandbari Distr.: same data as in

holotype, date 20, 21 or 22.IV.84 (ASCC, CNCC) 6; Kosi, Val. Arun ss/Num, 1050

or 1100 m, 20. or21.IV.84, Lobl-Smetana (BMNH, CNCC, MHNG) 19.

Revision of the Tribes Quediini and Atanygnathini

377

Geographical distribution. — Atanygnathiis hindii is at present known from

eastern Nepal and from Khasi Hills in Meghalaya (Map 38).

Bionomics. — Specimens of A. Hindu were taken in Nepal together with Quedius

assamcnsis, Q. nilo and A. chiso by sifting thin layer of soaking wet leaves and other

debris on a sandy bank of a creek. Specimens from Meghalaya were taken by sifting

leaf litter and other debris in a forest ravine and by sifting of fallen leaves in a fairly

dry forest.

Comparisons. — Atanygnathus Hindu can be positively identified only by the

unique shape of the aedoeagus (Figs. 370-372); the wide and deep emargination of

the apex of male sternite 8 is also diagnostic (Fig. 369). Atanygnathus Hindu occurs

in Khasi Hills together with A. paani in the same habitat and in eastern Nepal with

A. chiso (see under the respective species). Specimens from these areas have to be

very carefully screened; the females not associated with the males cannot be

positively determined.

Etymology. — The specific name is the Nepali noun bindu (point). It refers to the

shape of the aedoeagus.

7. Atanygnathus chiso spec. nov.

Figs. 373-375; Map 39

Description. — Piceous-black with apical margins of abdominal tergites and apex of abdomen

somewhat paler; or head piceous, pronotum piceous-brown and elytra piceous with feebly paler suture and

apical margins; mouthparts pale testaceous; antennae dark brownish, gradually becoming paler both toward

base and apex, latter pale testaceous; legs rufo-testaceous with paler tarsi. Head slightly wider than long

(ratio 1.17); surface with fine and extremely dense microsculpture of mostly slightly transverse meshes.

Antenna with segments 2 and 3 equal in length, segment 2 somewhat stronger than segment 3, following

segments much longer than wide, gradually becoming shorter and wider, segment 10 somewhat longer than

at apex wide, last segment slightly shorter than two preceding segments combined. Pronotum wide, with

ratio width; length 0.73, widely rounded basally and moderately narrowed anteriorly, evenly transversely

convex; surface with microsculpture similar to that on head. Scutellum densely and very finely punctate.

Elytra rather short, slightly widened posteriorly, at suture shorter (ratio 0.82), at sides as long as pronotum at

midline; punctation fine and very den.se, interspaces between punctures along transverse axis mostly

somewhat smaller than diameters of punctures; pubescence decumbent, dark brownish. Wings fully

developed. Abdomen with tergite 7 (fifth visible) bearing whitish apical seam of palisade setae; punctation

very dense and fine on basal portions of tergites, gradually becoming distinctly sparser toward apical margin

of each tergite and in general toward apex of abdomen; pubescence dark brownish.

Male. First three segments of front tarsus with long and den.se pale hairs ventromedially; apical margin

of sternite 8 with wide but not deep, obtusely triangular emargination (Fig. 373). Aedoeagus (Figs. 373-375)

moderately large; median lobe slightly dilated toward broadly arcuate apex; internal sac in situ with strongly

sclerotized, tweezer-like structure proximally and with distal sclerites complex, developed as in Fig. 375.

Female. Apical margin of 8th sternite obtusely arcuate.

Fength 3. 1^.0 mm.

Type material. — Holotype (male) and allotype (female): “NEPAL Khandbari

Dis. Arun Valley at Num main bridge 1050 m 21. IV. 84 Smetana & Lobl.” In the

collection A. Smetana, Ottawa, Canada.

Paratypes (11): INDIA: Meghalaya, Khasi Hills, Mewsynram-Balat, 1000 m,

27.x. 78, Besuchet-Lobl (MHNG) 3. NEPAL: Khandbari Distr.: same data as

Qiiaest. Ent., 1988, 24 (2)

378

Smetana

Map 39. Distribution records for: Atanygnathus hrevicollis { ^ Y, A. paani ( • ); chiso ( A ); and A. purha ( ■ ).

Revision of the Tribes Quediini and Atanygnathini

379

holotype, date 20. or 21. IV. 84, Lobl & Smetana (MHNG) 5; Arun River at Num,

1500-1600 m, 10.IV.82, A. & Z. Smetana (ASCC) 1; Khandbari, 1700 m, 23. III. 82,

A. & Z. Smetana (ASCC) 1; Kuwapani, 2100 m, 28. III. 82, A. & Z. Smetana

(ASCC) 1.

Geographical distribution. — Atanygnathus chiso is at present known from

eastern Nepal and from Meghalaya (Map 39).

Bionomics. — Specimens of A. chiso were taken in the Arun river valley,

together with Quedius assamensis, Q. nilo and Atanygnathus hindu, by sifting thin

layer of soaking wet leaves and other debris on a sandy bank of a creek; one

specimen also above the river by sifting wet layers of leaf litter in the forest. Other

specimens were taken by sifting various debris in a seepage among bushes in a

disturbed habitat; or under stones in wet habitats near irrigated fields.

Comparisons. — Atanygnathus chiso resembles in all external characters the four

preceding species and differs from them only insignificantly by the generally

somewhat smaller size and shorter elytra. The characters on the aedoeagus and the

internal sac are essential for the positive identifcation of this species. Atanygnathus

chiso occurs in the same habitats with A. hindu in Nepal and with A. hindu and A.

paani in Khasi Hills (see under the respective species). Specimens from these areas

have to be very carefully screened; females not associated with males are impossible

to distinguish from those of A. hindu and A. paani.

Etymology. — The specific name is the Nepali adjective chiso (cold). It refers to

the habitat of the specimens taken in Nepal at Arun river (see under “Bionomics”).

8. Atanygnathus purha spec, now

Figs 376-378; Map 39

Description. — in all characters extremely similar to A. chiso but different as follows: average size

smaller; colouration similar, but elytra with suture and apical margins always narrowly paler, abdomen

rufo-brunneous, bases of some tergites vaguely and indistinctly darkened in some specimens.

Male. First three segments of front tarsus with longer and denser pale ventromedian hairs less developed

and in general inconspicuous; apical margin of stemite 8 emarginate in similar way to that of A. chiso (Fig.

376). Aedoeagus (Figs. 377, 378) similar to that of A. chiso, but median lobe naiTower, slightly narrowed

towards obtuse apex; internal sac similar to that of A. chiso, but distal group of sclerites different,

particularly by presence of a pair of acute, triangular sclerites (Fig. 378).

Female-. Unknown.

Length 3. 7-3. 9 mm.

Type material. — Holotype (male); “E. NEPAL; KOSI Val. Arun ss/Num 1050

m, 21. IV. 84 Lobl-Smetana”. In the Museum d’Histoire Naturelle, Geneve,

Switzerland.

Paratypes (3): same data as holotype (ASCC, MHNG) 3.

Geographical distribution. — Atanygnathus purha is known only from the Arun

river valley in eastern Nepal (Map 39).

Bionomics. — Specimens of Atanygnathus purha were taken by sifting wet debris

along a creek.

Quaest. Ent., 1988, 24 (2)

380

Smetana

Comparisons. — Atanygnathus purha can be distinguished from A. chiso by the

smaller size and the colouration, as described in the description; however, the

characters on the aedoeagus are diagnostic.

Etymology. — The specific name is the Nepali noun purba (east).

CONCLUSIONS

The Himalaya

To aid in understanding the biogeography of the Himalayan Quediini and

Atanygnathini, the most important features of the Himalaya are briefly discussed.

More detailed information on the Himalaya can be found in several publications,

Burrard and Hayden (1907), Mani (1968) and Franz (1979).

The Himalaya is an enormous mass of Tertiary fold-mountains stretching

northwesterly from near the northeastern corner of India in the east to Pakistan in

the west (approximately between the longitudes 72° and 97°E, and the latitudes 27°

and 36°N. The Tertiary orogeny of the Himalaya, its high mean elevation and

enormous massiveness, the essentially east-west alignment of the mountain ranges,

and the Pleistocene glaciations are the main factors that have influenced the present

distributional patterns of the plants and animals of the region.

The main ecological/geographical divisions of the Himalaya are: 1) East or

Assam Himalaya (about 720 km long, from the southern bend of the Brahmaputra

river west to the Tista river in West Bengal), 2) Central or Nepal Himalaya (about

800 km long, from the Tista river west to the Kali river between Nepal and Uttar

Pradesh), 3) West or Kumaon Himalaya (about 320 km long, from the Kali river

west to the Sutlej river in Himachal Pradesh), and 4) Northwest or Punjab-Kashmir

Himalaya (about 560 km long, from the Sutlej river northwest to the southwestern

bend of the Indus river in Pakistan). While the first three divisions are to some

extent artificial, the separation of the Northwest Himalaya at the Sutlej river is

natural. The Sutlej river belongs to an ancient drainage pattern, established much

before the uplift of the Himalaya proper. The parts of the Himalaya west of the river

Sutlej are of more recent origin than the eastern part (see Mani 1968, 1986 for more

detail).

East Himalaya is relatively close to the Indian Ocean and plays an important role

in shaping the typical Indian monsoon climate. It is, in general, moist and therefore

relatively densely forested. The amount of moisture gradually diminishes toward the

continental and arid Northwest Himalaya. The general ecological conditions are

therefore drastically different at the eastern and western ends of the Himalaya.

While the Himalayan ranges east of the Sutlej river extend essentially in

east-west direction and lie between 27° and 29°N, those west of the river Sutlej

show an abrupt shift northward and lie between 30° and 36°N, and extend therefore

over a much wider area. There is an abrupt fall in the mean elevation of all ranges

west of the Sutlej river and unlike in the Himalaya east of the Sutlej river, there are

no rivers cutting through the main Himalayan range.

Revision of the Tribes Quediini and Atanygnathini

381

The tree line sinks from a mean elevation of about 3600 m in the east to almost

3000 m in the west. The permanent snowline in the east is at about 4500 m, but rises

to 5200 m in the arid northwest.

The east-west alignment of the ranges in most of the Himalaya results in

significant ecological differences between the southern and northern slopes of the

same range. Local climatic conditions may vary tremendously. For example, the

“rainshadow effect” is significant in some areas, e.g., in the valley of the Kali

Gandaki river in Nepal. The valley becomes increasingly arid north of the huge

massifs of Dhaulagiri in the west and Annapurna in the east, because these massifs

retain most of the monsoon rains. On the other hand, some areas on the south slope

receive enormous amounts of rain, e.g., the valley of the river Induwa in eastern

Nepal.

