The Genus Coptocarpus Chaudoir of

the Australian Region with Notes

on Related African Species

(Goleoptera: Carabidae: Oodini)

TERRY L. ERWIN

m

WUi.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 184
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 184

The Genus Coptocarpus Chaudoir of

the Australian Region with Notes

on Related African Species

(Coleoptera: Carabidae: Oodini)

Terry L. Erwin

SMITHSONIAN INSTITUTION PRESS

City of Washington

1974
ABSTRACT

Erwin, Terry L. The Genus Coptocarpus Chaudoir of the Australian Region

with Notes on Related African Species (Coleoptera: Carabidae: Oodini). Smithsonian Contributions to Zoology, number 184, 25 pages, 33 figures, 1 table,

1974.—The Australian-New Guinean genus Coptocarpus Chaudoir is revised

taxonomically. The 13 known species are presented in a key and then arrayed in

5 species groups. Four species, philipi, new species, chimbu, new species, yorkensis, new species, and grossus, new species, are newly described and illustrated. The

remaining nine species are redescribed and illustrated. Notes are presented on

probable phylogenetic relationships with the African genera Thryptocerus,

Orthocerodus, and Hoplolenus, and a preliminary zoogeographic scenario is

presented, based on these suspected relationships. The Indian genus Holcocoleus

is thought to be a possible key to the hypothesized relationships and is in need

of further study.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5164. SERIES COVER

DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Erwin, Terry L., 1940-

A Revision of the Australian-New Cuinean Genus Coptocarpus Chaudoir.

(Smithsonian contributions to zoology, no. 184)

Supt. of Docs, no.: SI 1.27: 184

1. Coptocarpus. 2. Carabidae. 3. Insects—Australia. 4. Insects—New Guinea. I. Title. II.

Smithsonian Institution. Smithsonian contributions to zoology, no. 184.

QL1.S54 no. 184 [QL596.C2] 591'.08s [595.7'62] 74-9923

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. - Price 85 cents (paper cover)
Contents

Page

Introduction 1

Methods 1

Terminology 2

Acknowledgments 2

Coptocarpus Chaudoir 2

Checklist of the Species of Coptocarpus Chaudoir 4

Key to the Species Groups and Species of Coptocarpus Chaudoir 5

Coptocarpus philipi, new species 5

Coptocarpus chimbu, new species 7

Coptocarpus yorkensis, new species • 7

Coptocarpus grossus, new species 8

Coptocarpus doddi Sloane 10

Coptocarpus nitidus Macleay . •. 11

Coptocarpus chaudoiri Macleay 12

Coptocarpus gibbus Chaudoir 12

Coptocarpus fuscitarsis (Blanchard) . 12

Coptocarpus championensis Chaudoir 13

Coptocarpus impar Sloane 14

Coptocarpus thoracicus (Castelnau) 15

Coptocarpus australis (Dejean) 16

Phylogenetic and Zoogeographic Notes . . 20

Literature Cited 25

1U

The Genus Coptocarpus Chaudoir of

the Australian Region with Notes

on Related African Species

(Coleoptera: Carabidae: Oodini)

Terry L. Erwin

Introduction

The Oodini, as a tribe, have never been systematically analyzed using modern techniques, although certain genera or generic groups were

recently treated taxonomically (Basilewsky, 1943,

1946, 1949; Darlington, 1968; Jeannel, 1949; Lindroth, 1956). Older papers (e.g., Blanchard, 1853;

Castelnau, 1868; Chaudoir, 1857, 1882; LaFerte,

1851; Macleay, 1873; Sloane, 1910, 1915) were regional compilations of species or "classical" taxonomic studies which did not assemble observed

morphological or natural history data into meaningful and predictive classification schemes; distribution, of course, was hardly mentioned, never

analyzed.

This study of the oodines, especially Coptocarpus, began when P. J. Darlington, Jr., requested

that I describe two of his new species of Coptocarpus from the Australian Region. Since I am

opposed in principle to single species descriptions

(Ball and Erwin, 1969; Erwin and Ball, 1972), I

undertook a basic revision of the Australian

Coptocarpus. This led to a preliminary morphological analysis of the oodines in general and to

the discovery of the apparent close relationship of

certain African species with Coptocarpus species.

Terry L. Erwin, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Thus, the purpose of this presentation is to taxonomically revise the species of Coptocarpus, to

analyze Coptocarpus species morphology in relation

to certain other oodine groups, and to provide

notes on the African-Australian relationships between Coptocarpus and the so-called "Thryptocerini" (Jeannel, 1949).

The only practical revision of Coptocarpus, that

of Sloane (1910), covers seven of the now known

13 species, provides a key for identification of specimens, and provides terse specimen descriptions

with little regard paid to species variation. In this

paper by Sloane, species described as Coptocarpus

riverinae Macleay (1873:329) and C. planipennis

Macleay (1878:216) were reassigned to genus Oodes

(Sloane, 1910:446). I have studied the types of

these two species (CSIRO) and agree with Sloane's

action. However, it must be pointed out that

Oodes, as presently conceived, is a "catch-all" genus

and must be studied from a worldwide viewpoint,

especially in regard to the placement of the Australian species. The same may be said for genus

Anatrichis, which is now composed of at least two

monophyletic groups that are not very closely

related.

METHODS.—This study is the result of the examination of 117 specimens of Coptocarpus and hundreds of specimens of other Oodini.

Methods of dissection and procedure are the

same as those used by me in the past (Erwin, 1970);
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

methods of genitalic illustration are, in part, those

I adopted from C. H. Lindroth (Erwin, 1972b,

1973, 1974). Note that the short line accompanying

the illustrations equals 1.0 mm unless otherwise

noted. My criteria for recognizing taxa at various

levels are the same as I outlined before (Erwin,

1970).

The abbreviations given under "Acknowledgments" indicate the museums in which studied

specimens are now housed.

I have seen all type-specimens unless otherwise

noted and all lectotypes have been so labeled by me.

TERMINOLOGY.—In my 1974 paper on Pericompsus, I introduced a new term for the longitudinal

elytral structures between the elytral intervals. That

word is "interneur" and I refer the reader to Erwin

(1974).

ACKNOWLEDGMENTS.—I wish to thank P. J. Darlington, Jr., for providing me the incentive and

specimens which launched this study. He has also

allowed me to place in the country of their origin

the holotypes of new species described herein, a

philosophy of which we both are strong advocates

(see also Darlington, 1971; Erwin, 1972a) when

there are suitable museum facilities in those countries. Darlington's specimens, which were not deposited in other countries, are either in the Museum

of Comparative Zoology, Cambridge, Massachusetts

(MCZ) or in the National Museum of Natural

History, Smithsonian Institution, Washington,

D.C. and are listed under the catalog numbers of

the old United States National Museum (USNM).

I also thank the following people for loan of

specimens, especially types, in their charge: Mme.

