H^-^o HARVARD UNIVERSITY Library of the Museum of Comparative Zoology bulletin OF THE Museum of Comparative Zoology ^ ^- Volume 142 1971 HARVARD UNIVERSITY CAMBRIDGE, MASSACHUSETTS 02138 U.S.A. CONTENTS No. 1. The Systematics, Distribution, and Zoogeography of the Marine Hatchetfishes (Family Stenioptychidae ) . By Ronald Clay Baird. October, 1971 1 No. 2. The Carabid Beetles of New Guinea. Part IV. General Considera- tions; Analysis and History of Fauna; Taxonomic Supplement. By P. J. Darlington, Jr. October, 1971 129 No. 3. A New Rhinoceros from the Late Miocene of Loperot, Turkana District, Kenya. By D. A. Hooijer. October, 1971 . 339 No. 4. Osteology of the Malaysian Phallostethoid Fish Ceratostethus hicornis, With a Discussion of the Evolution of Remarkable Struc- tural Novelties in its Jaws and External Genitalia. By Tyson R. Roberts. December, 1971 393 No, 5. Revision of North American Ciidae ( Coleoptera ) . By John F. Lawrence. December, 1971 . 419 ^^^^^^^^l^^-^;!?!l, o 18 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 \ V CO o U < X u > -Q O a o c Marine Hatchetfishes • Baird 19 O S3 CO D z z z < C/5 Z to ID a o ^ii£: 20 Bulletin Museum of Compavatwe Zoology, Vol. 142, No. 1 second neural spine to serve as a support- ing element. (In the P. spinosus species complex, the second neural spine resembles the first.) 2. Characteristic broadening and flat- tening of the haemal and neural spines in the posterior caudal region. 3. The presence of six or seven large, heavy, pleural ribs with relatively few reduced or vestigial ribs. This includes a low number (10-12) of abdominal verte- brae. 4. Development of the dorsal pterygio- phore system into a "blade" or spine. 5. A vertically oriented pelvic girdle, the basipterygia bearing spines, sometimes fused, and closely allied to the heavy pleural ribs. 6. A preopercle with a well-developed ventral spine. 7. A heavy, forked, post-temporal which is fused to the supracleithrum in Argijro- pelecus and Polyipnus, formmg a spiny extension dorsally. 8. A progressive migration forward of the suspensorium. 9. Reduction of the bony extension of the urohyal. 10. Epiotics meeting below the supra- occipital and the presence of well-de- veloped, ridged parietals. 11. Presence of a well-developed ab- dominal keel-like structure which is ossi- fied in Argijropelecus and Polyipnus. 12. Presence of a circular gap in the anal pterygiophore series, these pteiyg- iophores being enlarged. 13. Presence of ventral processes on the anteriormost anal pterygiophore in Ster- uoptyx and Polyipnus. 14. Marked similarity of photophore pattern and number. Some of these character complexes are not radically different from the gonosto- matids examined, and there is a degree of convergence and parallel evolution which is difficult to appraise. Taken as a whole, however, they strongly suggest that the stemoptychid genera have reached a com- mon evolutionary grade, typified by their peculiar body form, and by which they differ from the more generalized and primitive maurolicid gonostomatids. While acknowledging that the Sterno- ptychidae are a specialized offshoot of maurolicid or premaurolicid stock, for the following reasons I do not feel justified in combining the Gonostomatidae and Ster- noptychidae as some have suggested. The present family Gonostomatidae is an unwieldy one which involves many diverse types and requires extensive revision (Weitzman, personal conversation). The problem of gaps, their size and importance, cannot be adequately answered without further stud>- within the Gonostomatidae. Osteologically, the Sternoptychidae have reached an evolutionary grade peculiar to themselves and one quite distinct in several major ways from the gonostomatids ex- amined. Using for a guideline the family concept as it is generally employed by Mayr, Linsley, and Usinger (1953), it ap- pears that the Sternoptychidae do have an ecological, or at least adaptive, distinctness. The adaptive distinctness concerns the peculiar body shape and its possible func- tional significance. Tliere are at least two major adaptive features involved. The first deals with the ideas and evidence presented by Denton and Nicol ( 1965 ) and Nicol (1967) on the relationship between silvery color and body shape in teleost fishes. The midwater environment is one in which the distribution of daylight is in- dependent of the altitude of the sun and cloudiness of the sky, and light distribution is essentially symmetrical about a vertical to the surface. Furthermore, the Sterno- ptychidae have brilliant, silvery sides. All fish species with these features so far examined (Denton and Nicol, 1965; Nicol, 1967) have layers of reflecting platelets \\'hich are oriented to make the fish as invisible as possible. It may be assumed that the same is true with hatchetfish. Tliere is a change in reflectivity with body rotation in the several Stemoptychid Marine Hatchetfishes • Baird 21 species examined. A silveiy fish which is flattened laterally, having very little in- clined ventral surface will approach the ideal in camouflaging (see Denton and Nicol, 