Distributions of the species

The study of the distributional ranges of the Himalayan Quediini and

Atanygnathini and their affinities show that they in general originated either 1 ) east

of the Himalaya, or 2) west of the Himalaya.

1 . Species of eastern origin.

A. The bulk of the Himalayan species can be derived from the ancestors that

originated in the Tertiary fold-mountains in the areas east and southeast of the

Himalaya, in what is today China, Thailand, Malaya and Greater Sundas. While

some originated apparently in distant centres, the Greater Sundas, Malaya and

Thailand (c.g., Ctenandropus nigriceps), others apparently had their origin in closer

centres, in Yunnan, Burma and Assam (e.g., some species groups of the subgenus

Raphirus of Quedius, or the species of Acylophorus close to A. heesoni). All species

of this origin are basically forest species occurring in low elevations, up to about

2600 m and are mainly distributed over East and Central Himalaya, and only

sporadically reach West Himalaya {e.g., Quedius apicicornis, Q. stevensi,

Bolitogyrus vuhieratus, Indocpiedius sikkimensis, Atanygnathus paani). Only one of

these species {Quedius aureivenths) managed to spread westward as far as Pakistan

and is today distributed across the Himalaya, from Pakistan to Bhutan. A few other

species {Quedius milansaar, Indocjuedius daai, Anchocerus monticola) occur only in

West Himalaya.

The distributional ranges of most species of this first group are restricted to the

Himalaya {e.g., Quedius heesoni, Q. ripicola, Indoquedius fUiconiis)\ however,

those of some other species extend southeast from the Himalaya and reach the

Burma-Thailand border near the river Salween (Kayah State) (e.g.. Quedius

rugosus) or reach even further south to Tenasserim in southern Burma {Heterothops

oculatus, Anchocerus hinnanus). It is worth mentioning that some of the species

restricted to the Himalaya also occur in the isolated hills in Meghalaya (Garo and

Khasi Hills) and in Nagaland (Naga Hills). On the other hand, there are some

species occurring in Khasi Hills {Indocpiedius saathi and Acylophorus halchhi), in

Quaest. Ent., 1988, 24 (2)

382

Smetana

Garo Hills {Acylophorus khairo, Atanygnathus pictus), and in Naga Hills {Quedius

aureipilis) that are at present not known from the Himalaya proper. These latter

species may or may not be endemic to these hills; I suspect that most of them

acutally do occur in the Himalaya proper and that their seeming endemism is due to

inadequate collecting.

B. A large group of species of the genus Quedius can apparently be derived from

ancestors that originated in the more northern centres, in Yunnan and Sichuan. This

is an important group of species, dominant in the subgenus Raphirus and also

represented in the subgenera Microsaurus and Distichalius. All species of this origin

are endemic to the Himalaya; they occur mainly in East and Central Himalaya and

only very few occur west of Central Himalaya. In the subgenus Distichalius, Q.

kashmirensis reaches Kashmir in Northwest Himalaya; two species of the subgenus

Raphirus have this distribution pattern: Q. daksumensis, extending again all the way

to Kashmir, and Q. kaalo, occurring in West Himalaya in the Almora area (Uttar

Pradesh). Some of the species of the subgenus Raphirus show distinct relationships

to species described from high mountains in Sichuan {e.g., Q. maculiventris

Bernhauer 1934, Q. optahilis Bernhauer 1934, Q. reitteriamis Bernhauer 1934). I

am convinced that there is in fact a much closer relationship between these areas,

but unfortunately our knowledge of the fauna of the Quediini of the mountains in

Sichuan is only fragmentary. Quedius kashmirensis (and as a matter of fact also the

two remaining species of Distichalius), on the other hand, is related to a fairly large

group of species distributed in China, Japan, and through Kamchatka and across the

Bering Strait on the Pacific Coast of North America (this is the “Capucinus Group”

in North America, particularly the species Q. nevadensis (Casey 1915) through Q.

hakeri Hatch 1957 in Smetana 1971 :V).

Ecologically, most of the species occur in the middle and upper forest zone, from

about 2600 m to almost tree line {e.g., Q. goropanus, Q. vadhu, Q. gaarho, Q.

kanyasa), but some of them also ascend into habitats above tree line (e.g., Q.

angnimai, Q. franzi\ only one species {Q. taruni) is basically alpine, occurring in

alpine tundra up to about 4400 m, but occasionally descending just below tree line.

Only one species of this group (Q. vadhu) occurs also on the mountain Phulcoki, the

highest peak of the Mahabharat Lekh at the southern edge of the Kathmandu Valley

in Nepal. On the other hand, one of the species {Q. udagra) occurs only near the top

of Phulcoki and is almost certainly endemic to this mountain.

In summary, the species of Quediini and Atanygnathini of eastern origin clearly

dominate in the Himalaya, constituting some 93% of the fauna.

While only one genus {Paratolmerus) in the tribe Quediini seems to be endemic

to the Himalaya (another endemic genus may be erected in the future for Quedius

gardneri, associated here tentatively with the subgenus Raphirus), an impressive

portion of the species is endemic to the Himalaya. The figures are as follows:

Quedius: 41 of 52 (78.8%), Indocjuedius: 5 of 8 (62.5%); Heterothops: 7 of 9

(77.8%); Acylophorus: 5 of 8 (62.5%) and Anchocerus: 3 of 4 (75%). The situation

Revision of the Tribes Quediini and Atanygnathini

383

is reversed in the tribe Atanygnathini: only 1 of 5 species of Atanygnathus (20%) is

endemic to the Himalaya. Even considering the fact that these figures may not

exactly reflect the actual situation, due to inadequate collecting in many areas, they

are nevertheless convincing. Considering the known number of species involved, the

rate of endemism is highest in the subgenus Raphiriis of the genus Quedius: 24

species of 30 (80%), followed closely by the subgenus Microsaurus: 14 species of

19 (73.7%). It is also worth emphasizing that most of the endemic species of

Raphiriis and Microsaurus occur mainly in the middle and upper forest zone, with

only 1 species being alpine (see above). Species diversity distinctly diminishes

toward the west, which is clearly correlated with increasing aridity (see above).

2. Species of western origin.

The species that can be derived from the ancestors that originated in the areas

west of the Himalaya (mountainous areas of Middle Asia that are today considered

as belonging to the Turkmenian subregion of the Palaearctic Region), are very few

and they all belong to the genus Quedius, except for one species of the genus

Heterothops. Since these species dispersed from the west, they had to appear in the

Himalaya later than those species from the eastern centres, certainly after the

Northwest Himalaya had been sufficiently uplifted and the Tethys Sea had been

almost obliterated (see Mani 1986). Quedius martensi, Q. dui, Q. adjacens, Q.

ochripennis, Q. fluviatilis and Heterothops saano belong to this category. Quedius

ochripennis is widely distributed in the areas west of the Himalaya and reaches

Europe, but the remaining species are endemic to the Himalaya. They occur in

Northwest Himalaya and in the western portion of the West Himalaya; only one

species {Q. martensi) reaches Central Himalaya in the Khumbu area in Nepal.

The species of this group form only an insignificant portion of the fauna of the

Quediini and Atanygnathini of the Himalaya (about 7%) and they belong to only

two genera.

3. The following table gives the known numbers of species of the nine genera in

different countries and states of India covered in this revision to allow quick

orientation. However, two factors should be considered. 1 ) The Nepal Himalaya is

at present by far the best collected area. Therefore, the very high number of species

listed for Nepal is probably out of proportion to those listed for the other areas; for

example, the five species listed for Bhutan undoubtedly represent only a fraction of

the species actually occurring there. 2) Only species included in Cameron (1932) are

listed for Burma (see the Introduction).

ACKNOWLEDGMENTS

In preparing this revision I relied on the assistance of many individuals who at

different levels and by different means contributed to its eompletion. I thank all of

them in the same way, no matter how small or large their contribution.

Quaest. Ent., 1988,24 (2)

384

Smetana

Table 1. Geographical distribution of the genera and species of the subfamily

Quediinae in the Indian Subcontinent.

Revision of the Tribes Quediini and Atanygnathini

385

The types and/or authentic specimens were made available to me through the

kindness of Mr. M.E. Baccus, P.M. Hammond and Mrs. S.L. Shute (British

Museum, Natural History, London, England); Dr. L. Baert (Institut royal des

sciences naturelles de Belgique, Bruxelles, Belgium); Dr. M. Brancucci

(Naturhistorisches Museum, Basel, Switzerland); Dr. H. Coiffait (Toulouse,

Erance); Dr. H. Eranz (Mddling, Austria); Dr. R. Gaedike (Institut fiir Pflanzen-

schutzforschung, Abteilung Taxonomic der Insekten [ehemaliges Deutsches

Entomologisches Institut], Eberswalde, German Democratic Republic); Dr. F.

Janczyk (Naturhistorisches Museum, Wien, Austria); Dr. N. Nikitskij (Zoological

Museum, Moscow Lomonosov State University, Moscow, USSR); Dr. W. Schedl

(Institut fiir Zoologie der Universitiit Innsbruck, Innsbruck, Austria); Dr. R. zur

Strassen (Senckenberg Museum, Frankfurt a.M., Federal Republic of Germany) and

Dr. L.E. Watrous (Field Museum of Natural History, Chicago, U.S.A.).

I studied most of Cameron’s and Champion’s types during my visit to the British

Museum (Natural History), London, in September 1984. I thank my colleagues there

for the use of their facilities and other assistance during my visit.

My colleagues and coworkers from the Coleoptera Unit of the Biosystematics

Research Centre, Ottawa, provided significant help. Mr. Go Sato in his own spare

time inked with great care and patience at least two thirds of all line drawings. Mr.

E. Rickey provided the scanning electron photomicrographs and Dr. L. LeSage

prepared the base distributional maps. Drs. E.C. Becker, Y. Bousquet and J.M.

Campbell read the manuscript and their comments eventually led to its

improvement.