A. Bons, Museum National d'Histoire Naturelle,

Paris (MHNP), and Drs. E. B. Britton and B. P.

Moore, Division of Entomology, Commonwealth

Scientific and Industrial Research Organization

(CSIRO), Canberra City, Australia.

I heartily thank W. Donald Duckworth and

Donald R. Whitehead for reading and criticizing

the draft manuscript and Walt Brown for the excellent quality scanning electron micrographs of

Coptocarpus member's tarsal articles.

Coptocarpus Chaudoir

Coptocarpus Chaudoir, 1857:57. [Type-species: Oodes australis

Dejean, 1831:671, by original designation and monotypy.]

Oodes of authors.

DIAGNOSIS.—Coptocarpus species of Australia and

New Guinea may be recognized from other Oodini

in the Australian Region (Anatrichis, Oodes) by

the absence of ambulatory setae on the abdominal

sterna. Furthermore, Anatrichis have only two

male probasitarsomeres with squamate setae beneath, while Oodes members have three dilated,

but the third article of Oodes has squamate setae

on fully one-half of the article, whereas Coptocarpus members have only one-third clothed with

these setae.

Among the Oodini, only Coptocarpus, Thryptocerus, Orthocerodus, Hoplolenus, and Holcocoleus

members have the male anterior tarsi asymmetrically arranged (Figures 31, 32). Members of

Holcocoleus have the least asymmetric tarsal develFICURE 1—Scutellar region of elytra: A, Coptocarpus thoracicus, Margaret River, Australia; B, C. australis, Melbourne,

Australia.
NUMBER 184

opment. Of these five genera, only members of Holcocoleus possess ambulatory setae on the abdominal

sterna, a feature also common to all other Oodini

members. Members of all of these genera have an

enlarged scutellum in relation to other Oodini.

DESCRIPTION.—Form (Figures 19-30) broad and

convex, the prothorax quite large in relation to

the elytra as compared to many other ground

beetles. Color: Black to rufopiceous, some members quite iridescent beneath, appendages generally

paler than color of dorsum. Head: Size various in

proportion to pronotal width; labrum with six

setigerous pores arrayed across anterior sixth, inner

four slightly closer to each other than to lateral

ones; clypeus with one setigerous pore at each

anterior angle or not; mandibles smooth dorsally;

penultimate article of labial palp without setae,

other articles of labial palps and maxillary palps

also without setae or pubescence; mentum with

anterior edge emarginate and unidentate, tooth

broad and apically rounded or truncate, margin of

mentum with raised bead extended from right to

left lobe; submentum bisetose, setigerous pores at

posterior lateral angles; antennal pubescence begins at basal sixth of article 4; one supraorbital

seta per eye or none. Prothorax: Broad and convex, nearly as wide or wider than elytra; lateral

setae absent; proepisterna and prosternum laterally sparsely punctate or not; male protarsal articles 1-3 (Figure 2) asymmetrically dilated, 2 and 3

fully clothed beneath with squamate setae which

are transversely arranged, article 1 with only apical

third clothed with these setae, article 4 very small

and slightly bilobed, no articles with natatorial

setae; protibial spur long and stout, subequal in

length to length of probasitarsus or longer. Mesothorax: Scutellum large (Figure 1); scutellar interneur (Figure 1) present or not, when present striate,

punctate, or striatopunctate; interneurs 1-7 present

externally or not, when present, striate or striatopunctate; apex of elytra moderately to strongly

carinatesulcate, sulcus multituberculate; medial

groove of mesosternum deeply sulcate; mesepisternum punctate or not; mesotarsus without natatorial

setae. Metathorax: Flight wings present or not,

when absent, metasterna shortened longitudinally;

metepisternum (Figure 3) punctate or not; metatarsus without natatorial setae. Abdomen: Sterna

I—III rugosely punctate, sternum IV punctate or

not; ambulatory setigerous pores of sterna II-V

FIGURE 2.—Male anterior tarsomeres: A, Coptocarpus australis, Melbourne, Australia;

B, C. grossus, Margaret River, Australia.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

absent; setigerous pores of sternum VI, 4 in female,

2 in male. Genitalia: Male (Figures 4, 6-15) with

broadly lobed parameres, the left larger than the

right; median lobe with distal orifice symmetrically

placed on dorsal surface near apical third; internal

sac with numerous spines, plates, and/or microtricheal fields; female stylus a short, arcuate blade

with one stout and rather long medial spine and

one or two shorter lateral spines located basally,

and with a small setigerous pore located lateroventrally at apical third. Microsculpture: Dorsally

with very fine isodiametric reticulation interspersed with micropunctulae, ventrally either the

same as dorsally or with longitudinally stretched

reticulations which produce iridescent coloration.

Size: Length, 7.37 to 15.06 mm; width, 2.83 to 6.39

mm. Secondary sexual characteristics: Male with

strongly dilated anterior tarsal articles 1-3, each

with squamate setae beneath, and sternum VI with

two setigerous pores. Female without dilated anterior tarsi or squamate setae and sternum VI with

four setigerous pores. Pupae may be sexed by observing the anterior tarsi for dilated articles (Ball

and Erwin, 1969).

DISTRIBUTION (Figures 16-18).—The genus as

here defined has species with a combined range

extending from New Guinea at 5°S latitude to

southern Australia at 38°S latitude and which are

apparently confined to those two land masses.

Checklist of the Species of Coptocarpus Chaudoir

The philipi group

1. C. philipi, new species

The chimbu group*

2. C. chimbu, new species

3. C. yorkensis, new species

The grossus group

4. C. grossus, new species

The chaudoiri group

5. C. doddi Sloanc, 1910:450

6. C. nitidus Macleay, 1873:330

7. C. chaudoiri Macleay, 1873:329

The australis group

8. C. gibbus Chaudoir, 1882:511

9. C. fuscitarsis (Blanchard), 1853:39

10. C. championensis Chaudoir, 1882:510

11. C. impar Sloane, 1910:451

12. C. thoracicus (Castelnau), 1868:151

13. C. australis (Dejean), 1831:671

FIGURE 3.—Metastcrnum, metepisternum, and base of abdominal sterna: A, Coptocarpus thoracicus, Margaret River, Australia; B, C. australis, Melbourne, Australia.
NUMBER 184

Key to the Species Groups and Species of Coptocarpus Chaudoir

1. Elytron with well-impressed striate interneurs, easily visible to the unaided eye 2

Elytron smooth, if striate interneurs present then only discernible at high magnification ..