1965: 717). The Sternoptychidae could thus serve as a living model for such a body fonn. The second adaptive feature concerning body shape is the development of heavy structural ossifications and spines, espe- cially the dorsal "blade." Spines have developed in fish, presumably, for pro- tection. The sternopt)'chids have several extensive spine complexes: post-abdominal, post-temporal, preopercle, and dorsal. The spines are rigidly braced and the whole body strongly ossified, resulting in a compact rigid body shape. A spinous dor- sal has developed somewhat analogous to that of the higher Perciform fishes. This, coupled with the expanded abdominal region, results in a high length-to-depth ratio ( Table 1 ) . In an environment populated by a host of predators, many with special adap- tations for ingesting large prey items, an increase in the length-to-depth ratio of a prey should be advantageous. A predator nomially capable of swallowing Valen- ciennelhis would require an approximate threefold increase in mouth diameter in order to accommodate A. hemigijmnus of the same length (Table 1). Ossification also takes place quite early. Juveniles or prejuveniles of about 10 mm have well- developed spines and are ossified. Pliylogenetic relationships. The question of a monophyletic origin of the hatchetfish is unanswerable. The three genera show a great deal of divergence and independent evolution even within genera. Using the character complexes examined, some com- ments about generic relationships can be made, however. The family appears primitive and prob- ably originated from a i^remaurolicid an- cestor, possibly something between the very early Vinciguerria and Maiiroliciis. Most of the characters examined could have been derived from a form somewhat intermediate to the above genera. The genus Sternoptijx seems to have di- verged quite early from the line or lines leading to Pohjipniis and Argyropelecus. It then continued to evolve independently, resulting in the present highly specialized form. In almost every case, Sternoptijx shows marked differences. The presence of a basisphenoid, the characteristically shaped, enlarged, first anal pterygiophore; the simple anterior, dorsal pterygiophores; possibly the meeting of the parietals, and the unfused post-temporal and supra- cleithrum all appear primitive. These char- acters are also shared with Polyipnus with the exception of the unfused post-temporals and meeting parietals. The presence of a small premaxilla and large maxilla as the major jaw bone in the gape are generally regarded as primitive. However, jaws and dentition have varied considerably in gonostomatids (Grey, 1964), and this may be a secondary phenomenon. The dis- appearance of the anterior pedicels of the premaxilla and loss of the second supra- maxilla can be explained in the same way, especially since the orbital region seems to have undergone considerable expansion. The resemblance of the urohyal to Argy- ropelecus may again be the result of parallel or convergent evolution involving feeding ecology which is similar in these genera. Evolution from a premaurolicid ancestor can be traced somewhat more directly in the case of Polyipnus and Argyropelecus. Polyipnus and Argyropelecus share several character complexes: the characteristic blade-shaped, caudal haemal spines; the presence of the double pterygiophore as the major element in the "blade"; the presence of ossified, bony keel j)lates; the fusion of the post-temporal and supra- cleithrum; and separation of parietals by the supraoccipital (known to be variable in the Gonostomatidae). Polyipnus ap- pears intermediate between Maurolicus and Argyropelecus in several characters: 22 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Table 1. Body depth and standard length MEAsuIlE^'rENTS. Family Species SL ( mm ) Maximum Body Depth* ( mm ) Gonostomatidae Stemoptychidae Danaphos aculatus Valenciennellus tripunctatiis Argtjropelecus hemigymus Argyropelecus hemigymmis 38.0 26.0 38.0 26.0 9.2 5.4 24.0 16.9 * Includes dorsal blade. the axial skeleton in general; anterior dorsal pterygiophore development; pelvic girdle modification; evolution of the cleithrum, first branchiostegal rays, and urohyal; the opercular series, especially the preopercle and interopercle; suspensorium develop- ment; parasphenoid curvature, and pro- gressive deepening and shortening of the body with reduction in a long unbroken series of ventral photophores. PoJyipmis has characters that are not shared with Argyropelecus in addition to those which are shared with Sternoptyx. These include: hypurals 5 and 6 unfused (3 and 4 also in the P. spinosus complex); small, relatively unmodified dentition; and, a urostylar element with several unfused post-terminal centra. Polyipmis also has several highly special- ized characters: the peculiar jaw morphol- ogy; a greatly enlarged otic region with characteristically shaped, large otoliths (Kotthaus, 1967); and, the peculiar de- velopment of the cleithrum (pectoral shield ) . The divergence of Polyipnus and Argy- ropelecus has involved the continued evo- lution of many