My wife Zdena accompanied me on one of the Nepal expeditions; she helped

with the collecting and braved the strenuous trekking and the primitive living

conditions. My friend Ivan Ldbl provided exciting company and support on three of

the five Nepal expeditions. The Nepalese guides, particularly Ang Nima Sherpa,

who led three expeditions, Dorjee Sherpa Makalu and Sardar Tejbahadur Magar

(known as “TBS”), who each led one expedition, provided not only indispensable

guidance during the treks, but also took care of all the countless chores that

eventually led to the success of these expeditions. And last, but not least, the

contribution of the cooks, kitchen boys and numerous porters has to be gratefully

acknowledged. They took care of all our needs, so that we could spend most of our

time collecting. We had to admire the way they handled with ease and laughter

difficult and some dangerous situations, particularly at higher elevations, that would

upset most of us Westerners.

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(N.S.)No. 54:188-218.

Hatch, M.H. 1957. The beetles of the Pacific Northwest, Part II: Staphyliniformia.

University of Washington Publications in Biology 16, IX + 384 pp.

Herbst, J.F.W. 1784. Kritisches Verzeichnis meiner Insecten-Sammlung. In:

Fuessly: Arch. Naturg. 5:1-151, pi. 19-30.

Horn, G.H. 1878. Synopsis of the Quediini of the United States. Transactions of the

American Entomological Society 7:149-167.

Jakobson, G.G. 1909. Zhuki Rosii i zapadnoy Evropy. St. Petersbourg, 11 fasc.,

1024 pp., 83 pi. Unfinished.

Kraatz, G. 1856-1858. Staphylinii. Naturgeschichte der Insecten Deutschlands. II.

Berlin 8 + 1080 pp.

Kraatz, G. 1858. Einige neue und ausgezeichnete Staphylinen-Gattungen. Berliner

Entomologische Zeitschrift 2:361-368.

Kraatz, G. 1859. Die Staphylinen-Fauna von Ostindien, insbesondere der Insel

Ceylon. Archiv fiir Naturgeschichte 25:1-193, 3 pi. Sep. Berlin, 196 pp., 3 pi.

Kuhnt, P. 1913. Illustrierte Bestimmungs-Tabellen der Kafer Deutschlands.

Stuttgart, 10 + 1 138 pp.

Lea, A.M. 1925. On Australian Staphylinidae (Coleoptera). Part II. Transactions and

Proceedings of the Royal Society of Southern Australia 49:213-253.

Lohse, G.A. 1964. Staphylinidae I. (Micropeplinae bis Tachyporinae). In: Freude,

Harde, Lohse: Die Kafer Mitteleuropas. Vol. 4. Krefeld, 264 pp.

Lucas, R. 1920. Catalogus alphabeticus generum et subgenerum Coleopterorum

orbis terrarum totius (faml., trib., subtrib., sect. inch). Pars I. Berlin, XXXI + 696

pp.

Mani, M.S. 1968. Ecology and biogeography of high altitude insects. Series

entomologica. Vol. 4. Dr. W. Junk N.V. Publishers. The Hague. XIV + 527 pp.

Mani, M.S. 1986. Butterflies of the Himalaya. Series enomologica. Vol. 36. Dr. W.

Junk Publishers, Dortrecht. The Netherlands, and Oxford 104 Publishing Co.,

New Delhi, India. 181 pp.

Revision of the Tribes Quediini and Atanygnathini

389

Mannerheim, C.G. 1831. Precis d’un nouvel arrangement de la famille des

Brachelytres de I’ordre des insectes Coleopteres. Memoires presentes a

FAcademie Imperiale des Sciences de St. Petersbourg 1:415-501. Sep. 87 pp.

Menetries, E. 1832. Catalogue raisonne des objects de Zoologie recueillis dans un

voyage au Caucase et jusqu’aux frontieres actuelles de la Perse. St. Petersburg,

32 + 271 + 5 pp.

Motschulsky, V. 1858a. Enumeration des nouvelles especes de Coleopteres

rapportes de ses voyages. (Continuation). Bulletin de la Societe Imperiale des

naturalistes de Moscou 31, 2:634-670.

Motschulsky, V. 1858b. Enumeration des nouvelles especes de Coleopteres

rapportes de ses voyages. (Continuation). Bulletin de la Societe Imperiale des

naturalistes de Moscou 31, 3:204-264, 1 pi.

Mulsant E., C. Rey, 1875. Histoire naturelles des Coleopteres de France. Tribu des

Brevipennes. Staphyliniens. Annales de la Societe (Imperiale) d’Agriculture,

Histoire Naturelle et Arts Utiles de Lyon (8): 145-856, 6 pi.

Nordmann, A. 1837. Symbolae ad monographiam Staphylinorum. Petropoli, 167

pp., 1 pi.

Portevin, C. 1929. Histoire naturelle des Coleopteres de France, II. Adephaga,

Polyphaga: Staphylinoidea. Paris, 630 pp., 5 pi.

Reitter, E. 1909. Fauna Cermanica. Die Kafer des Deutschen Reiches. II. Stuttgart,

392 pp., pi. 41-80.

Scheerpeltz, O. 1965. Wissenschaftliche Ergebnisse der Schwedischen Expedition

1934 nach Indien und Burma - Coleoptera Staphylinidae (except.

Megalopsidiinae et Steninae). (5. Beitrag zur Kenntnis der orientalischen

Staphyliniden). Arkiv for Zoologi 17:93-371.

Scheerpeltz, O. 1974. Studien an der Cattung Cyrtothorax Kraatz. Mit Beschreibung

neuer Arten sowie einer Dichotomik aller bisher bekannt gewordenen Arten

dieser Cattung (Coleoptera, Staphylinidae, Staphylininae, Quediini). (14. Beitrag

zur Kenntnis der orientalischen Staphyliniden, gleichzeitig 27. Beitrag zur

Kenntnis der neotropischen Staphyliniden). Reichenbachia 15:175-192.

Scheerpeltz, O. 1976a. Wissenschaftliche Ergebnisse der von Prof. Dr. H.

Janetschek im Jahre 1961 in das Mt. Everest-Cebiet Nepals unternommenen

Studienreise. Khumbu Himal, Innsbruck 5:3-75.

Scheerpeltz, O. 1976b. Wissenschaftliche Ergebnisse entomologischer

Aufsammlungen in Nepal. Khumbu Himal, Innsbruck 5:77-173.

Schubert, K. 1908. Beitrag zur Staphylinidenfauna Ostindiens (West-Himalaya).

(Col.). Deutsche Entomologische Zeitschrift :609-625.

Sharp, D. 1874. The Staphylinidae of Japan. The Transactions of the Royal

Entomological Society of London : 1-103.

Sharp, D. 1889. The Staphylinidae of Japan. The Annals and Magazine of Natural

History, Series 6, 3:28^4, 108-121 , 249-267, 319-334, 406^19, 463-J76.

Shibata, Y. 1984. Provisional check list of the family Staphylinidae of Japan. IV.

Quaest. Ent., 1988, 24 (2)

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Smetana

(Insecta;Coleoptera). Annual Bulletin of the Nichidai Sanko No. 22:79-141.

Smetana, A. 1958. Fauna CSR. Vol. 12. Staphylinidae I. Praha, 437 pp.

Smetana, A. 1971. Revision of the tribe Quediini of America north of Mexico

(Coleoptera;Staphylinidae). Memoirs of The Entomological Society of Canada

No. 79. VI + 303 pp.

Smetana, A. 1975. A collection of Quediini from Nepal (Coleoptera:Staphylinidae).

Oriental Insects 9:323-342.

Smetana, A. 1976. Revision of the tribe Quediini of America north of Mexico

(Coleoptera:Staphylinidae). Supplementum 3. The Canadian Entomologist

108:169-184.

Smetana, A. 1977a. Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen

Museums in Basel. Coleoptera: Earn. Staphylinidae. Tribus Quediini.

Entomologica Basiliensia 2:242-250.

Smetana, A. 1977b. The Nearctic genus Beeria Hatch. Taxonomy, distribution and

ecology (Coleoptera, Staphylinidae). Entomologica Scandinavica 8:177-190.

Smetana, A. 1984. Le “culte de I’edeage”: reflexions additionelles, suives d’une

discussion sur le concept de la sous-tribu Heterothopsi Coiffait 1978

(Coleoptera, Staphylinidae). Nouvelle Revue d’Entomologie (N.S.) 1:277-282.

Stephens, H.F. 1829. The nomenclature of British Insects, ... London, 68 pp.

Stephens, H.F. 1832-1835. Illustrations of British Entomology ... Mandibulata.

London, 448 pp., pi. XXIV-XXVII.

Szujecki, A. 1980. Klucze do oznaczania owadow Polski. Part XIX. Coleoptera.

Issue 24e. Staphylinidae-Staphylininae. Warszawa, 164 pp.

Tottenham, C.E. 1955. Studies in the genus Philonthus Stephens (Coleoptera:

Staphylinidae). Parts II-IV. The Transactions of the Royal Entomological

Society of London 106:153-195.

Wagler, J.G. 1832. Monographia psittacorum. Abhandlungen der Mathematisch-

Physikalischen Classe der Koniglichen Bayerischen Akademie der

Wissenschaften 1:463-750, pi. 22-27.

Westwood, J.O. 1838. An introduction to the modern classification of Insects; ...

Vol. 2. Synopsis of the genera of British Insects. London, pp. 1^8.

Revision of the Tribes Quediini and Atanygnathini

391

Figs. 1-5. Quedius inquietus: 1, apical portion of male stemite 8; 2, aedoeagus, ventral view; 3, apical

portion of median lobe, ventral view; 4, apical portion of median lobe, lateral view; 5, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 6-9. Quedius apicicornis: 6, apical portion of male stemite 8; 7, aedoeagus, ventral view; 8, apical

portion of median lobe, with internal sac; 9, apical portion of paramere, underside. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

393

Figs. 10-13. Quedius beesoni: 10, apical portion of male stemite 8; 11, aedoeagus, ventral view; 12, apical

portion of median lobe, with internal sac; 13, apical portion of paramere, underside. Scale in mm.

Quaest. Ent., 1988, 24 (2)

Smetana

394

Figs. 14—17. 14-16, Quedius flavocaudatus: 14, apical portion of male stemite 8; 15, aedoeagus, ventral

view; 16, apical portion of paramere, underside with sensory peg setae. 17, Q. antennalis: aedoeagus, ventral

view. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

395

Figs. 18-21. 18-20, Quedius anteiuialis: 18, apical portion of male stemite 8; 19, apical portion of median

lobe, with internal sac; 20, apical portion of paramere, underside. 21, Q. martensi: apical portion of male

stemite 8. Scale in mm.