(chimbu group) 12

2(1). Mctepisternum with large coarse punctures (Figure 3) at least in apical half; mesepisternum usually also coarsely punctate (Figure 3) 3

Metepisternum and mespisternum without punctures, surface (at least disc of mesepiSternum) smooth and shiny 9

3(2). Sutural interneur (Figure 1A) continuous from elytral base adjacent to umbilicate setigerous pore to apex; form narrow and elongate (australis group, in part) 4

Sutural-scutellar interneur divided, thus not continuous (Figure 1B) adjacent to umbilicate setigerous pore; form large and robust 5

4(3). Size large, length more than 10.0 mm; male genitalia (Figure 13) 11. C. impar Sloane

Size small (Figure 29), length less than 9.5 mm; male genitalia (Figure 14)

12. C. thoracicus (Castelnau)

5(3). Interneurs striate and finely punctate, punctures easily visible at X 50; male probasitarsus

transverse (Figure 2A), its length subequal to or less than that of article 2 (not seen

in C. gibbus) (australis group, in part) 6

Interneurs striate, smooth, without fine punctures; male probasitarsus elongate (Figure

2B), longer than article 2 (grossus group) 4. C. grossus, new species

6(5). Pronotum with lateral bead narrow throughout, much narrower than apical antennal

articles (viewed from dorsal surfaces) 7

Pronotum with lateral bead wide (Figure 26), especially in basal half, fully as wide as

dorsal surfaces of apical antennal articles 8. C. gibbus Chaudoir

7(6). Mesepisternum rugose and coarsely punctate, punctures widespread; pronotum with

lateral bead evenly narrow throughout 8

Mesepisternum smooth, without rugosities or punctures; pronotum with lateral bead

wide at middle, narrower toward base and apex 9. C. fuscitarsis (Blanchard)

8(7). Mesepisternum with numerous fine and coarse punctures (Figure 3B), punctures easily

visible among the finely engraved microsculpture; male probasitarsus narrow at apex,

less than 0.75 as wide as article 2 (Figure 2A) 13. C. australis (Dejean)

Mesepisternum not or very shallowly and sparsely punctate, but finely rugose; male

probasitarsus wide at apex, fully as wide as article 2 10. C. championensis Chaudoir

9(2). Prosternum laterally with numerous coarse punctures; head rather large (Figure 23-25)

(chaudoiri group) 10

Prosternum laterally without punctures, surface smooth; head very small and deeply

silicate transversely behind eyes) Figure 19) . . (philipi group) 1. C. philipi, new species

10(9). Pronotum with lateral bead continuous throughout, well developed to base 11

Pronotum with lateral bead effaced in basal half 7. C. chaudoiri Macleay

11(10.) Elytral intervals slightly convex, interneurs striate and coarsely punctate, deeply impressed; length 9.23 mm 5. C. doddi Sloane

Elytral intervals flat, interneurs striate and finely punctate, shallowly impressed; length,

10.20 to 11.66 mm 6. C. nitidus Macleay

12(2). Abdominal sterna II coarsely punctate laterad of metacoxa; pronotum (Figure 20) very

broad, wider than elytra across humeri 2. C. chimbu, new species

Abdominal sternum II rugose but not punctate; pronotum (Figure 21) long and narrow,

subequal to elytral width 3. C. yorkensis, new species

1. Coptocarpus philipi, new species TYPE-LOCALITY.—The Crater, near Atherton Pla-

_ . _ ,D ,„ teau, Queensland, Australia.

FIGURES 4, 5A, 18, 19 ' ^ _ , , , ,

DESCRIPTION (Figure 19).—Form broad and deTYPE-SPECIMENS.—The holotype male is in pressed, with a very small head about one-third

CSIRO; it was collected by P. J. Darlington, Jr., in the width of the pronotum. Easily recognized by

1957. Three para types are listed below. the small, transversely sulcate head and the pecu-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

liar form of the mesbasitarsus (Figure 5A). Color:

Black dorsally, black iridescent ventrally; appendages rufopiceous, antennae and palpi testaceous.

Head: Narrow and small, with transverse sulcus

posterior to eyes, width across eyes subequal to

one-third width of pronotum; labrum with two

long setae, one at each anterior angle and with

four shorter setae closely spaced at middle of anterior edge; mandible with shorter scrobe, dorsal

surface broader, otherwise as in C. australis. Prothorax: Pronotum broad and depressed, wider than

elytra, lateral margin narrowly beaded, with bead

extended from hind angle around anterior angle

nearly to middle, otherwise as in C. australis; venter

without punctation, otherwise as in C. australis.

Mesothorax: As in C. australis except elytral interneurs shallowly striate-punctate, scutellar interneur nearly effaced, intervals flat, humeral tooth

well developed and prominent; mesepisternum

without punctures; mesobasitarsus elongate and

bent (Figure 5A) with serrate carina beneath. Metathorax: Flight wings absent; metepisternum without punctures, shape quadrate; metabasitarsus

elongate. Abdomen: Sterna not punctate, otherwise

as in C. australis. Genitalia: Male (Figure 4).

Microsculpture: As in C. australis. Size: Length,

9.47 to 10.12 mm; width, 4.13 to 4.45 mm.

VARIATION.—The four specimens studied are

quite homogeneous.

NATURAL HISTORY.—Darlington (1960) indicates

that his specimens were collected in rain forest

beneath stones or logs; one was at waterside, the

others were not. Specimens were collected in December and February. None were teneral.

ETYMOLOGY.—The genitive patronym, philipi,

honors my friend and mentor who collected the

types of this species, Philip J. Darlington, Jr.

LOCALITY RECORDS (Figure 18).—I have seen

four specimens from the following localities: AUSFIGURE 4.—Male genitalia with everted endophallus, left

lateral profile, and apex, ventral aspect of Coptocarpus

philipi, holotype from the Crater, Queensland, Australia.

FIGURE 5.—Mesobasitarsus, distal lateral aspect; A, Coptocarpus

philipi, Lake Eacham, Queensland, Australia; B, C. australis,

40 miles ENE of Glen Innes, New South Wales, Australia.
NUMBER 184

TRALIA: QUEENSLAND: Lake Eacham near Atherton Plateau (MCZ), Longlands Gap near Atherton

Plateau (MCZ), "Mts. above Atherton" (USNM),

The Crater near Atherton Plateau (CSIRO).

2. Coptocarpus chimbu, new species

FIGURES 6, 16, 20

TYPE-SPECIMENS.—The holotype male is in

CSIRO; it was collected by F. Parker in 1967.

TYPE-LOCALITY.—Dumun, Northeast New Guinea (ca. 145°W 6°S).

DESCRIPTION (Figure 20).—Form broad and convex with a wide and nearly quadrate pronotum

and a moderately large head that is subequal

across eyes to half the width of the pronotum.