intermediate characters mentioned above. Other major develop- ments in Argyropelecus are: fusion of hypurals 5 and 6 and the post-tenninal centra; the presence of seven rather than six heavy pleural ribs; development of a lateral preopercular spine (one species of Polyipnus has this); development of a fanglike dentition; loss of the basisphenoid; loss of the flangelike process on the first anal pterygiophore; and, the development of telescopic eyes. Because the number of character com- plexes examined was limited, the suggested phylogeny is only a tentative one. The family consists of three divergent, inde- pendently specialized genera. Polyipnus ajDpears the most primitive, Sternoptyx the most highly specialized and the most diffi- cult to place, while Argyropelecus falls somewhere in between. THE FOSSIL RECORD The earliest reported stemoptychid fossil is from the Eocene of the Dabakhan beds of Georgia, USSR ( Daniltshenko, 1962). Tlie fossil, Polyipnoidcs levis, is not well preserved and many important characters cannot be appraised. It does have long pleural ribs and a characteristic broadening of the body anteriorly. Tlie post-temporal, however, is unlike any modem stemopty- chid. The dorsal "blade" or pterygiophore development is absent and the jaws seem more gonostomatidlikc, although this is difficult to determine with certainty. The neural and haemal spines show little char- acteristic flattening, and the frontals do not exhibit the heavy development char- acteristic of the hatchetfish. Consequently, it appears that while this fossil could be a proto-stemoptychid fish, I cannot accu- rately place it with the present Stemopty- chidae or Gonostomatidae. Polyipnus sobnioviensis was reported from the Jaslo shales of Poland (Jerz- maiiska, 1960; Jerzmanska and Jucha, 1963) and dates as late Eocene-early Oligocene. Enlarged pleural ribs with a general broad- ening of the body anteriorly are present in this species. Pterygiophore development Marine Hatchetfishes • Baird 23 anterior to the dorsal fin rays is definite, above the dorsal fin origin, low; last pleural and there is evidence of very slight, dorsal rib only slightly reinforced; anterior haemal blade development. Some photophore spines not greatly flattened, postabdominal groups conform roughly to modem Poly- spines short, symmetrical, not markedly ipmis, although the fossil supra-abdominal curved; transitional vertebrae two in num- group is more numerous. The cleithrum ber; anal pterygiophores relatively simple, displays the marked ventral curve typical not markedly broadened at distal end; anal of the Stenioptychidae and the maurolicid- pterygiophore gap contains t\vo haemal gonostomatids. There is, however, little spines; number of anal pterygiophores be- flattening of haemal and neural spines. The fore gap, seven, after gap, four to five; pelvic girdle, while partially vertical, is hypurals 1 and 2 separate; number of still below the rib line, and the body shape, vertebrae from posterior margin of dorsal while somewhat broad, is more similar to blade to last neural spine, 29. the maurolicid gonostomatids. Tliere is no spine on the preopercle, the orbit shows no Fossil B (Figure 13) great expansion, and there are no signs of . , r,. -i t-. i * i keel plates. Tliis fish, while it has some Description. Similar to Fossil A above; sternoptychid characters, appears essen- f ^™ber of vertebrae from dorsal blade to tially to be maurolicid-gonostomatid. Con- ^^'^ "^^^^1 spine 30; hypurals 1 and 2 sequently, its place in the genus PoZy^-pm/s separate; postabdommal spines simple, is questionable, although it may be near symmetrical; posteriormost pleural ribs not the basal stock which gave rise to modem greatly enlarged, hatchetfishes. Pauca (1931) described Stemoptyx Fossil C (Figure 14) prisca from the lower Oligocene deposits Description. SL 60 mm, body depth 40 of Piatra Neamt. The presence of a well- mm; number of vertebrae from posterior developed dorsal "blade," heavy cleithmm, dorsal blade to last neural spine 26, pos- and pleural rib characteristics place it in sibly 27; both abdominal and trunk regions the genus Argyropelecus. If the dating is greatly broadened; hypurals 1 and 2 fused; correct, it represents the earliest known anterior haemal spines broad, flat, blade- fossil of this genus. lilce; distal end of anal pterygiophores By Ohgocene, and certainly by Miocene broad, gap well developed, circular, and times, several examples of the genus includes two neural spines. Argyropelecus were evident in Tethys Fossils A and B are indistinguishable in deposits of Europe (Arambourg, 1929; both key characters and meristics from the Daniltshenko, 1960), and in various de- modem species A. affinis (Fig. 15) and posits of California (David, 1943). All of can be assigned to this species complex, these fossils clearly represent members of Fossil A seems broader than the modern the above genus, and A. logearti (Aram- form, but the fossil appears distorted ven- bourg, 1929) appears to be closely related trally and there are no other obvious dif- to the modern A. hemigymniis. ferences. In the present study, three remarkable Fossil C is a member of the A. lychnus fossils from Miocene deposits in California complex (Figs. 11 and 16). Osteologically were examined and compared with modern there is little difference between A. olfersi relatives. and A. lychnus. However, the fossil has a relatively low dorsal blade, measured Fossil A (Figure 12) from the origin of the dorsal fin rays, a Description. SL 50 mm, body depth 26 characteristic of A. lychnus (Fig. 16). The mm; dorsal blade from its extension second transitional vertebra of Fossil C 24 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 a U J2 O u o CN < Marine Hatchetfishes • Baud 25 1 ' ' V ■1 , \ V s -% '"> r^ ^ 1 V I ■!