Quaest. Ent., 1988, 24 (2)

396

Smetana

Figs. 22-26. 22-23, Quedius martensi: 22, apical portion of median lobe, ventral view; 23, apical portion of

paramere, underside with sensory peg setae. 24-26, Q. dui\ 24, apical portion of male stemite 8; 25,

aedoeagus, ventral view; 26, apical portion of median lobe with internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

397

Figs. 27-31. 27, Quedius dui: apical portion of paramere underside with sensory peg setae. 28-31, Q.

adjacens: 28, apical portion of male stemite 8; 29, aedoeagus, ventral view; 30, apical portion of median

lobe with internal sac; 31, apical portion of paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

398

Smetana

Figs. 32-36. 32-34, Quedius ochhpennis: 32, apical portion of male stemite 8; 33, aedoeagus, ventral view;

34, apical portion of paramere, underside with sensory peg setae. 35, 36, Q. ptacidus: 35, apical portion of

male stemite 8; 36, aedoeagus, ventral view. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

399

Figs. 37, 38, Quedius placidus: 37, apical portion of median lobe with internal sac; 38, apical portion

of paramere, underside with sensory peg setae. 39-41, Q. lesagei: 39, apical portion of male stemite 8; 40,

aedoeagus, ventral view; 41, apical portion of paramere underside with sensory peg setae. 42, Q. stevensi:

male genital segment, dorsal view. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 43^7. 43, 44, Qiiedius stevensi: 43, apical portion of male stemite 8; 44, aedoeagus, ventral view.

45^7, Q. ripicola: 45, apical portion of male stemite 8; 46, aedoeagus, ventral view; 47, apical portion of

median lobe with internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

401

Figs. 48-54. 48, 49, Quediiis ripicola: 48, apical portion of median lobe, lateral view; 49, apical portion of

paramere, underside. 50-54, Q. milansaar: 50, apical portion of male stemite 8; 51, aedoeagus, ventral view;

52, apical portion of median lobe, with internal sac; 53, apical portion of median lobe, lateral view; 54,

apical portion of paramere, underside. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 55-60. 55-59, Quedius franzi: 55, apical portion of male stemite 8; 56, aedoeagus, ventral view; 57,

apical portion of median lobe with internal sac; 58, apical portion of median lobe, lateral view; 59, apical

portion of paramere, underside with sensory peg setae. 60, Q. goropanus: apical portion of male stemite 8.

Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

403

Figs. 61-65. 61-64, Quedius goropanus: 61, aedoeagus, ventral view; 62, apical portion of median lobe with

internal sac; 63, apical portion of median lobe, lateral view; 64, apical portion of paramere, underside with

sensory peg setae. 65, Q. tanderi: apical portion of male stemite 8. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 66-72. 66-68, Quedius tanderi: 66, aedoeagus, ventral view; 67, apical portion of median lobe with

internal sac; 68, apical portion of paramere, underside. 69-72, Q. kailo. 69, apical portion of male stemite 8;

70, aedoeagus, ventral view; 71, apical portion of median lobe with internal sac; 72, apical portion of

paramere, underside. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

405

Figs. 73-77. Quedius angnimai: 73, apical portion of male stemite 8; 74, aedoeagus, ventral view; 75, apical

portion of median lobe with internal sac; 76, 77, apical portions of parameres, undersides. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 78-82. 78-81, Quedius kashmirensis: 78, apical portion of male stemite 8; 79, aedoeagus, ventral

view; 80, apical portion of median lobe, ventral view; 81, apical portion of paramere, underside with sensory

peg setae. 82, Q. chatterjeei: apical portion of male stemite 8. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

407

Figs. 83-88. 83-86, Qiiedius chatterjeei: 83, aedoeagus, ventral view; 84, apex of median lobe, ventral view;

85, apex of median lobe, lateral view; 86, apical portion of paramere, underside with sensory peg setae. 87,

88, Q.fluviatilis: 87, apical portion of male stemite 8; 88, aedoeagus, ventral view. Scale in mm.

Quaest. Ent., 1988, 24 (2)

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Smetana

Figs. 89-94. 89, 90, Quedius fluviatilis: 89, apical portion of median lobe with internal sac; 90, apical

portion of paramere, underside with sensory peg setae. 91-94, Q. kaalo: 91, apical portion of male stemite 8;

92, aedoeagus, ventral view; 93, apical portion of median lobe with internal sac; 94, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

409

Figs. 95-102. Quedius vadhu: 95, 96, apical portions of male stemites 8; 97, 98, aedoeagi, ventral view; 99,

apical portion of median lobe with internal sac; 100, apical portion of median lobe, lateral view; 101, 102,

apical portions of parameres, undersides with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

Smetana

410

Figs. 103-111. 103-106, Quedius gaarho: 103, apical portion of male stemite 8; 104, aedoeagus, ventral

view; 105, apical portion of median lobe with internal sac; 106, apical portion of paramere, underside with

sensory peg setae. 107-1 \ \, Q. daksumensis: 107, apical portion of male stemite 8; 108, aedoeagus, ventral

view; 109, apical portion of median lobe with internal sac; 1 10, apical portion of median lobe, lateral view;

111, apical portion of paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

411

view; 1 14, apical portion of median lobe with internal sac; 1 15, apical portion of median lobe, lateral view;

116, apical portion of paramere, underside with sensory peg setae. 117-120, Q. aureiventris. 117, apical

portion of male stemite 8; 118, aedoeagus, ventral view; 119, apical portion of median lobe with internal

sac; 120, apical portion of median lobe, lateral view. Scale in mm.

Quaest. Ent., 1988, 24 (2)

412

Smetana

Figs. 121-129. 121, Quedius aureiventris: apical portion of paramere, underside with sensory peg setae.

122-129, Q. muscicola: 122, 123, apical portions of male stemite 8; 124-126, aedoeagi, ventral views;

127-129, apical portions of median lobes, ventral view, 129 with evaginated internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

413

132

134

0.4

0.2

Figs. 130-136. 130-132, Qiiedius muscicola: apical portions of parameres, undersides with sensory peg

setae. 133-136, Q. hliari: 133, apical portion of male stemite 8; 134, aedoeagus, ventral view; 135, apical

portion of median lobe with internal sac; 136, apical portion of paramere, underside with sensory peg setae.

Scale in mm.

Quaest. Ent., 1988, 24 (2)

414

Smetana

Figs. 137-145. 137-140, Quedius eklai: 137, apical portion of male stemite 8; 138, aedoeagus, ventral view;

139, apical portion of median lobe with internal sac; 140, apical portion of paramere, underside with sensory

peg setae. 141-145, Q. sundar. 141, apical portion of male stemite 8; 142, aedoeagus, ventral view; 143,

apical portion of median lobe with internal sac; 144, apical portion of median lobe, lateral view; 145, apical

portion of paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

415

Figs. 146-150. Quedius udagra: 146, apical portion of male stemite 8; 147, aedoeagus, ventral view; 148,

apical portion of median lobe with internal sac; 149, apical portion of median lobe, lateral view; 150, apical

portion of paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

416

Smetana

Figs. 151-156. 151-155, Quedius satoi: 151, apical portion of male stemite 8; 152, aedoeagus, ventral view;

153, apical portion of median lobe with internal sac; 154, apical portion of median lobe, lateral view; 155,

apical portion of paramere, underside with sensory peg setae. 156, Q. kanyasa: apical portion of male

stemite 8. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini 417

Figs. 157-164. 157-160, Quedius kanyasa: 157, aedoeagus, ventral view; 158, apical portion of median lobe

with internal sac; 159, 160, apical portions of parameres, undersides with sensory peg setae. 161-164, Q.

kanyasa (holotype of Q. dhaulagirensis): 161, apical portion of male stemite 8; 162, aedoeagus, ventral

view; 163, apical portion of median lobe with internal sac; 164, apical portion of paramere, underside with

sensory peg setae. Scale in mm.

Quaest. Ent., 1988,24 (2)

418

Smetana

Figs. 165-171. 165-169, Quedius naati: 165, apical portion of male stemite 8; 166, aedoeagus, ventral view;

167, apical portion of median lobe with internal sac; 168, apical portion of median lobe, lateral view; 169,

apical portion of paramere underside with sensory peg setae. 170, 171,0. dewar 170, apical portion of male

stemite 8; 171, male genital segment, dorsal view. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

419

Figs. 172-179. 172-174, Qnedius dewar: 172, aedoeagus, ventral view; 173, apical portion of median lobe

with internal sac; 174, apical portion of paramere, underside with sensory peg setae. 175-179, Q. tikta: 175,

apical portion of male stemite 8; 176, aedoeagus, ventral view; 177, apical portion of median lobe with

internal sac; 178, apical portion of median lobe, lateral view; 179, apical portion of paramere, underside with

sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

420

Smetana

Figs. 180-189. 180-184, Quedhis tonglw. 180, apical portion of male stemite 8; 181, aedoeagus, ventral

view; 182, apical portion of median lobe with internal sac; 183, apical portion of median lobe, lateral view;

184, apical portion of paramere, underside with sensory peg setae. 185-189, Q. pharak: 185, apical portion

of male stemite 8; 186, aedoeagus, ventral view; 187, apical portion of median lobe with internal sac; 188,

apical portion of median lobe, lateral view; 189, apical portion of paramere, underside with sensory peg

setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

421

Figs. 190-197. 190-194, Qiiedius atchala: 190, apical portion of male stemite 8; 191, aedoeagus, ventral

view; 192, apical portion of median lobe with internal sac; 193, apical portion of median lobe, lateral view,

with internal sac; 194, apical portion of paramere, underside with sensory peg setae. 195-197, Q. durgaa:

195, apical portion of male stemite 8; 196, aedoeagus, ventral view; 197, apical portion of median lobe with

internal sac. Scale in mm.

Quae St. Ent., 1988, 24 (2)

422

Smetana

Figs. 198-201. 198, Quedius durgaa: apical portion of paramere, underside with sensory peg setae.

199-201, Q. anomalus: 199, apical portion of male stemite 8; 200, aedoeagus, ventral view; 201, apical

portion of median lobe with internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

423

Figs. 202-205. 202, Quedius anomalus: apical portion of paramere, underside with sensory peg setae.

203-205, Q. assamensis: 203, apical portion of male stemite 8; 204, aedoeagus, ventral view; 205, apical

portion of aedoeagus with internal sac, lateral view. Scale in mm.

Qiiaest. Ent., 1988, 24 (2)

424

Smetana

Figs. 206-210. 206, Quedius assamensis: apical portion of paramere, underside with sensory peg setae.

207-210, Q. himalayiciis: 207, apical portion of male stemite 8; 208, aedoeagus, ventral view; 209, apical

portion of median lobe, lateral view; 210, apical portion of paramere, underside with sensory peg setate.

Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

425

Figs. 21 1-215. 21 1-214, Qiiedius nilo: 211, apical portion of male stemite 8; 212, aedoeagus, ventral view;

213, apical portion of median lobe, lateral view; 214, apical portion of paramere, underside with sensory peg

setae. 215, 0. kiuro: apical portion of male stemite 8. Scale in mm.

Quaest. Ent., 1988, 24 (2)

426

Smetana

222

Figs. 216-223. 216, 217, Quediiis kuiro: 216, aedoeagus, ventral view; 217, apical portion of paramere,

underside with sensory peg setae. 218-223, Q. aureipilis'. 218, apical portion of male sternite 8; 219, tergite

10 of male genital segment; 220, stemite 9 of male genital segment; 221, aedoeagus, ventral view; 222,

apical portion of median lobe, lateral view; 223, apical portion of paramere, underside with sensory peg

setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

427

Figs. 224—227. Qiiedius haryio: 224, apical portion of male stemite 8; 225, aedoeagus, ventral view; 226,

apical portion of median lobe with internal sac; 227, apical portion of paramere, underside with sensory peg

setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

428

Smetana

Figs. 228-235. 228-231, Quedius rugosus: 228, apical portion of male stemite 8. 229, aedoeagus, ventral

view; 230, apical portion of median lobe with internal sac; 231, apical portion of paramere, underside with

sensory peg setae. 232-235, Q. gardneri: 232, apical portion of male stemite 8; 233, aedoeagus, ventral

view; 234, apical portion of median lobe with internal sac; 235, apical portion of one branch of paramere.

Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

429

Figs. 236-242. Quediiis lineipennis: 236, apical portion of male stemite 8; 237, tergite 10 of male genital

segment; 238, stemite 9 of male genital segment; 239, aedoeagus, ventral view; 240, apical portion of

median lobe, with evaginated internal sac; 241, apex of median lobe, lateral view; 242, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

Smetana

430

Figs. 243-246. Indoquedius sikkimensis: 243, apical portion of male stemite 8; 244, aedoeagus, ventral view;

245, apical portion of median lobe with internal sac; 246, apical portion of paramere, underside with sensory

peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

431

Figs. 247-250. Indoqiiedius baliyo: 247, apical portion of male stemite 8; 248, aedoeagus, ventral view; 249,

apical portion of median lobe, with evaginated internal sac; 250, apical portion of paramere, underside with

sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

432

Smetana

Figs. 251-257. 251-254, Indoquedius bipunctatus: 251, apical portion of male sternite 8; 252, aedoeagus,

ventral view; 253, apical portion of median lobe with internal sac; 254, apical portion of paramere, underside

with sensory peg setae. 255-257, /. daai: 255, apical portion of male sternite 8; 256, aedoeagus, ventral

view; 257, apical portion of paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

433

Figs. 258-262. 258, Indoqiiediiis daai: apical portion of median lobe with internal sac. 259-262, 1.filicornis:

259, apical portion of male stemite 8; 260, aedoeagus, ventral view; 261, apical portion of median lobe with

internal sac; 262, apical portion of paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

434

Smetana

view; 265, apical portion of median lobe with internal sac; 266, apical portion of paramere underside with

sensory peg setae. 267-269, 1, aberrans: 261, apical portion of male stemite 8; 268, aedoeagus, ventral view;

269, apical portion of paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

435

Figs. 270-274. Bolitogyrus vulneratus: 270, apical portion of male stemite 8; 271, tergite 10 of male genital

segment; 272, stemite 9 of male genital segment; 273, aedoeagus, ventral view; 274, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Qiiaest. Ent., 1988, 24 (2)

436

Smetana

Figs. 275-283. 275-280, Heterothops oculatus: 275, 276, male stemite 8; 211 , 278, aedoeagus, ventral view;

279, 280, apical portion of median lobe with internal sac. 281-283, H. indicus: 281, male stemite 8; 282,

aedoeagus, ventral view; 283, apical portion of median lobe with internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

437

Figs. 284—292. 284—287, Heterothops pusillus: 284, male sternite 8; 285, aedoeagus, ventral view; 286,

apical portion of median lobe with internal sac; 287, apex of median lobe, with evaginated internal sac.

288-290, H. hindustanus: 288, male sternite 8; 289, aedoeagus, ventral view; 290, apical portion of median

lobe with internal sac. 291, 292, H. khairo: 291, aedoeagus, ventral view; 292, apical portion of median lobe

with internal sac. Scale in mm.

Quaest. Ent., 1988, 24 (2)

438

Smetana

Figs. 293-300. 293-295, Heterothops persimilis: 293, male stemite 8; 294, aedoeagus with evaginated

internal sac; 295, apical portion of median lobe, with evaginated internal sac. 296-300, H.franzi: 296, male

stemite 8; 297, male genital segment, dorsal view; 298, aedoeagus, ventral view; 299, 300, apex of median

lobe with partially and fully evaginated internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

439

Figs. 301-307. 301-303, Heterothops saphaa: 301, male stemite 8; 302, aedoeagus, ventral view; 303,

apical portion of median lobe, with partially evaginated internal sac. 304-307, Ctenandropiis nigriceps: 304,

male tergite 8; 305, male stemite 8; 306, male genital segment, dorsal view; 307, aedoeagus with internal

sac. Scale in mm.

Quaest. Ent., 1988,24 (2)

440

Smetana

hiiri

Figs. 308-311. Paratolmerus pilosiventris: 308, male genital segment, dorsal view; 309, aedoeagus, ventral

view; 310, apical portion of median lobe with internal sac; 311, paramere, underside with sensory peg setae.

Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

441

Figs. 312-319. 312, 313, Acylophorus beesoni: 312, aedoeagus, ventral view; 313, apical portion of

aedoeagus, with internal sac, and apical portion of paramere with sensory peg setae. 314, 315, A. chillo: 314,

aedoeagus, ventral view; 315, apical portion of aedoeagus, with internal sac, and apical portion of paramere

with sensory peg setae. 316, 317, A. khairo: 316, aedoeagus, ventral view; 317, apical portion of paramere,

underside with sensory peg setae. 318, 319, A. siyo\ 318, aedoeagus, ventral view; 319, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Quaest. Ent., 1988, 24 (2)

442

Smetana

322, apical portion of median lobe with internal sac; 323, apical portion of paramere, underside with sensory

peg setae. 324-327, A. daar. 324, male genital segment, dorsal view; 325, aedoeagus, ventral view; 326,

apical portion of median lobe with internal sac; 327, apical portion of paramere, underside with sensory peg

setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

443

Figs. 328-335. 328-330, Acylophorus ruficollis\ 328, aedoeagus, ventral view; 329, apical portion of median

lobe with internal sac; 330, apical portion of paramere, underside with sensory peg setae. 331-333, A.

halchhi: 331, aedoeagus, ventral view; 332, apical portion of median lobe with internal sac, ventral view;

333, same, lateral view. 334, 335, A. raato: 334, aedoeagus, ventral view; 335, apical portion of median lobe

with internal sac. Scale in mm.

Quae St. Ent., 1988, 24 (2)

444

Smetana

Figs. 336-343. 336, 337, Acylophorus microcephalus: 336, aedoeagus, ventral view; 337, apical portion of

median lobe with internal sac. 338, 339, A. charaa: 338, aedoeagus ventral view; 339, apical portion of

median lobe with internal sac. 340, 341, A. flavipes: 340, aedoeagus, ventral view; 341, apical portion of

median lobe with internal sac. 342, 343, A. tibialis: 342, aedoeagus, ventral view; 343, apical portion of

paramere, underside with sensory peg setae. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

445

Figs. 344-350. 344-346, Acylophonis puncticeps: 344, aedoeagus, ventral view; 345, apical portion of

median lobe with internal sac; 346, right (in Fig. 344) branch of paramere with sensory peg setae. 347, 348,

Anchoceriis monticola: 347, aedoeagus, ventral view; 348, internal sac, evaginated. 349, 350, A. hinnanus:

349, aedoeagus, ventral view; 350, apical portion of median lobe with internal sac. Scale in mm.

Qiiaest. Ent., 1988, 24 (2)

446

Smetana

Figs. 351-356. 351-353, Anchocerus nepalicus: 351, male genital segment, dorsal view; 352, aedoeagus,

ventral view; 353, apical portion of median lobe with internal sac. 354, 355, Atanygnathus pictus: 354,

aedoeagus; 355, apical portion of aedoeagus with internal sac. 356. A. sasuraa: male stemite 8. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

447

Figs. 357-362. 357-359, Atanygnathus sasuraa: 357, male genital segment, dorsal view; 358, aedoeagus,

ventral view; 359, apical portion of aedoeagus with internal sac. 360-362, A. hrevicollis: 360, male stemite

8; 361, aedoeagus, ventral view; 362, apical portion of aedoeagus with internal sac. Scale in mm.

Quaest. Ent., 1988, 24 (2)

448

Smetana

Figs. 363-371. 363-365, Atanygnathus piceus: 363, male stemite 8; 364, aedoeagus, ventral view; 365,

apical portion of aedoeagus with internal sac. 366-368, A. paani: 366, male stemite 8; 367, aedoeagus,

ventral view; 368, apical portion of aedoeagus with internal sac. 369-371, A. bindw. 369, male stemite 8;

370, aedoeagus, ventral view; 371, apical portion of aedoeagus with internal sac. Scale in mm.

Revision of the Tribes Quediini and Atanygnathini

449

Figs. 372-378. 372, Atanygnathiis hindu: apex of aedoeagus with evaginated internal sac. 373-375, A. chiso:

373, male stemite 8, 374, aedoeagus, ventral view; 375, apical portion of aedoeagus with internal sac.

376-378, A. purba: 376, male stemite 8; 377, aedoeagus, ventral view; 378, apical portion of aedoeagus with

internal sac. Scale in mm.

Quaest. Ent., 1988,24 (2)

450

Smetana

Figs. 379-384. Qeudius apicicornis: 379, head dorsally; 380. base of left antenna; 381, male front

tarsus; 382, apical margin of tergite 7 with palisade setae; 383, same, detail; 384, female genital segment

dorsally.

Figs. 379-384. Qeudius apicicornis: 379, head dorsally; 380, base of left antenna; 381, male tront tarsus;

382, apical margin of tergite 7 with palisade setae; 383, same, detail; 384, female genital segment dorsally.