Easily recognized by the absence of striatepunctate interneurs and by the punctate abdominal

sternum II. Color: Nigropiceous throughout; appendages rufopiceous. Head: As in C. australis

except clypeus and vertex without setae. Prothorax: As in C. australis except pronotum wider than

elytra, venter without punctures. Mesothorax:

Elytra smooth, interneurs effaced externally, dorsal

setigerous pores of interval 3 absent, humeral angle

dentate, tooth prominent, otherwise as in C. australis; mesepisternum without punctures; mesobasitarsi elongate, not serrate-carinate beneath as

in C. philipi. Metathorax: Flight wings absent;

metepisternum not punctate, shape square; otherwise as in C. australis. Abdomen: Sternum II

coarsely punctate, other sterna smooth or slightly

rugose; otherwise as in C. australis. Genitalia: Male

(Figure 6); female unknown. Microsculpture: As

in C. australis. Size: Length, 8.58 mm; width, 3.72

mm.

ETYMOLOGY.—The noun, chimbu, refers to the

name of the river in New Guinea from which area

the type was collected.

LOCALITY RECORDS (Figure 16).—I have seen only

the holotype from Dumun, New Guinea (CSIRO).

3. Coptocarpus yorkensis, new species

FIGURES 7, 18, 21

TYPE-SPECIMENS.—The holotype male is in

CSIRO; it was collected by P. J. Darlington, Jr., in

1958. One para type is listed below.

FIGURE 6.—Male genitalia with everted endophallus, left lateral profile, and apex, ventral aspect

of Coptocarpus chimbu, Dumun, New Guinea.
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPE-LOCALITY.—Vicinity of Tozers Gap on Cape

York, Queensland, Australia.

DESCRIPTION (Figure 21).—Form narrow and

subconvex with a moderately small head. Most

easily recognized by the nearly smooth elytra and

the lack of punctures on abdominal sternum II.

Color: Rufopiceous throughout; legs more rufous;

male anterior tarsi, antennae, and palpi testaceous.

Head: As in C. australis except relatively much

narrower and with small, nearly flat eyes. Prothorax: Pronotum as in C. australis except narrower

and less convex; venter as in C. australis except

punctures absent. Mesothorax: Elytral interneurs

very faintly impressed, striate; scutellar interneur

almost effaced; intervals flat, interval 3 with two

faintly impressed setigerous pores posterior of

middle; otherwise elytra as in C. australis; mesepisternum without punctures, surface slightly rugose;

middle tarsal articles 2 to 4 with moderately long

golden setae ventrally, one row each side. Metathorax: Flight wings absent; sterna without punctures;

tarsal article 4 only, with 2 rows of moderately

long golden setae, otherwise with bilateral rows of

spinose setae ventrally. Abdomen: Sterna without

punctures, otherwise as in C. australis. Genitalia:

Male (Figure 7). Microsculpture: As in C. australis.

Size: Length, 8.67 to 9.87 mm; width, 3.56 to 4.05

mm.

NATURAL HISTORY.—P. J. Darlington, Jr., and

family collected the two known specimens in

savannah woodland or rain forest beneath logs or

stones (Darlington, 1960). Both specimens were

collected in January and neither was teneral.

ETYMOLOGY.—The adjective, yorkensis, refers to

Cape York, the general area where these beetles

live.

LOCALITY RECORDS (Figure 18).—I have seen two

specimens, both from Cape York, Queensland; one

from near Tozers Gap (CSIRO) and the other

from the Iron Range (MCZ).

4. Coptocarpus grossus, new species

FIGURES 2B, 8, 17, 22

TYPE-SPECIMENS.—The holotype male and allotype are in CSIRO; both were collected by P. J.

Darlington, Jr., in 1931. Ten paratypes are listed

below.

FIGURE 7.—Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, right lateral aspect; and apex, ventral aspect of Coptocarpus yorkensis, vicinity of Tozer

Gap, Queensland, Australia.
NUMBER 184

TYPE-LOCALITY.—Margaret River, Western Australia.

DESCRIPTION (Figure 22).—Form very board and

convex with a transverse pronotum, which is somewhat narrower than the inflated elytra, and with a

very large head, which is wider across the eyes than

half the width of the pronotum. Easily recognized

by the smooth striate interneurs and large size.

Color: Piceous to nigropiceous throughout; antennae, palpi, and tarsi rufopiceous. Head: Very

broad across eyes, wider than half the width of

pronotum; mental tooth broadly truncate, otherwise head as in C. australis. Prothorax: As in C.

australis except probasitarsus of male (Figure 2B).

Mesothorax: Elytral interneurs striate, striae well

impressed, numbers 1, 4-7 not quite extended to

basal margin; scutellar interneur a row of unconnected punctures removed from interneur 1; intervals nearly flat, interval 3 with two small setigerous

pores posterior of middle; humeral angle with welldeveloped, acute tooth that protrudes beyond margin; otherwise as in C. australis. Metathorax: Flight

wings absent; metepisternum coarsely yet sparsely

punctate; otherwise as in C. australis. Abdomen:

Sterna I to III rugosely punctate laterally; otherwise as in C. australis. Genitalia: Male (Figure 8).

Microsculpture: Head, pronotum, and elytra with

very small isodiametric reticulation throughout;

FIGURE 8 Male genitalia with everted endophallus, left lateral profile, and apex, ventral aspect of Coptocarpus grossus, Margaret

River, Australia.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

venter with more longitudinally stretched reticulation, slightly iridescent. Size: Length, 11.34 to 15.06

mm; width, 4.61 to 6.39 mm.

VARIATION.—As in C. australis the size variation

is remarkable even within a single population sample. However, all character states studied are constant and the male genitalia with all of its intricate

internal details are constant.

NATURAL HISTORY.—P. J. Darlington, Jr., collected one example near the Margaret River in

October and several others at the same place in

November. Other specimens seen were collected in

October and February. None were teneral.

ETYMOLOGY.—The Latin adjective, grossus, meaning "big," in reference to the broadness of these

beetles.

LOCALITY RECORDS (Figure 17).—I have seen 12

specimens from the following localities: AUSTRALIA: WESTERN AUSTRALIA: Margaret River (CSIRO,

MCZ, MHNP, USNM), Manjimup (MCZ), 30 miles

SW Nannup (CSIRO).

5. Coptocarpus doddi Sloane

FIGURES 9, 18, 23

Coptocarpus doddi Sloane, 1910:450. [Holotype, a male, in

CSIRO. Type-locality: Port Darwin, Northern Territory,

Australia.]

DESCRIPTION (Figure 23).—Form broad and convex with a transverse pronotum subequal in width

to width of elytra, and with a moderate-sized head

which, across the eyes, is slightly smaller than half

the width of the pronotum. Easily recognized by

the slightly convex elytral intervals, continuous

lateral pronotal bead, punctate prosternum laterally, and smooth met- and mesepisterna. Color:

Piceous to rufopiceous throughout; appendages

slightly paler. Head: As in C. australis except vertex transversely shallowly sulcate. Prothorax: As

in C. australis except probasitarsus of male shorter

than apical tibial spur. Mesothorax: As in C. australis except punctures of interneurs more coarse,

scutellar interheur absent, and mesepisternum

FIGURE 9.—Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook; left oblique aspect; and apex, ventral aspect of Coptocarpus doddi, holotype from Port

Darwin, Australia.
NUMBER 184 11

without punctures. Metathorax: Flight wings absent, metepisternum without punctures, otherwise

as in C. australis. Abdomen: Sterna I to III only

punctate laterally, not rugose, otherwise as in C.

australis. Genitalia: Male (Figure 9); female unknown. Microsculpture: As in C. australis, venter

with more longitudinally stretched reticulation.