>*- ¥ o U CD > o I u o 2 < 26 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 a U o en _o o o u o u < u Marine Hatchetfishes • Baud 27 •o o •f CM CO X / - £3iW SHI ^ ^^^pJMP^rA. L i b^^v Art o V3 U < X u ID a o >.. 28 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 ^0 CM 1/1 3 Z o z < c -c a o Marine Hatchetfishes • Baird 29 Key to the Genera of Sternoptychidae la. Abdominal photophores 12; telescopic, dorsally oriented eyes; several dorsal pterygio- phores fomi extensive blade anterior to dorsal rays genus Argijropelecus {-p. 31). b. Abdominal photophores 10; eyes normal; dorsal blade consisting of only one or two spines from a single or two fused pterygiophores 2 2a. Anal photophores 3; no supra-abdominal photophores; single large dorsal spine with anterior serrate extension; first anal pterygiophore greatly enlarged, supporting tri- angulate transparent membrane above anal fin rays genus Sternoptyx (p. 67). 1). Anal photophores 6 or greater; 3 supra-abdominal and a lateral photophore; dorsal blade reduced; no large transparent membrane above anal fin rays genus Poltjipmis (p. 79). has a pair of vestigial ribs whereas the one time also. Many modern gonostomatid modem A. hjchnus examined does not. genera were present during that time, and One of the specimens of A. olfersi ex- Miocene faunas have distinctly modem amined has small vestigial ribs on this resemblances (David, 1943; Grey, 1964; vertebra; otherwise, all of the fossil char- Crane, 1966; Daniltshenko, 1960). The acters and vertebral counts are identical to salmonoid-derived midwater fauna appears these modern species. Fossil C is probably to have replaced earlier forms during the A. hjchnus or at least its immediate prede- early Tertiary, and it remains the dominant cessor. element today. By mid-Miocene times evolution within the genus Ar^ijropehciis was essentially SYSTEMATICS complete and species distributions show modem characteristics. This genus with '^^'^''y STERNOPTYCHIDAE its many specializations must have origi- ^/P® Genus: Siernopiyx Hermann 1781 nated by the late Eocene at the latest and Diagnosis. Neural spine of first pre- possibly as far back as the Paleocene or terminal vertebra vertically oriented, late Cretaceous. broadened, with triangulate paddle shape, Durmg or prior to the early Cretaceous, no fin rays attached; second preterminal some members of the early salmonoid vertebra modified for support of first (ex- fishes began to adapt to a deep water eept F. splnosus complex); basiptervgia envn-onment. After the basic adaptations verticallv oriented, spine bearing, con- to this environment were acquired (at tained dorsally within, and closely joined latest mid-to-late Cretaceous) there was to the ventral margin of the posterior considerable stomiatoid radiation which pleural ribs; pelvic fin ravs xertically continued into the late Eocene to early oriented; six to se\'en i^leural ribs enlarged Ohgocene. This radiation led to many to form an expanded rib cage; epiotics diverse forms, of which the maurolicid- meet below supraoccipital; parietals well gonostomatids ^^'ere one. Within the latter, developed, bearing dorsolateral ridges; one an ancestor, possibly resembling P. sohnlo- or more dorsal pterN giophores enlarged to viensis, gave rise to a form or series of form blade or spinelike extension anterior forms with many features of the modem to dorsal rays; anal ptervgiophores form genus Pohjlpmis. From this basic stock the characteristic gap below anal photophore modern genera evolved, conceivably quite group; preopercle bearing well-developed rapidly. By the Miocene, evolution was ventral spine. practically complete in the specialized Description. Bright silverv colored, Argyiopclcciis and possibly the other small fishes; standard length usually less genera as well. The stomiatoid-gonosto- than 90 mm; body deep" strongly com- matid radiations of the early tertiary show pressed; bony scalelike plates forni keel evidence of being fairly complete by that below ventral photophore groups (except 30 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 PTO PRO Figure 17. Photophore and spine characteristics: top — Po/yipnus; bottom — Argyropelecus. Abbreviations — photophores: see p. 6; spines: DB = dorsal blade; PAS ^ postcbdominal spine; PTS z= post-temporal spine. Sternoptyx); 10-12 abdominal, and four muscular stomach, five or more plyroic subcaudal