Revision of the Tribes Quediini and Atanygnathini

451

Figs. 385-391. 385-387, Quedius apicicornis: 385, female genital segment dorsally, detail; 386, female

genital segment ventrally; 387, same, detail. 388-391, Q. tikta: 388, base of left antenna; 389, last segment

of middle tarsus with claws; 390, scutellum; 391, apical margin of tergite 7.

Quaest. Ent., 1988, 24 (2)

452

Smetana

Figs. 392-395. Quediits tikta: 392, female genital segment dorsally; 393, same, detail; 394, female genital

segment ventrally; 395, same, detail.

Revision of the Tribes Quediini and Atanygnathini

453

Figs. 396-399. Qiiediiis stevensi: 396, female genital segment dorsally; 397, same, detail; 398, female

genital segment ventrally, 399, same, detail.

Qiiaest. Ent., 1988, 24 (2)

454

Smetana

Figs. 400-404. 400-403, Indoquedius filicornis: 400, left half of head dorsally; 401, base of left antenna;

402, female genital segment dorsally; 403, female genital segment ventrally. 404. Heterothops persimilis,

female genital segment dorsally.

Revision of the Tribes Quediini and Atanygnathini

455

Figs. 405-408. 405^07, Heterothops persimilis: 405, female genital segment dorsally, detail; 406, female

genital segment ventrally; 407, same, detail. 408, Acylophorus furcatus: head dorsally.

Quaest. Ent., 1988, 24 (2)

456

Smetana

Figs. 409^14. Acylophorus furcatiis: 409, base of left antenna. 410, apical margin of tergite 7, with

detail of palisade setae; 411. palisade setae at apical margin of tergite 7 and punctation of tergite 8; 412,

hind tarsus; 413, hind tarsal claws; 414, front tarsal claws.

Figs. 409^14. Acylophonis furcatus: 409, base of left antenna. 410, apical margin of tergite 7, with detail of

palisade setae; 411, palisade setae at apical margin of tergite 7 and punctation of tergite 8; 412, hind tarsus;

413, hind tarsal claws; 414, front tarsal claws.

Revision of the Tribes Quediini and Atanygnathini

457

Figs. 415-418. 415, Acylophorus fiircatus: lateral portion of pronotum. 416, A. raato: female genital

segment dorsally. 417, 418, Atanygnathus sasuraa: 417, female genital segment dorsally; 418, same, detail.

Quaest. Ent., 1988,24 (2)

458

Smetana

Figs. 419^21. 419, 420, Atanygnathiis sasuraa: 419, female genital segment ventrally; 420, same, detail. 421, Quedius tikta: head dorsally.

Revision of the Tribes Quediini and Atanygnathini

459

INDEX TO NAMES OE TAXA

(Synonyms in italics)

EAMILY GROUP TAXA

Atanygnathini, 167, 178, 368-369

Quediini, 167-168, 178-179, 334,

336, 382

Staphylini, 336

Staphylinidae, 169

Staphylininae, 169, 177

Staphylinini, 179, 336

Tanygnathini, 368

Tanygnathinini, 368

Xantholininae, 169, 177

Xanthopygi, 179

GENERA AND SUBGENERA

Acylophorus Nordmann, 168, 179,

181,336-338,361-362,381

Algon Sharp, 168, 179

Anchocerus Eauvel, 168, 181,

361-362

Atanygnathus Jacobson, 168,

178- 179,318,368

Barypalpus Cameron, 168, 179

Bolitogyrus Chevrolat, 167,

179- 180,314

Ctenandropus Cameron, 168,

179-180, 334

Cyrtothorax Kraaiz, 167-168, 314

Distichalius Casey, 181, 183-184,

227, 382

Erichsonius Eauvel, 339

Gabronthus Tottenham, 319

Heterothops Stephens, 168,

179-180,318,334

Indoquedius Cameron, 179-180,

300

Microsaurus Dejean, 181,

183-185, 192,382

Paratolmerus Cameron, 179, 181,

335-336, 338, 382

Quedius Stephens, 168, 179-181,

381

Raphirus Stephens, 181, 183-185,

231,252,267, 381-382

Rientis Shaip, 179

Sauridus Mulsant et Rey, 181, 183

Securipcilpus Schubert, 168, 179

Tachyporus Gravenhorst, 368

Tanygnathimis Reitter, 368

TcmygmitJms Erichson, 368

Tolmerinus Bernhauer, 336

Velleius Mannerheim, 168, 192

SPECIES AND SUBSPECIES

aberrans (Cameron), Indoquedius,

301,311

aherrans Cameron, Quedius, 3 1 1

adjacens Cameron, Quedius, 168,

185, 188,205-206,383

ahrensii Nordmann, Acylophorus,

337

akalita (Smetana), Indoquedius,

303

akalita Smetana, Quedius, 169,

302

analoka Smetana, Quedius, 169,

212-213

angnimai sp. nov., Quedius, 182,

186, 189,222-224,382

anomalus Cameron, Quedius, 168,

234, 274-275

antennalis Cameron, Quedius, 182,

185-186, 201-202

apicicornis Eppelsheim, Quedius,

168-169, 182, 184-185, 187,

192-194, 196, 198,381

assamensis Cameron, Quedius,

Quaest. Ent., 1988,24 (2)

460

Smetana

234, 284-285, 287, 379

atchala Smetana, Quedius, 239,

264, 271

attenuatus Gravenhorst,

Staphylinus, 231

aureipilis Cameron, Quedius, 234,

236, 285, 287-289, 382

aureiventris Bernhauer, Quedius,

168,250-252, 381

bakeri Hatch, Quedius, 382

balchhi sp. nov., Acylophorus,

339-340, 350-351,381

baliyo sp. nov., Indoquedius, 301,

304-305

basalis Motschulsky, Erichsonius,

339

beesoni Cameron, Acylophorus,

168,339,341-344, 381

beesoni Cameron, Quedius,

182-186, 196, 198, 381

bhari sp. nov., Quedius, 236,

256-257

bindu sp. nov., Atanygnathus, 369,

376-377, 379

binotatus Gravenhorst,

Staphylinus, 318

bipunctatus (Eppelsheim),

Indoquedius, 305-308

bipunctatus Cameron,

Acylophorus, 174, 338-339,

358-359

bipunctatus Eppelsheim, Quedius,

168,305

birmanus Cameron, Quedius, 185,

187,202

birmanus Fauvel, Anchocerus,

168,361-366, 381

brevicollis (Fauvel),

Atanygnathus, 168, 370,

373-376

brevicollis Fauvel, Tanygnathus,

373

buphthalmus (Erichson),

Bolitogyrus, 314

buphthalmus (Erichson),

Cyrtothora.v, 3 1 5

buphthalmus Erichson, Quedius,

314

cachemiricus Coiffait, Quedius,

239, 241

capucinus Gravenhorst,

Staphylinus, 227

charaa spec, nov., Acylophorus,

339-340, 355-356

chatterjeei Cameron, Quedius,

168,227,229-231

chillo sp. nov., Acylophorus, 339,

341,343

chiso sp. nov., Atanygnathus, 370,

377, 379

chlorophanus Erichson, Quedius,

167, 183-184

coeruleus Coiffait, Quedius, 169,

295

conicus Champion, Quedius, 168,

309

cribripennis Chevrolat,

Bolitogyrus, 314-315

cyanipennis Coiffait, Algon,

302-303

cyanopterus Sheerpeltz, Quedius,

169, 192-193

daai sp. nov., Acylophorus, 339,

341,345,348-349

daai sp. nov., Indoquedius, 301,

307-310, 381

daksumensis Coiffait, Quedius,

169, 237,246-248,252, 382

deceptor Cameron, Quedius, 168,

227,231

decipiens Cameron, Quedius, 168,

250-252

deuvei Coiffait, Quedius, 228

dewar sp. nov., Quedius, 239,

Revision of the Tribes Quediini and Atanygnathini

461

265-266

dhaulagirensis Coiffait, Quedius,

262-263

dohertyi Cameron, Quedius,

253-255

dui sp. now, Quedius, 185, 187,

204-205, 383

durgaa sp. now, Quedius, 234,

273-274

eklai sp. now, Quedius, 237, 257

filicomis (Eppelsheim),

Indoquedius, 301-302,

307-310, 381

filicomis Eppelsheim, Quedius,

168,309

flaviceps Motsehulsky,

Acylophorus, 167, 174, 340,

356-357

flavicollis Motsehulsky,

Heterothops, 319

flavocaudatus Cameron, Quedius,

168, 185-186, 200-201

flavotenninatus Cameron,

Quedius, 192, 194

fluviatilis Cameron, Quedius, 168,

233,239-241,243,383

franzi Coiffait, Heterothops, 320,

326, 328

franzi Smetana, Quedius, 181, 186,

189,217,219-221,382

frigidus Smetana, Quedius, 244

fulgidus (Eabricius), Quedius, 167,

185, 188,209-210

fulgidus Eabricius, Staphylinus,

209

fulvicollis Stephens, Quedius, 244

furcatus Motsehulsky,

Acylophorus, 167, 339, 341,

345, 348-349, 352

fuscus (Kraatz), Atanygnathus,

167, 374-375

fuscus Kraatz, Tanygnathiis,

374-375

gaarho sp. now, Quedius, 237,

246-248, 264, 382

gardneri Cameron, Quedius, 169,

179, 182-184, 233,295-297,

382

glaherrinus Herbst, Staphylinus,

337

goropanus Smetana, Quedius, 186,

189, 220-223,382

hariyo sp. now, Quedius, 233,

291-292

heterogaster Cameron, Quedius,

168,253-254

himalayicus Bernhauer, Quedius,

168,235,285-288

liindustana Cameron, Heterothops,

325

hindustanus Cameron,

Heterothops, 320, 325-326

indica Cameron, Heterothops, 322

indicus Cameron, Heterothops,

168,319,321-323,328

inquietus (Champion), Quedius,

183, 185-186, 189-190

inquietus Champion, Velleius, 168,

189

intricatus Fauvel, Quedius, 168,

233, 295

juno Shai-p, Indoquedius, 300, 306

kaalo sp. now, Quedius, 235,

243-244, 382

kailo sp. now, Quedius, 182, 186,

189, 222-223,288

kanyasa Smetana, Quedius, 238,

262-265, 382

kashmirensis Cameron, Quedius,

168-169, 181,227-229,382

kashmiricus Cameron, Quedius,

239, 241

khairo sp. now, Acylophorus, 339,

341,344-345,382

Quaest. Ent., 1988, 24 (2)