Size: Length, 9.23 mm; width 4.05 mm.

LOCALITY RECORDS (Figure 18).—I have seen only

the holotype from Port Darwin, N.T., Australia

(CSIRO).

6. Coptocarpus nitidus Macleay

FIGURES 10, 18, 24

Coptocarpus nitidus Macleay, 1873:330. [Type lost according

to B. P. Moore, in litt. Type-locality: Cape York, Queensland, Australia.]

Coptocarpus oviformis Chaudoir, 1882:511.—Sloane, 1910:

451. [Lectotype, a female, herein selected in MHNP. Typelocality: Cape York, Queensland, Australia.]

DESCRIPTION (Figure 24).—Form broad and moderately convex with a transverse pronotum nearly

as wide as the elytra, and with a moderately large

head, which is slightly narrower than half the

width of the pronotum. Recognized best by the

nonpunctate venter, complete pronotal lateral

bead, well-impressed striate-punctate interneurs,

and flat intervals. Color: As in C. australis. Head:

As in C. australis except smaller relative to rest of

body and with four medial labral setae located

closer to center of labrum. Prothorax: As in C.

australis except male probasitarsus more elongate.

Mesothorax: As in C. australis except interneurs

less impressed, intervals flatter, venter not punctate. Metathorax: Flight wings absent, venter not

punctate, otherwise as in C. australis. Abdomen:

As in C. australis except sterna I and II less punctate and less rugose. Genitalia: Male (Figure 10).

Microsculpture: As in C. australis. Size: Length,

10.20 to 11.66 mm; width, 4.05 to 4.94 mm.

VARIATION.—The small sample is quite homogeneous.

FIGURE 10 Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, left oblique aspect; and apex, ventral aspect of Coptocarpus nitidus, Bamaga, Queensland,

Australia.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NATURAL HISTORY.—P. J. Darlington, Jr., and

family collected these beetles in light rain forest

or savannah woodland in two localities. I have

seen specimens collected only in January; none

were teneral.

LOCALITY RECORDS (Figure 18).—I have seen 10

specimens from the following localities: AUSTRALIA: QUEENSLAND: "Cape York" (MHNP), Bamaga (CSIRO, MCZ, USNM), Lockerbie (MCZ).

NOTES.—Although the type is apparently lost we

have Sloane's (1910) statement that he examined

the "one type-specimen (female)" in the Macleay

Museum. He considers C. nitidus synonymous with

C. oviformis (of which the type is in MHNP) and

provides additional character states to add to

Macleay's terse description (1873). Based on these

observations of both Macleay and Sloane, I am

confident that Darlington's specimens are of this

species.

7. Coptocarpus chaudoiri Macleay

FIGURES 18, 25

Coptocarpus chaudoiri Macleay, 1873:329. [Lectotype, a female, here selected in CSIRO. Type-locality: Clarence

River, New South Wales, Australia.]

DESCRIPTION (Figure 25).—Form very broad and

moderately convex with a transverse pronotum

nearly as wide as the elytra, and with a large head

which, across the eyes, is slightly narrower than

half the width of the pronotum. Easily recognized

by the nonpunctate postventer (except abdominal

sternum I) and the effaced lateral bead along the

basal half of the pronotum. Color: As in C. australis. Head: As in C. australis. Prothorax: As in

C. australis except punctation of posternum coarser; lateral bead of pronotum effaced in basal third;

male tarsi unknown. Mesothorax: As in C. australis

except intervals less convex medially and flat laterally, interneurs less impressed and venter not

punctate. Metathorax: Flight wings absent, venter

not punctate. Abdomen: Sternum I rugosely punctate, otherwise as in C. australis. Genitalia: Male

unknown; female not dissected. Microsculpture: As

in C. australis. Size: Length, 12.60 mm; width,

5.58 mm.

LOCALITY RECORDS (Figure 18).—I have seen only

the lectotype from the Clarence River, New South

Wales, Australia (CSIRO).

8. Coptocarpus gibbus Chaudoir

FIGURES 17, 26

Coptocarpus gibbus Chaudoir, 1882:511. [Lectotype, a female,

here selected in MHNP. Type-locality: "Australie" as

originally given by Chaudoir.]

DESCRIPTION (Figure 26).—Form broad and convex with a very large head which is wider than half

the width of the pronotum. Easily recognized by

the very wide pronotal bead, which is wide throughout its length. Color: Nigropiceous throughout;

appendages rufopiceous. Head: As in C. australis

except head much broader in relation to width of

pronotum. Prothorax: As in C. australis except

lateral bead wider throughout its length. Male

anterior tarsi not seen. Mesothorax and Metathorax: As in C. australis except flight wings absent;

metepisternum shorter, almost quadrate. Abdomen:

As in C. australis except sternum III with fewer

punctures and more longitudinal rugosities. Genitalia: Male unknown; female as in C. australis.

Microsculpture: As in C. australis. Size: Length,

11.74 to 11.90 mm; width, 4.69 to 4.86 mm.

LOCALITY RECORDS (Figure 17).—I have seen two

specimens, one not labeled (MHNP) and one from

Queensland, Australia (MHNP).

NOTES.—Sloane (1910) recorded specimens from

western Australia at Albany, Bunbury, and Mount

Barker, but I have not seen these specimens. One

specimen in MHNP is labeled "Queensland" and

it matches the unlabeled type. Sloane's records

must be regarded as based on some other species

with which he confused C. gibbus. It is doubtful

that C. gibbus occurs both in Queensland and

southernmost Western Australia.

9. Coptocarpus fuscitarsis (Blanchard)

FIGURES 11, 18, 27

Oodes fuscitarsis Blanchard, 1853:39. [Lectotype, a male, here

selected in MHNP. Type-locality: Rafles Bay, Northern

Territory, Australia.]

DESCRIPTION (Figure 27).—Form broad and convex with pronotum wider than widest part of

elytra. Easily recognized by the smooth mesepisternum and the wide pronotal bead medially,

which narrows anteriorly and posteriorly. Color:

Nigropiceous, appendages rufopiceous except male

anterior tarsi rufous, and antennae and palpi testa-
NUMBER 184 13

FIGURE 11 Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, left oblique aspect; and apex, ventral aspect of Coptocarpus fuscitarsis, lectotype from

Rafles Bay, Australia.

ceous. Head: As in C. anstralis except narrower

than half the width of the pronotum and eyes more

depressed posterodorsally. Prothorax: Pronotum

broad and convex, slightly wider than elytra at

widest point, lateral margin with bead wide at

middle, tapering posteriorly and anteriorly; posterolateral depression very shallow; otherwise as in

C. australis. Mesothorax: As in C. australis except

mesepisternum smooth, without punctures. Metathorax: As in C. australis except flight wings absent.