photophores always present (see caecae, and short straight intestine; eyes Figs. 17 and 18 for photophore and spine large, well developed; gape vertical; adi- characteristics ) ; nasal lamallae well de- pose fin usually present; scales thin or veloped; digestive tract simple, with thick absent except along ventral surface; swim Marine Hatchetfishes • Baiid 31 Figure 18. Photophore characteristics: genus Sternoptyx. Abbreviations: see p. 6. bladder present ( see above for osteological description ) . Genus ArgyropeJecus Cocco, 1829 Argijropelecus Cocco, 1829: 146 (type species: ArgyropeJecus hemigijmnus Cocco, 1829, bv monotypy ) . Fleurothysls Lowe, 1843: 64 (type species: Sterno- ptyx olfersi Ciivier, 1843, by original designa- tion ) . Sternoptychides Ogill^y, 1888: 1313 (type spe- cies: Sternoptychides amahUis Ogilby, 1888, by monotypy ) . Diapiosis. Twelve abdominal, six supra- abdominal and two suprapectoral photo- phores; eyes telescopic, dorsally oriented; frontal ridges compressed dorsally above eyes; basisphenoid absent; several teeth directed anteriorly on posterior maxillary margin; dorsal "blade" consisting of several broadened pterygiophores anterior to dor- sal rays; seven enlarged pleural ribs. Description. Photophorcs: PO 1; PTO 1; BR 6; I 6; AB 12; PRO 1; so 1; SP 2; SAB 6; PAN 4; AN 6; SC 4 (for anatomical details see Brauer, 1908; Bassot, 1966). Spines: Post-temporals extended pos- teriorly to form a small spine; preopercle bears one ventrally and one posteriorly directed spine; retroarticular bears ven- trally directed spine; basipterygia extended ventrally bearing one or two postabdominal spines; cleithrum extends ventrally form- ing preabdominal spine; spiny scales present in adults of some species below subcaudal and preanal photophorcs. Eyes: Large, well developed, telescopic, lens dorsally oriented, fitting into dorsal grooves in the frontal bone. Gill Rakers: Total 15-24; rakers well developed with rough toothlike surface; epi- and ceratobranchials bear well-de- veloped spines on internal surface. Jaws and Dentition: Jaws somewhat vertically oriented; premaxilla well de- veloped, toothed, and majoi- upper jaw bone in gape; maxilla also somewhat in- 32 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 eluded in gape, toothed, the posteriormost teeth eurved markedly forward; lower jaw sturdy, heavily toothed, oecasionally with large eanines; dentition eonsisting of multi- rowed single cusped, curved caninelike teeth; palatine teeth present, often well developed; epibranchial of third and fourth arch extends ventrally and laterally to form toothed plates. Meristies: Vertebrae 35-40; C. 9+10; D. 8-10; A. 6-8 + 5-6. Color: Bright silvery in life, quickly lost in formalin preservative; dark pigmenta- tion often striated posteriorly; stable for long periods in preservation. Internal Anatomy: Relatively thin-walled swim bladder (see Marshall, 1960) and gas gland well developed; digestive tract simple, consisting of heavily pigmented, double compartmented stomach; the an- terior internal lining very thick walled and covered with rasping tubercles; posterior lining thin and distensible, five to seven thick-fingered pyloric caecae, large liver, and a short straight intestine; caelomie cavity lined with heavily pigmented mem- brane; gonads when mature fill the dorsal and lateral posterior half of the body cavity; nephritic tissue moderately well developed. Species complexes. There has been sub- stantial radiation within the genus and even to some extent within the species complexes. The A. af finis complex appears to be the most primiti\'e and other forms can be derived from it. Primitive char- acters of this complex include: three hypural elements in lower caudal lobe; posterior ventral photophores in an almost unbroken series; glandular photophore ar- rangement simple, the posterior photo- phores not joined in glandular clusters; little reinforcement of posteriormost pleural rib; body not markedly deepened ante- riorly; basipterygia lacking support ami for keel plates; generally unspecialized axial skeleton, including lack of marked broad- ening of anterior haemal spines; vertebral number 38-40. The more advanced members of the genus are characterized by two hypural elements in the lower caudal lobe; pos- terior photophores joined into distinct glandular clusters; a general deepening of the anterior body region with subsequent reduction in vertebral number; increased complexity of structural ossification espe- cially in the axial skeleton, including a marked reinforcement of the last large pleural rib, and a keel supporting ex- tension on the left basipterygia. A. he mi [1,1/771)1 lis appears slightly more primitive than the A. h/climis complex and is highly specialized. Important characters include: primitive transitional vertebrae (like A. af finis):, dwarfism (maximum length 38-40 mm); fused basipterygia forming single postabdominal spine; 38 vertebrae; epiotics with dorsal extensions (Fig. 7); peculiar dorsal blade shape, often \\'ith supplementary spiny spurs on the major element. The most specious group, the