462

Smetana

khairo sp. nov., Heterothops, 320,

326

kuiro sp. now, Quedius, 234,

286-288

lama Coiffait, Quedius, 262-263

lateralis Gravenhorst, Staphylinus,

185

leptocephalus Coiffait, Quedius,

189

lesagei sp. now, Quedius, 185,

187,211-212

limbatus Heer, Quedius, 243

lineipennis Cameron, Quedius,

179, 181-184, 190, 297-298

maculiventris Bernhauer, Quedius,

382

magniceps (Bernhauer),

Ctenandropus, 334

mandra Smetana, Quedius, 169,

228

martens! Smetana, Quedius, 185,

188, 203-205

microcephalus Cameron,

Acylophorus, 339-340, 352,

355-356

microcerus Fauvel, Acylophorus,

167

milansaar 5'p. nov., Quedius, 182,

185, 188,215,217,381

mimeticus Cameron, Quedius,

196-198

monticola Cameron, Anchocerus,

168, 362-363,381

montivagus Smetana, Quedius,

229

multipunctatus Sharp, Quedius,

289

muscicola Cameron, Quedius, 168,

236, 253-257, 259, 262, 383

mussooriensis Cameron, Quedius,

292-293

naati sp. now, Quedius, 238-239,

264-266

nemoralis Baudi, Quedius, 243

nepalicus Coiffait, Acylophorus,

339

nepalicus sp. now, Anchocerus,

362, 365-366

nevadensis Casey, Quedius, 229,

382

nichinaiensis Casey, Quedius, 239,

241

nigriceps Cameron, Ctenandropus,

168,334-335,381

nilo sp. now, Quedius, 235,

286-287, 379

notahilis Cameron, Quedius,

196-198

ochripennis Menetries, Quedius,

167,207,209-210,383

ochripennis Menetries,

Staphylinus, 207

oculatus (Fauvel), Indoquedius,

301, 306-307

oculatus Fauvel, Heterothops, 168,

319-322,381

oculatus Fauvel, Quedius, 168,

300, 306

optabilis Bernhauer, Quedius, 382

paani sp. now, Atanygnathus, 370,

375-376,379,381

paschim sp. now, Quedius, 237,

249, 252

perqffinis Cameron, Quedius,

196-198

persimilis Cameron, Heterothops,

320, 326-328

pharak sp. now, Quedius, 238,

268-269

piceus (Motschulsky),

Atanygnathus, 167, 174, 370,

374-375

piceus Motschulsky, Tanygnathus,

370, 374

Revision of the Tribes Quediini and Atanygnathini

463

pictus (Motschulsky),

Atanygnathus, 1 67, 369-37 1 ,

382

pictus Motschulsky, Tanygnathus,

371

pilosiventris Cameron,

Paratolmerus, 335-337, 361

placidus Cameron, Quedius, 182,

185, 187,210-212

praeditus Shaip, Indoquedius, 300,

306

prostans Horn, Quedius, 272

pulchellus Kraatz, Gabronthus,

319

punctatissimus Smetana,

Anchocerus, 169, 362,

365-366

puncticeps Fauvel, Acylophorus,

168,338-339, 341,359, 361

purba sp. now, Atanygnathus, 370,

379

pusilla Coiffait, Hetevothops, 323

pusillus Coiffait, Heterothops, 319,

323-324

queinneci Coiffait, Quedius,

2 1 2-2 1 3

raato sp. now, Acylophorus,

339-340, 351-352

reitterianus Bernhauer, Quedius,

382

ripicola Cameron, Quedius, 168,

182, 185, 188,215-217,381

ruficoUis (Kraatz), Atanygnathus,

167,371-372

ruficoUis Kraatz, Acylophorus,

349-350

ruficoUis Kraatz, Tanygnathus,

370-371

ruficoUis Motschulsky,

Acylophorus, 167, 339,

349-350

ruficomis Cameron, Barypalpus,

179

rugosus Cameron, Quedius, 233,

292-295,381

saano sp. nor., Heterothops, 319,

329, 383

saathi sp. now, Indoquedius, 302,

310,381

saphaa sp. now, Heterothops, 320,

330

sasuraa sp. now, Atanygnathus,

369, 372-373

satoi sp. now, Quedius, 235, 256,

260-262

seriatus Horn, Quedius, 272

sherpai Coiffait, Quedius,

192-193, 196

sikkimensis (Cameron),

Indoquedius, 169, 301-305,

381

sikkimensis Cameron, Quedius,

302-303

siyo sp. now, Acylophorus, 339,

341, 344-345, 348

sodalis Cameron, Quedius, 168,

215-216

spectabilis Kraatz, Quedius, 167,

183

stevensi Cameron, Quedius, 169,

185, 188,212-213,215,217,

381

sundar sp. now, Quedius, 236,

258-259

tanderi sp. now, Quedius, 182,

186, 189, 222-223

taruni sp. now, Quedius, 181, 184,

233,243,271-272,382

terminalis Erichson, Atanygnathus,

369

terminalis Erichson, Tanygnathus,

368

tetricus Smetana, Quedius, 229

tibialis Cameron, Acylophorus,

Quaest. Ent., 1988, 24 (2)

464

Smetana

339-340, 357-358

tikta Smetana, Quedius, 238, 262,

266- 269

tonglu sp. now, Quedius, 238,

267- 269

tristis Gravenhorst, Staphylinus,

181

trisulensis Coiffait, Quedius, 217,

219

udagra Smetana, Quedius, 239,

259-260, 262, 382

vadhu Smetana, Quedius, 238,

245-248, 252, 382

ventralis Coiffait, Acylophorus,

342

vulneratus (Fauvel), Bolitogyrus,

167,315-316,318, 381

vulneratus Fauvel, Cyrtothorax,

315

Book Review

465

BOOK REVIEW

JAMIESON, B. G. M. 1987. The Ultrastructure and Phylogeny of Insect

Spermatozoa. Cambridge University Press, Cambridge, xv + 320 pp., 159 figs.,

author and subject indices. 30 pounds , U. K.

Most zoologists probably recall animal spermatozoa as resembling the miniature

tadpoles figured in many introductory textbooks of biology or zoology (our own

having a ghostly little bloke squeezing his knees inside). But, as emphasized by B.

A. Afzelius in the foreword to this book, and as vividly illustrated on its pages, these

cells are as diverse as the male animals from which they come. This comprehensive

new book rigorously surveys the known ultrastructure of this diversity within a

cladistic framework for members of the phylum ‘Uniramia’ (Onychophora,

Myriapoda, Hexapoda) of Tiegs, Manton and Anderson but with particular emphasis

on the sperm of insects.

The book is dedicated to Baccio Baccetti of the University of Siena in Italy, who,

with his students and colleagues, has probably described the ultrastructure,

physiology and behaviour of the sperm of more insect species than have all other

workers combined. He, too, has written extensively about the phylogenetic

significance of his findings, but not within a cladistic framework, and with little

regard for previous phylogenies reconstructed on the basis of comparative

morphology and life style.

Jamieson believes that information about spermatozoa can act as an

“independent arbiter” for resolving contentious problems of relationship. He

attempts to prove this by reviewing knowledge of hexapod sperm structure for

known members of each order and by examining congruence between his

cladograms, based on analysis of these character sequences, and Niels Kristensen’s

recent cladogram of hexapod relationships (1981 -Phylogeny of Insect Orders. Ann.

Rev. Ent. 26: 135-157) reconstructed on the basis of holomorphology (reproduced

as Eig. 4.1 on page 81).

He ‘sets the stage’ for his attempt in the first three chapters of the book which

provide comparative information about the sperm of other arthropods. In Chapter 1

(25 pp.), Jamieson defines the phylum Uniramia, summarizes the structural,

physiological, biochemical, embryological and fossil evidence for and against its

existence as a monophyletic taxon, and also for arthropod polyphyly vs. monophyly.

He concludes that the Uniramia does form a monophyletic lineage, albeit of

problematic position, but that the issue of arthropod polyphyly or monophyly cannot

be settled yet (however, the emphasis of his words about the latter topic will

convince any reader that he is an ardent polyphyleticist).

This chapter also contains brief, but rich and fully illustrated summaries of

chelicerate and crustacean sperm diversity (incredible) wherein he concludes that

the ground plan spermatozoan for both taxa is an “aquasperm” (an aquatic cell

generally associated with external fertilization and, like the ‘tadpole’ mentioned

Quaest. Ent., 1988,24 (2)

466

Book Review

Fig. 1. Phylogeny of the Uniramia as interpreted by Jamieson showing some aspects of sperm diversity.

Book Review

467

above, having a round nucleus, an apical acrosomal vesicle, a few large

mitochondria surrounding the base of the axoneme, the latter with a 9+2

arrangement of 9 doublet microtubules and 2 central singlets; and 2 centrioles

behind the head, of 9 triplet microtubules each, the distal one comprising the basal

body of the axoneme). However, such a sperm is unknown for any extant

crustacean, those of most species being aflagellate. He also emphasizes that sperm

ultrastructure of neither taxon provides evidence for close relationship with

Uniramia.

Sperm of onychophorans (Chapter 2 [13 pp.]- filiform, helically twisted cells; w

ith midpiece mitochondria separating the nucleus from the single centriole; a

flagellum containing a 9+2 axoneme + 9 peripheral accessory tubules and, external

to these, a persistent sheath [=manchettej of microtubules [i.e., n+9+9+2])- share

several characters considered by Jamieson to be synapotypic with those of

euclitellate annelids (Fig. 2.1). These, along with other, non spermatozoal,

characters, suggest to him that these taxa are sister groups descended from a

common, protoclitellate ancestor near the base of the Uniramia (Fig. 2.6, my Fig. 1).

This, of course, and as he mentions, would result in these worms being included

within the Uniramia; the remaining Annelida constituting a paraphyletic phylum.

At the end of this chapter, and seeming to this reviewer to have little to do with

spermatozoa, Jamieson considers the odd possibility that the trochophore larva is not

primitive to the annelids but was developed secondarily in the polychaete line as a

mechanism for dispersal (Fig. 2.6). This causes problems for the origin of molluscs

“as it would have to be postulated: ( 1 ), that molluscs developed a trochophore

independently of annelids; or (2), that molluscs arose from an ancestor, shared with

or ... referable to the polychaetes, which had a trochophore; or (3) that molluscs

developed by neoteny of a trochophore and have retained this in their life cycle. . .”