Abdomen: As in C. australis except only sternum

I and II with fine punctures, sternum III finely

rugose. Genitalia: Male (Figure 11); female as in

C. australis. Microsculpture: As in C. australis.

Size: Length, 10.04 to 11.34 mm; width, 4.61 to

4.69 mm.

LOCALITY RECORDS (Figure 18).—I have seen 2

specimens from Rafles Bay, Northern Territory,

Australia (MHNP).

10. Coptocarpus championensis Chaudoir

FIGURES 12, 17, 28

Coptocarpus championensis Chaudoir, 1882:510. [Lectotype,

a male, here selected in MHNP. Type-locality: "Australie"

as originally given by Chaudoir, herewith restricted to

Eradu, near Champion Bay, Western Australia on the

basis of Sloan's (1910) records, and on specimens listed

below.]

DESCRIPTION (Figure 28).—Form short, broad,

and convex with strongly depressed posterolateral

sides of the pronotum. Easily distinguished from

the similar C. australis members by the rugose,

rather than punctate, mesepisterna and by the

nearly smooth striate interneurs. Color: Rufopiceous to nigropiceous; head and pronotum in

some specimens rufous and elytra nigropiceous, in

others concolorous; appendages rufous or piceous

except tarsi, antennae, and palpi testaceous or rufotestaceous. Head and Prothorax: As in C. australis

except posterolateral sides of pronotum more depressed, anterolateral sides of prosternum less punctate, and probasitarsus of male subequal in width

to article 2. Mesothorax: As in C. australis except

interneurs 3 to 7 with extremely small punctures

hardly visible at X 50, scutellar interneur striatepunctate, intervals barely convex, and mesepisternum finely rugose but not punctate. Metathorax:

As in C. australis except metepisternum with smaller and more rugose punctation. Abdomen: As in
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 12. Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, left oblique aspect; and apex, ventral aspect of Coptocarpus championensis, Geraldton,

Australia.

C. anstralis. Genitalia: Male (Figure 12); female

as in C. anstralis. Microsculpture: As in C. australis.

Size: Length, 9.39 to 11.74 mm; width, 3.96 to

4.86 mm.

VARIATION.—See description of color above;

otherwise the samples available are quite homogeneous.

NATURAL HISTORY.—I have seen specimens collected in April, October, and December. Some specimens collected in October were quite teneral, the

rest subteneral when collected. The April and

December specimens, one collected each month,

are fully pigmented and hard. It is likely that immatures could be found in September.

LOCALITY RECORDS (Figure 17).—I have seen 24

specimens from the following localities: AUSTRALIA: (MHNP); WESTERN AUSTRALIA: Cranbrook

(MCZ), Geraldton (CSIRO, MCZ, USNM), "Mt.

Casino ? v. Perth" (MCZ), Rockingham (MCZ),

Swan River (MHNP).

NOTE.—Sloane (1910) records specimens from

Eradu, near Champion Bay.

11. Coptocarpus impar Sloane

FIGURES 13, 17

Coptocarpus impar Sloane, 1910:451. [Holotype, a headless

male, in CSIRO. Type-locality: Roebuck Bay, Western

Australia.]

DESCRIPTION.—Form similar to C. thoracicus but

more elongate and proportionately larger. Easily
NUMBER 184 15

\

FIGURE 13.—Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, left oblique aspect; and apex, ventral aspect of Coptocarpus impar, holotype, Roebuck

Bay, Australia.

distinguished from other members of the genus by

the absence of a scutellar interneur and the elongate, narrow form. Color: Piceous; appendages

paler with rufotestaceous anterior tarsi (male).

Head: Missing from type, the only specimen I have

seen. Prothorax: As in C. australis except prosternum anterolaterally with fewer but coarser punctures. Mesothorax: As in C. australis except elytra

without scutellar interneur and mesepisternum

coarsely punctate. Metathorax: As in C. australis

except flight wings absent. Abdomen: Sterna I to

III with scattered coarse punctures, otherwise as in

C. australis (male only, female not seen). Genitalia:

Male (Figure 13). Microsculpture: Pronotum with

very small isodiametric reticulations, surface rather

dull; elytral reticulation larger, surface shiny. Size:

Length (without head), 8.58 mm; width, 3.88 mm.

Sloane (1910) gives the length of the type with the

head as 10.0 mm and the width as 3.80 mm. The

small discrepancy in width size is probably due to

refinement of measuring devices since 1910.

LOCALITY RECORD (Figure 17).—I have seen only

the type-specimen from Roebuck Bay, Western

Australia.

NOTE.—Sloane (1910) recorded a female specimen from Shark Bay, Western Australia.

12. Coptocarpus thoracicus (Castelnau)

FIGURES 1A, 3A, 14, 18, 29, 33

Oodes thoracicus Castelnau, 1868:151. [Lectotype, a male,

here selected in MHNP. Type-locality: Swan River, Western Australia.]

Coptocarpus parvus Sloane, 1915:461, [Holotype, a male, in

CSIRO. Type-locality: Karri Forest near Manjimup, Western Australia. New synonymy.]

DESCRIPTION (Figure 29).—Form much narrower

and flatter than C. australis members, but with

proportions similar. Recognized immediately on

small size and absence of scutellar interneur. Color:

Rufopiceous to piceous; head, pronotum, and sutural interval of elytron paler than elytral disc;

antennae and palpi testaceous, legs bright rufous,

tarsi rufotestaceous. Head and Prothorax: As in
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Male genitalia with everted endophallus, left lateral profile; anterior endophallus hook, left oblique aspect; and apex, ventral

aspect of Coptocarpus thoracicus, Margaret River, Australia.

C. australis. Mesothorax: As in C. australis except

scutellar interneur absent (Figure 1A), punctures

of interneurs 1 to 8 smaller, punctures of mesepisternum doarser. Metathorax: As in C. australis

except flight wings absent, metepisternum smaller

and more quadrate with fewer punctures (Figure

3A). Abdomen: As in C. australis except sterna I

to III coarsely and more sparsely punctate. Genitalia: Male (Figure 14); female as in C. australis.

Microsculpture: As in C. australis except reticulation of elytra much larger and coarser. Size: Length,

7.37 to 9.15 mm; width, 2.83 to 3.64 mm.