A. hjchmis complex, shows a high degree of structural ossification in the axial skeleton, including the dorsal and anal pterygiophore systems; some species have developed long, fang- like canines in the lower jaw; there is a marked deepening of the anterior body region with reduction in vertebral number. Key to the Species of Argijwpelectis la. Snpra-abdominal, pieanal, anal, and subcaudal photophores in a nearly continuous straight line; subcaudal photophores separate, an appreciable gap between each photo- phore; .3 separate hypural elements in lower caudal lobe A. affinis complex 3 b. Supra-al)dominal, preanal, anal, and subcaudal photophores not in a continuous straight line; subcaudal photophore group in a single glandular cluster, no appreciable gap be- tween each photophore; two separate hypural elements in lower caudal lobe 2 A single posteriorly directed, serrate postabdominal spine; gap between anal and sub- caudal photophore groups greater than 2.2 times gap between preanal and anal groups; dorsal rays 8 A. heiiiigyiuiius (id. 42). 'Sa Marine Hatchetfishes • Baird 33 b. Two separate postabdominal spines; anal subcaudal gap less than 2.0 times anal-pre- anal gap; dorsal rays 9 A. lijchntis complex 4 3a. Dorsal blade low, its height less than one-third its length; body margin not markedly raised posterior to dorsal blade; ventral keel scales do not extend far below abdominal photophores; no laterally directed sphenotic spine near dorsal, posterior edge of orbit A. affinis (p. 34 ) . b. Dorsal blade high, its height greater than one-third its length; body margin markedly raised posterior to dorsal blade; ventral keel scales extend well below alsdominal photo- phores forming flaplike process; prominent laterally directed sphenotic spine near dorsal, posterior edge of orbit A. gigas (p. 38). a. b. 4a. Posterior postabdominal spine directed posterioventrally and markedly larger tlian the anterior; anterior margin of posteriormost abdominal keel scale slants markedly forward; standard length less than 3.4 times body depth; pair of enlarged canine teeth present in lower jaw; subcaudal spines present A. aculeatus (p. 48). h. Both postabdominal spines of about equal length and size; anterior margin of posterior- most abdominal keel scale almost vertical; SL greater tlian 3.5 times body depth; pair of enlarged canines may or may not be present in lower jaw; subcaudal spines may or may not be present 5 5a. Pair of enlarged canine teeth in lower jaw; sharply pointed anteriormost postabdominal spine curved markedly and evenly forward; upper preopercular spine short, not extend- ing much beyond posterior border of preopercle; outermost tips of dorsal and ventral- most caudal rays streaked with dark pigment (fish greater than 35 mm) - — - A. olfersi (p. 52). b. No pair of enlarged canines in lower jaw; anteriormost postabdominal spine squared or blunt (except very small individuals), not curving evenly foi^ward; upper preopercular spine extends well beyond posterior border of preopercle; no pigment on outermost caudal rays 6 34 BiiUetin Museum of Comparative Zoology, Vol. 142, No. 1 fia. Dorsal blade low, blade height less than 2.5 mm for SL 25-50 mm, less than 3.1 mm for SL 50-70 mm; body narrow (see regression, body depth, Table 11); no spines on scales below subcaudal photophores; dark well-developed pigment spots fonn line along posterior midline ( especially in smaller sizes ) ; upper preopercle spine usually ciuved dorsally, never ventrally — . A. .sladeni (p. 56). b. Dorsal blade high, blade height greater tlian 2.7 mm for SL 25-50 mm, greater than 3.8 mm for SL greater than 50 mm; body robust (see regression, body depth, Table 14); spiny scales present below subcaudal photophores (fish greater than 30 mm SL); pigment spots minute along posterior midline; upper preopercle spine usually ciuved ventrally, never dorsally A. hjchnus (p. 63). a. Argyropelecus affinis Garman Figure 19 Argyropelecus affinis Garman, 1899: 237 (holo- type USNM 44593; tropical North Adantic; not seen); Brauer, 1901: 120; 1906: 103 (fig. larvae); Regan, 1908: 218; Barnard, 1925: 153; Norman, 1930: 301 (fig.); Jespersen, 1934: 15 (fig.); Fowler, 1936: 221; Beebe, 1937: 201; Parr, 1937: 49; Norman, 1939: 19; Nybehn, 1948: 23; Misra, 1952: 367; Smith, 1953: 102; Haig, 1955: 321; Fowler, 1956: 67; Schultz, 1961: 597 (fig.): Bahamonde, 1963: 83; Blache, 1964: 71 (fig.); Schultz, 1964: 241 (fig.); Backus et al., 1965: 142; Bussing, 1965: 185; Bright and Paquegnat, 1969: 27. Argyropelecus pacificus Schultz, 1961: 599 (fig.); 1964: 241; Berry and Perkins, 1965: 625; Lavenberg and Ebeling, 1967: 185. Species distinction. Differs from A. gigas (in addition to key characters) by its nar- rower body depth and trunk (see regres- sion, body depth. Tables 2 and 3); less distinct trunk striations; relatively longer teeth in lower jaw; less well-developed post-temporal spines; smooth dorsal body surface; and less well-developed neuro- cranial crests (frontals, sphcnotics, and parietals ) . Description. D. 9; A. 12-13; P. (10) 11; total gill rakers 18-22; vertebrae 38-39 (40). Medium size species rarely exceeding 70 mm SL; body more evenly tapered than others in genus; body depth at end of dorsal greater than 3.5 times into SL; caudal peduncle long and narrow, its depth Marine Hatchetfishes • Baud 35 /-':'' -^''iV" ■•^'■. / :■::::■:■[: y^-''4>U i ^ '0.1 J Figure 19. Argyropelecus altinis; R/V CHAIN, Cruise 60; Station 1257; SL 51 mm. less than length of siibcaudal photophore opercle spine directed latero-anteriorly; group; dorsal spine low, its height less ventral preopercle spine long, curved an- than one-third its length; post-temporal tcriorly; jaws large; teeth short, recurved, spine short; postabdominal spines of equal better developed in lower jaw; gill rakers size, with no marked curving; dorsal pre- long, closely set; in preservative, trunk Table 2. Regression statistics for various populations of A. affinis. Character Indian Ocean (5°-12°N, 160°-168°E) Body depth Jaw length Gtilf of Guinea Body depth Jaw lengtli Jaw width NW Atlantic (30°-33°N, 73°-78°W) Body depth Jaw length Jaw width S£ Pacific (Chile) Body depth Jaw length NE Pacific (California) Body depth Jaw lengtli Regression -1.83 -0.78 -2.44 -0.27 -0.23 2.59 -0.78 -1.35 0.55 0.64 -1.02 0.27 0.49 ± .146 0.25 ± .065 N — 11 0.46 ± .121 0.22 ±: .060 0.12 ±2 .059 N = 10 0.37 ±2 .176 0.24 ^ .107 0.14 ^ .087 N 7 0.42 + .086 0.21 ^ .047 N — 13 0.49 ^ .062 0.22 ± .029 N = 19 36 Bulletin Mitseuni of Comparative Zoology, Vol. 142, No. 1 o -o > -a (U E o □ o c o o X Marine Hatchetfishes Baird 37 D E P H A • B ▲ C O D • FISH HR Figure 21. Diurnal vertical distribution of A. alfinis determined by rate of capture with depth during the day (D) and night (N). A = Pacific (California); B = Pacific (Chile]; C = Gulf of Guinea; D = Gulf of Mexico and Caribbean. region exhibits cross pigment striations with well-defined, midlateral line. Distribution. Horizontal distribution (Fig. 20): Taken abundantly in the Gulf of Guinea, off California, Chile, and in the northern Indian Ocean; moderate catches are recorded from the northern Gulf of Mexico and the coast of Venezuela in the Caribbean; smaller catches w^iich may rep- resent possible populations are recorded southeast of Hawaii, south of Java, and off the southeast coast of the United States; scattered samples representing this species appear in the Bay of Bengal, Gulf of Aden, tropical Atlantic, and off the southeast coast of Brazil. (Additional records: At- lantic, occasional catches between Azores and Madeii-a; Pacific, moderate catches near coast of northern Peru. ) Vertical distribution (Fig. 21): Appears concentrated between 350 m and 600 m by day with the highest concentrations in the vicinity of 400 m; by night the dis- tribution is somewhat more shallow, major 38 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 —I— 18 I iO I 22 Figure 22. Geographic variation in gill raker number in A. affinis. A := Gulf of Guinea; B ^ Caribbean; C ;= Indian Ocean; D = Pacific (Chile); E = Pacific (Cali- fornia); F = NW Atlantic (NW Atlantic pocket). Numbers refer to sample size. concentrations occnning from 170 m to 400 m. With the possible exception of the Gulf of Guinea, there are no indications of marked geographic variation in depth dis- tribution, although Appendix C indicates slightly shallower daytime depths off Cali- fornia than in the tropical Atlantic. Geo^ i (U c o o I 48 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 aO B D C O □ • E -Q E o 0) o u "o -^ 'o ~5 o X 54 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 • • • •0- -ttH ♦ Zoo ■♦t n* * 400 ♦ • • • t>00 . D 8oo D lOOO 15 00 N A O B O C • D -d E f F D 10 20 30 FISH HR Figure 37. Diurnal vertical distribution of A. aculeatus determined by rate of capture with depth during day (D) and nighf (N). A = Pacific (Chile); B = N Central Pacific; C = Caribbean and Tropical Atlantic; D = NW Atlantic; E = Gulf of Mexico; F = NE Atlantic. Holt and Byrne, 1913: 120; Jespersen, 1915: 23; 1934: 15; Roule and Angel, 1933: 46; Buen, 1935: 52; Parr, 1937: 49 (spines); Bertin, 1940: 314 (holotype); Nybelin, 1948: 23; Bertelsen and Grontved, 1949: 163 (light organs); Maul, 1949b: 13; Dollfus, 1955: 1; Holgersen, 1958: 120 (population density); Koefoed, 1961: 10; Schultz, 1961: 610; 1964: 241; Wheeler, 1969: 136. Species distinction. Differs from A. aculeatus by absence of subcaudal spines, less deep body (see regressions, body dcptli, Tables 8 and 10), lower dorsal spine, higher \'ertebral count and post- abdominal spine characteristics; differs from A. lychnus by presence of enlarged canines, lighter pigment, no subcaudal spines, preopercle and post-temporal spine characteristics and first anal photophore; differs from A. sladeni by presence of enlarged canines; no definite pigmented Marine Hatchetfishes • Baird 55 Table 8. Regression statistics for various populations of A. aculeatus. Regression Character SE Pacific (Chile) Body depth Jaw length Gulf of Mexico (24°N, 83°W) Body depth Caudal peduncle depth Jaw length Caribbean (13°N, 71°W) (Sample 1) Body depth Caudal peduncle depth Jaw length Caribbean (13°N, 71°W) (Sample 2) Body depth Caudal peduncle