(p. 38). His Figure 2.6, which includes this possibility, will be difficult for many

invertebrate zoologists to swallow, since it shows the zany (at least to this reader)

phylogenetic relationship: [Chelicerata+[Crustacea+[Mollusca +[Polychaeta+

Uniramia]]]! I fear that many may not read his discussion which is intriguing, but

will accept his figure at face value-perhaps even using it in a course-parish the

thought!

Sperm ultrastructure in myriapods (Chapter 3-40 pp.) is characteristic for

members of each class (Fig. 3.1). Jamieson believes the myriapod ground plan for

sperm to resemble those of extant pauropods and chilopods and, to a lesser extent of

symphylans, with adaptation for increased length and motility in those of most

pauropods and chilopods and for immotility and a ribbon- or biscuit-shape in those

of extant diplopods. The ground plans for both myriapod and hexapod sperm are

synapotypic for a centriolar adjunct (a mass of material surrounding the remains of

the centriole at the junction between nucleus and axoneme and perhaps holding the

two together), a 9+2 axoneme with a tendency for addition of peripheral singlets,

peripheral axonemal material, and loss of the peripheral manchette of onychophoran

Quaest. Ent., 1988, 24 (2)

468

Book Review

sperm (Fig. 2.6).

The next 16 chapters summarize information about sperm ultrastructure and

function in investigated hexapods and indicate that Jamieson agrees with Knstensen

in including each of the entognath (Collembola, Protura, Diplura), apterygote groups

and Insecta as separate classes within a monophyletic superclass Hexapoda. His

chapters mostly treat what he believes to be monophyletic assemblages of orders

(Chapter 4-entognath apterygotes, 9 pp.; 5-ectognath apterygotes, 8 pp.; 6-summary

of pterygote sperm, 8 pp.; 7-Palaeoptera, 5 pp.; 8-Blattodea and Isoptera, 11 pp.;

9-Orthoptera and Phasmatodea, 18 pp.; 10-Embioptera, Dermaptera, Plecoptera and

Grylloblattodea, 6 pp.; 1 1-Psocoptera, Phthiraptera and Thysanoptera, 6 pp.;

12-Homoptera, 14 pp.; 13-Heteroptera, 9 pp.; 14-Holometabola-summary and

Neuroptera, 4 pp.; 15-Coleoptera, 19 pp.; 1 6-Mecoptera and Siphonaptera, 5 pp.;

17-Diptera, 38 pp.; 18-Amphiesmenoptera (Trichoptera and Lepidoptera), 24 pp.;

and 19-Hymenoptera, 6 pp.), many to the family level. References provided at the

end of each chapter, or at the end of some orders, vary in number from 6 (chapter 7)

to 132 (chapter 17).

From reading these chapters, one can see that knowledge of sperm ultrastructure

varies tremendously among higher taxa: from 0 species investigated (Zoraptera,

Mantodea, Megaloptera, Rhaphidioptera and Strepsiptera), through 1 (Phasmatodea,

Embioptera, Dennaptera, Plecoptera, Grylloblattodea and Thysanoptera), 2

(Symphyla, Pauropoda, Archaeognatha, Zygentoma, Psocoptera and Phthiraptera), 3

(Protura, Odonata and Siphonaptera), 4 (Collembola, Diplura, Ephemeroptera and

Mecoptera), 5 (Blattodea), 7 (Onychophora and Neuroptera), 10 (Hymenoptera), 1 1

(Chilopoda and Isoptera), 13 (Trichoptera), 18 (Heteroptera), 23 (Diplopoda), 27

(Lepidoptera), 33 (Orthoptera), 36 (Coleoptera), 57 (Homoptera), and 68 (Diptera).

In some taxa (Chilopoda, Diplopoda, Isoptera, Homoptera, Coleoptera, Diptera and

Lepidoptera), representatives have been selected from species occurring within most

lineages of the taxon; in others, from only one or two (Orthoptera and Heteroptera).

Such spotty coverage, as Jamieson states, makes phylogenetic reconstruction based

on analysis of spenn ultrastructure alone (he coins the term ‘spermiocladistics’ for

this enterprise) highly speculative. This is particularly so when one considers the

remarkable diversity in sperm structure known to occur within some well-studied

taxa such as Homoptera {e.g.„ some psylloids have binuclear sperm with spinose,

acrosomal appendages; some aleyrodids, immotile sperm with mitochondria and

axoneme vestigial; and coccoids, immotile or secondarily motile sperm with

acrosomes, mitochondria, centrioles and axonemes absent, the latter replaced with

circlets or spirals of 20 to >400 microtubules) and Diptera {e.g.„ Bibionidae-

axoneme 9-+-9+0 or 9+0; Mycetophilidae- only one mitochondrial derivative,

axoneme 9+2 or with variable number of outer singlets, rarely 9+9+3; Sciaridae-

many axonemal doublets; Cecidomyiidae- acrosome lost; mitochondrial derivatives

one or two, and cristate, crystalline or not; axonemes one or two of 9+0

microtubules, or doublets multiplied, some greatly, outer singlets lost but outer

Book Review

469

microtubules may be many; Simuliidae- axoneme 9+9+3; Culicidae- axoneme

9+9+1; Psychodidae- immotile, aflagellate, bifurcate sperm). Such diversity may,

eventually, be found to be true of other, now inadequately known, taxa even though

this seems not to be so of beetles.

In Chapter 20 (24 pp.), Jamieson integrates all this information together by first

postulating a detailed ground plan for the hexapod spermatozoan and then

summarizing evolution within member taxa of the Insecta. This ground plan gamete

he postulates to have the following characteristics (p. 282): elongate, filamentous

and motile; acrosome with bilayered or possibly trilayered perforatorium; nucleus

elongate with condensed chromatin; centriole with doublets only and surrounded by

a centriole adjunct; accessory bodies absent (2 elongate bodies typically flanking the

axoneme in the flagellum); two elongate, at least partly cristate, non-crystalline

mitochondrial derivatives; additional smaller mitochondria present in variable

numbers; axoneme of the 9+9+2 type, with regularly arranged peripheral singlets;

and peripheral ‘coarse fibre’ material present but ill defined.

In table 20.1, he lists the 57 derived character states he identified in the ground

plans of the spermatozoa of each higher group of hexapods and, before summarizing

the distribution of these on appropriate portions of Kristensen’s cladogram, provides

his own (Fig. 20.1) derived solely from consideration of these states. This tree bears

some resemblance to Kristensen’s tree but the relationships of many orders are

inadequately discriminated.

In the rest of this chapter he derives the spermatozoal ground plans for each

higher taxon treated in chapters 4 through 19, emphasizing in italics the character

states he believes to be apotypic for that taxon. These apotypies he then

superimposes on the appropriate portion of Kristensen’s cladogram (Figs. 20.

2-20.8), in some instances (entognath and ectognath Apterygota, Polyneoptera, and

Acercaria) supplying alternate hypotheses of relationship. These arguments are far

too complex to detail in this review but I include some differences in the attached

phylogenetic tree as Figure 1 .

The book is well produced on good quality paper, is relatively free of

typographical errors, and is profusely illustrated with fully labelled line drawings,

and transmission (and a few scanning) electron micrographs, partly from others

publications and partly from his own laboratory. Many of the figures are full page

plates comprising many separate drawings or photomicrographs so that the book

actually contains about 490 illustrations! The amount of information available about

sperm of some species is mind-boggling (c.g.,, the summary of sperm structure in

Drosophila spp. covers 1 1 pages and includes not only information about

ultrastructure, but also about function of the mitochondrial derivatives, axonemal

orientation during locomotion, developmental gradients in the axoneme, and genetic

control of spemiiogenesis).

My principal criticisms of the book are that Jamieson failed to provide: (1), a

glossary to aid non-specialist readers in coping with the profusion of special terms

Quaest. Ent., 1988,24 (2)

470

Book Review

for spermatozoa! organelles; and (2), a general discussion of factors possibly

influencing sperm structure and function, such as mode of transmission, duration of

survival in male and female genital tract or within spermatophore or spermatheca,

type of fluid they swim in and complexity of envelopes they must penetrate. The

latter details are known for sperm of few species, but a general summary of

possibilities such as that in Baccetti and Afzelius (1976, The Biology of the Sperm

Cell. Karger) would surely result in readers having these ideas in mind as they

plunge into the complex universe of spermatology.

Jamieson also missed a critical reference about thrips spermatozoa (Bode, W.

1983. Spermienstruktur und Spermatohistogenese bei Thrips validus Uzel [Insecta,

Thysanoptera]. Zool. Jb. Anat. 109: 301-318.) showing the males of this species to

produce sperm having an axoneme with 18 doublet intermixed with 9 singlet

microtubules that is fornied, during spermiogenesis, by development, alteration and

fusion of three, separate, 9-1-0 cilia present in early spermatids. In addition, the

Habilitationsschrift of this author (1986. Auswertung von Ultrastrukturmerkmalen

fiir die Phylogenetische Systematik der Thysanopteren. Fakultat fur Biologic,

Universitat Bielefeld-pp. 49-70) treats the ultrastructure of spermatozoa in 5 other

species of thrips in 4 families suggesting a ground plan axoneme for Thysanoptera

of 2x(9-t-2) or 2x(9-i-9-i-2) with those of the suborder Terebrantia 3x(9-i-0) and of

Tubulifera 2x(9-f2) or 2x(9-i-3).

The book is expensive but no more so than others its size now being published in

Great Britain. It will enable the cell biologist interested in the function of cell

organelles to select the best experimental material for his investigations and will

help these and other process-oriented biologists to understand and appreciate

evolutionary thinking-an understanding often seeming to be in short supply. It will

provide systematic entomologists, and other invertebrate zoologists and comparative

biologists entry to a valuable new source of taxonomic characters and will provide

guidance as to which taxa to examine. Finally, it should open a door for some

ecologists interested in sexual selection, since many seem to forget the little gametes

that make it all work.

B. S. Heming

Department of Entomology

University of Alberta

iVs

Quaestiones

Entomolosicae

A periodical record of entomological investigations,

published at the Department of Entomology,

University of Alberta, Edmonton, Canada.

VOLUME 24

NUMBER 3

SUMMER 1988

Publication of Quaestiones Entomologicae was started in 1965 as part of a

memorial project for Professor E.H. Strickland, the founder of the Department

of Entomology at the University of Alberta in 1922.

It is intended to provide prompt relatively low cost publication for compre-

hensive accounts of entomological research of greater than average length.

However, shorter papers about insects in the Prairie Provinces of Canada are

acceptable. Page charges are normally levied, the rate determined by printer’s

charges. Eor information about current page charges, consult the Editor.