VARIATION.—This is a remarkably variable species. The width and general shape of the pronotum

vary in relation to the elytra, the pronotum being

narrower, subequal, or wider than the elytra and

the microsculpture of the elytra may be exceedingly coarse or quite smooth. There is apparently

no correlation with sex and microsculpture coarseness, as in some other ground beetles.

NATURAL HISTORY.—I have seen specimens collected in October, November, and January. The

January specimen appears to have been subteneral

when collected.

LOCALITY RECORDS (Figure 18).—I have seen 19

specimens from the following localities: AUSTRALIA: WESTERN AUSTRALIA: King George Sound

(MHNP); Manjimup in Karri Forest (CSIRO); Margaret River (CSIRO, MCZ, MHNP, USNM); Swan

River (MHNP).

13. Coptocarpus australis (Dejean)

FIGURES 1B, 2A, 3B, 5B, 15, 18, 30, 31

Oodes australis Dejean, 1831:671. [Lectotype, a female, here

selected in MHNP. Type-locality: "Nouvelle-Holande" as
NUMBER 184 17

given originally by Dejean, herewith restricted to Melbourne, Victoria, Australia, based on hand-labeled specimens in the Chaudoir collection in MHNP.]

Oodes reichei LaFerte, 1851:271 [type(s) lost].—Chaudoir,

1857:58.

Oodes convexus Castelnau, 1868:151. [Lectotype, a male,

here selected in MHNP. Type-locality: King George Sound,

Victoria, Australia. New synonymy.]

DESCRIPTION (Figure 30).—Form broad and convex with a transverse pronotum nearly as wide as

the elytra, and with a large head which, across the

eyes, is subequal to half the width of the pronotum.

Recognized easily by the character states mentioned

in the key. Color: Piceous to nigropiceous throughout; appendages, especially palpi, slightly paler.

Head: Broad across eyes, subequal to half the width

of pronotum; clypeus with setigerous pore at each

anterior angle; labrum with six setigerous pores

evenly spaced across anterior margin; mandibles

smooth dorsally; palpi glabrous; ligula bisetose,

medial carina broad and flat; mentum bisetose,

setigerous pores at base of marginal tooth; mentum

with anterior edge emarginate and unidentate,

tooth broad and apically rounded, margin with

raised bead extended from right to left lobe; submentum bisetose, setigerous pores at posterior lateral angles. Prothorax: Pronotum broad and convex, slightly narrower than elytra at widest point,

lateral margin narrowly beaded with bead extended

from hind angle around anterior angle halfway to

midpoint of anterior margin, disc with shallow

stria directed longitudinally at middle and disc

slightly depressed posterolaterally; proepisterna

finely and sparsely punctate; prosternum more

densely punctate anterolaterally; prosternal process

well developed, sulcate at middle and with raised

bead around apex, apex acute or rounded and

received into deep mesosternal groove; male tarsal

(Figure 2A) articles 1 to 3 broadly dilated and

asymmetric medially, ventrally (Figure 31) apical

FIGURE 15.—Male genitalia with everted endophallus, left lateral profile; anterior endophallus

hook, left oblique aspect; and apex, ventral aspect of Coptocarpus auslralis, north of Coonabarabran, Australia.
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 16.—Distribution map.

third of basitarsus and all of articles 2 and 3 with

numerous squamate setae (Figure 33) arranged in

transversely arcuate rows of 4 to 12 setae per row,

articles laterally with long and slightly flattened

setae, the lateral ones longer than the medial ones,

article 4 small and emarginate apically; apical tibial

spur long and stout, slightly arcuate, as long as

basitarsus. Mesothorax: Elytral interneurs striatepunctate, punctures small and widespread, striae

well impressed and not quite extended to basal

margin; scutellar interneur (Figure 1B) a row of

unconnected punctures extended apically to base

of interneur 1; intervals moderately convex, interval 3 with two small setigerous pores posterior of

middle; humeral angle with a very small tooth

which barely protrudes beyond margin; lateral

channel with numerous setigerous pores and small

tubercles anteriorly and posteriorly, less so medially; mesepisternum with numerous fine punctures; middle tarsal articles 1 to 4 (Figure 5B) with

bilateral rows of spinose setae ventrally. Metathorax: Flight wings fully developed with reflexed

apex; metepisternum coarsely punctate in apical

half (Figure 3B); metasternum smooth; hind tarsal

articles as middle ones. Abdomen: Sterna I to IV

rugosely punctate laterally; sterna III to V without

ambulatory setae; sternum VI bisetose in males,

quadrisetose in females. Genitalia: Male (Figure

15). Microsculpture: Head, pronotum, and elytra

with very small isodiametric reticulation throughout. Size: Length, 9.23 to 13.12 mm; width, 4.05

to 5.34 mm.

VARIATION.—The remarkable size difference between individuals has led to confusion among the

early students of the genus. All external character

states studied and the male genitalia indicate,

however, that there is probably but one species

involved.

NATURAL HISTORY.—P. J. Darlington, Jr., and

family collected members of this species in five

different localities, all of which are in savannah

woodland. According to Darlington (1960) specimens were collected from beneath logs or stones

in all these localities. I have seen specimens collected in May, June, October, and November; none

were teneral.

LOCALITY RECORDS (Figure 18).—I have seen 35

specimens from the following localities: AUSTRALIA: AUSTRALIAN CAPITAL TERRITORY: Gudgenby

River (MCZ, USNM). NEW SOUTH WALES: about

20 miles E of Armidale (CSIRO, MCZ), about 40

miles ENE of Glen Innes (MCZ), Ebor on Dorrigo

Plateau (MCZ), Manar Creek on Canberra Coast

Road (CSIRO), north of Coonabarabran (MCZ,

USNM), Paterson (MCZ, CSIRO). VICTORIA: Mel-
NUMBER 184 19

ahampionen8ie

? gibbus

FIGURE 17.—Distribution map.

bourne (MHNP), Ringwood (MCZ). WESTERN

AUSTRALIA: Cape DuBoulay (MHNP), Fremantle

(MHNP).

DISTRIBUTIONAL NOTE.—Sloane (1910) recorded

specimens from Western Australia, Mount Barker,

Beverley, Roebuck Bay.

NOMENCLATORIAL NOTES.—LaFerte (1851:271)

described Oodes reichei in a footnote to his table

of Oodes where he listed "Reichei (nobis). (1) id.

Swan river." Chaudoir (1857:58) stated that O.

reichei was a variation of Coptocarpus anstralis

and not "une espece distincte." Later, Chaudoir

(1882:510) stated that O. reichei was not discretely

described (more or less a nomen nudum) but equals

C. convexus. LaFert^'s description, although short

and inconcise, is a valid piece of work under the

present nomenclatorial code. Sloane (1910:446) apparently read Chaudoir's (1882) statement and

reported O. reichei as a nomen nudum, but equal to

C. convexus Castelnau. Since the type appears to

be lost, I am regarding O. reichei as synonymous

with C. convexus, in view of Chaudoir's (1882)

opinion, and therefore with C. australis.

Coptocarpus convexus Castelnau is based on a

very small specimen of C. australis; in all ways,

including male genitalia, it matches C. australis

Dejean; therefore I am regarding the two as

conspecific.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

+ australis

Qyorkensis

thoracicii8 D nitidus gj ahaudoiri

i philipi Q <&xJ2£ |£ fusaitareus

FICURE 18.—Distribution map.

Phylogenetic and Zoogeographic Notes

The striking similarities between the Australasian

Coptocarpus and the African "Thryptocerini" are

more than convergences. After analyzing many

character states of members of 23 genera or subgenera of Oodini, I believe that the apomorphic

trends set out in Table 1 amply indicate relationship among Coptocarpus, Thryptocerus, Orthocerodus, and Hoplolenus. An overall oodine study now

underway will clarify this situation. In the interim,

the following notes express my current opinions

with regard to Coptocarpus.

The 13 known species of Coptocarpus are arrayed in five species groups. The members of the

australis group are the most generalized of Coptocarpus, the most widespread, and two species have

winged members. These members, while having

objective synapomorphies 10, 15, 23 (see Table 1),

retain an Oodes form. Members of the other four

groups are variously apomorphic in form, they are

wingless, and they have some or all of the objective

synapomorphies 1, 2, and 4 (see Table 1).

The members of the philipi and chimbu groups

are most similar to Orthocerodus and Hoplolenus

members, respectively. The members of Thryptocerus are similar to members of Orthocerodus (thus

also, philipi group) except for the unique (among

oodines) elongate antennal scape and the oblique

insertion of article 2 into the scape. Hoplolenus

obesus members have peculiar mandibles (among

oodines), but in all other aspects are similar to
NUMBER 184 21

TABLE 1.—Distribution of character states in the four Oodine genera, Coptocarpus, Thryptocerus, Orthocerodus, Hoplolenus (asterisk = objective apomorphic condition, i.e., highly derived when distribution of state is examined

throughout the oodines; plus = character present; minus = character absent

or character state is different)

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

Character state

Head:

Labrum, number and pattern of

setae

Clypeus, number of setae

Labial palp, number of setae

Front, number of supraorbitai

setae per eye

Antennal scape, elongate

Antennal article 2, obliquely

inserted into scape

Prothorax:

Pronotum, number of setae

Proepisternum, punctate

Prosternum, punctate

Male, 3 basitarsomers

asymmetrically dilated

Male, tarsomere vestiture

transversely arranged

Tarsomere, laterally with

natatorial setae

Tibial spurs, elongate

Prosternal process, triangulate

and produced

Mesothorax:

Scutellum, very large

Ombilicate setal pore adjacent

to scutellum, foveate

Male, basitarsomere elongate

Mesosternal groove, deep

Metathorax:

Metepisternum, punctate

Metasternum, punctate

Male Genitalia:

Apex of median lobe, toothed

laterally

Right paramere, reduced to

nubbin

Abdomen:

Ambulatory seta absent

Coptocarpus

6, or 1-4-1*

0

#

,or2

0

0*,or 1

-

-

0

-.4 -

- , 4-

+ *

+ •

-

+

+ *

+ •

+ •

+ *

+ *

4-, -

4-, -

-

4-*

Thryptocerus

1-4-1*

0*

0

1

4-*

4-*

0

-

-

+ •

4-*

-

4-

+ •

+ •

4-*

+ •

+ •

-

-

+ , -

4-

4-*

Orthocerodus

1-4-1*

0*,or2

0

1

+ / "

-

0

-

-

+ •

+ •

-

4-

4-*

4-*

4-*

+ •

4-*

-

-

-

4-

4-*

Hoplolenus

1-4-1*

0*

0

1

-

-

0

4-

4-

4-*

+ • (Fig. 32)

-

4-

4-*

4-*

4-*

+ •

4-*

+

4-

+

?

4-*

Coptocarpus philipi and also C. nitidus of the

chaudoiri group. This indicates to me that if

Coptocarpus, Orthocerodus, Thryptocerus, and

Hoplolenus are each monophyletic groups, they

would have attained similar forms through convergent processes. Alternatively, if they had a common ancestral stock similar in form to australis

group members, and this ancestral stock was wide-
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

20

FIGURES 19-21.—Habitus, dorsal aspect: 19, Coptocarpus philipi, female, Mountains above

Atherton, Queensland, Australia; 20, C. chimbu, male, Dumun, New Guinea; 21, C. yorkensis

female, Iron Range, Queensland, Australia.

23

FIGURES 22-24.—Habitus, dorsal aspect: 22, Coptocarpus grossus, male, Margaret River, Australia;

23, C. doddi, holotype, male, Port Darwin, Australia; 24, C. nitidus, female, Bamaga, Australia.
NUMBER 184 23

\

25

FIGURES 25-27 Habitus, dorsal aspect: 25, Coptocarpus chaudoiri, lectotype, female, Clarence

River, Australia; 26, C. gibbus, lectotype, female, Australia; 27, C. fuscitarsus, lectotype, male,

Rafles Bay, Australia.

29

FIGURES 28-30 Habitus, dorsal aspect: 28, Coptocarpus championensis male, Geraldton, Australia; 29, C. thoracicus, male, Margaret River, Australia; 30, C. australis, male, Melbourne,

Australia.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spread throughout the Oriental, Ethiopian, and

Australian Regions, then similar forms and synapomorphies 3, 7, 10, 11, 12, 13, 14, 15, 16, 17, 18, and

23 (see Table 1) would be easily explained through

relationship. Perhaps with the evolution and

spread of the more successful Oodes members this

primitive stock of Coptocarpus-like animals was

replaced in the northern Ethiopian and Oriental

Regions, leaving the relict groups in the southern

Ethiopian and Australian Regions.

Because of the overall oodine study already mentioned, I do not want to rerank the "thryptocerine"

genera here. Also, I wish to point out that if the

alternate scenario above is correct we might expect

to find a relict or two of the Coptocarpus lineage

in the Oriental tropics. It is very possible that further study of the Indian Holcocoleus latus LaFert6

will prove to be the key to this problem. This

species has members strongly resembling the australis group members of Coptocarpus; the most

significant difference is the presence of ambulatory

setae on the abdominal sterna in Holcocoleus

members.

K

1

FIGURES 31-33—Right anterior tarsus, ventral aspect: 31,

Coptocarpus australis, male (X 74); 32 Hoplolenus obesus,

male (X 68); 33, suction cup setae of tarsus, Coptocarpus

thoracic us (X 295).
NUMBER 184 25

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Their Phylogenetic and Zoogeographic Significance

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1956. The American Group of Oodes. Coleopterists' Bulletin, 10:63-66.

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1873. Notes on a Collection of Insects from Gayndah.

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