depth Jaw length N\V Atlantic (42°N, 47°W) (9/64) Body depth Caudal peduncle depth Jaw length NW Atlantic (41°N, 62°W) (9/64) Body depth Caudal peduncle depth Jaw length NW Atlantic (42°N, 62°W) (9/62) Body depth Caudal peduncle def)th Jaw length NE Atlantic (32°N, 13° W) Body depth Caudal peduncle deptli Jaw length -1.64 1.27 0.49 0.12 0.70 2.32 0.20 0.70 2.85 0.39 0.66 0.69 -0.06 -0.05 0.60 0.23 -0.17 0.25 0.57 -0.47 1.32 1.34 -0.72 0.77 ± .274 0.22 ± .087 N = 8 0.67 ± .084 0.12 ± .015 0.23 ± .032 N = 23 0.64 ± .064 0.12 ± .013 0.24 ± .025 N=r23 0.63 ± .080 0.11 ± .018 0.25 ± .034 N = 26 0.66 ± .087 0.12 ± .020 0.26 ± .031 N = 28 0.67 ± .061 0.12 ± .012 0.26 ± .025 N = 30 0.67 ± .055 0.11 ± .010 0.27 ± .038 N = 40 0.69 ± .123 0.10 ± .019 0.29 ± .042 Nzz29 Table 9. Slope comparisons of regressions of various characters between two populations of A. ACULEATUS. ThE CARIBBEAN POPULATION CONSISTS OF TWO SUBSAMPLES ( SEE TabLE 8). Character PopulaHon 1 Population 2 Caudal peduncle depth NE Atlantic Caribbean 2 2.009 .05 Jaw length NE Atlantic Caribbean 1 2.266 .038 Jaw lengtli NE Atlantic Caribbean 2 2.059 .048 56 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 15 A B Table 10. Regression statistics for two popu- lations OF A. OLFERSI. □ 1 15 11 15 1 Character Regression ^51^ A B NE Atlantic Body depth Jaw length Southern Ocean Body depth Jaw length (Pacific) 0.51 0.56 1.74 -0.19 0.64 ± .231 0.26 ± .099 K = 8 0.61 ± .158 0.28 ± .075 N = 10 T-» • . .7 1 1 -I . . \h 18 Figure 38. Geographic variation in gill raker count in A. aculeatus. A = NW Atlantic; B = NE Atlantic; C = N Central Pacific; D = Pacific (Chile); E = Caribbean; F =1 Gulf of Mexico. Numbers refer to sample size. midline; deeper body (see regression, body depth, Table 11), spine characteristics, anal photophores, and lower gill raker count. Description. D. 9; A. 12; P. 10-11; total gill rakers (15) 16-17; vertebrae 36-37 (38). Large species often exceeding 70 mm SL; body deep, depth at end of dorsal usually greater than 1.5 times into SL; first preanal photophore with pointed dorsal margin; dorsal spine high, its height nearly one-half its length; post-temporal spines well developed; postabdominal spines nearly equal, anteriormost spine curves smoothly forward; lower preopercle spine long, curving fonvard, upper very short; jaws large; teeth recurved with two large canines in lower jaw and a somewhat smaller pair in the upper jaw; pigment diffuse over whole of trunk; no marked midline f)igment spots; less marked con- centration of pigment in caudal peduncle; dark pigment present on outermost caudal rays (this often lost in handling). Distribution. Horizontal distribution (Fig. 36): Restricted to the northeast Atlantic between latitudes 35°N and 65°N and east of longitude 35°W; occurs in a broad band across the southern Pacific between 30° S and 50° S from Chile to New Zealand; re- ported southwest of the Cape of Good Hope suggesting a bipolar distribution in the Atlantic; not reported from the North Pacific or southern Indian Ocean. Vertical distribution (Fig. 41): Data variable by day with relatively low con- centrations from 200 m to 750 m; by night depths are concentrated between 200 m and 450 m with most records from LSO m to 300 m; no indications of marked geo- graphic variation in depth. Geographic variation. Analysis of small sample sizes from the two major widely separated populations indicate no statisti- cal differences and little evidence of separation (Table 10; Fig. 42). Argyropelecus sladeni Regan Figure 43 Ar'^ijropelccus sladeni Regan, 1908: 218 (liolotype BMNH; Central Indian Ocean; not seen); Jespersen, 1934: 15; Fowler, 1936: 1208; Parr, 1937: 49 (fig., incorrectly cites Norman, 19.30 as original description); Norman, 1939: 19; Marine Hatchetfishes • Baird 57 45 15- 20 BO S L Figure 39. Geographic variation in the regression of body depth (BD) on standard length (SL) in A. aculeatus. A _ NW Atlantic; B = Gulf of Mexico; C = NE Atlantic; D = Caribbean; E = Pacific (Chile). Marr, 1948: 140; Misra, 1952: 367; Haig, 1955: 321; Fowler, 1956: 27; Koefoed, 1961: 1. \rgtiropdecus olfersi: Barnard, 1925: 153; Smith, 1957: 37 (?); Bright and Paquegnat, 1969: -29. Argtjropelccus Itjchntis hjchnus Schultz, 1961: 587 (in part); 1964: 241;" Blache, 1964: 71; Backus et al., 1965: 139; Bright and Paquegnat, 1969: 30. Argijwpelecus lycliniis sladcni Schultz, 1961: 587; 1964: 241 (incorrectly cites Norman, 1930, as original description); Kotthaus, 1967: 22 (photo., otoliths). Argyropelecus hjchnus hawaicnsis Schultz, 1961: 587; 1964: 241. Argyropclcciis hawaicnsis Berry and Perkins, 1965: 625; Lavenberg and Ebeling, 1967: 185. Species distinction. See A. olfersi (p. 52) and A. lychnus (ix 63) . Description. D. 9; A. 12; P. 10-11; total gill rakers 17-21; vertebrae 35-37. Medium size species seldom exceeding 60 mm SL; body less deep, depth at end of dorsal about two or more times into SL; dorsal blade low, height about three or more times into its length; postabdominal spines of equal size, anterior one occasion- ally straight, usually squared or blunted; upper preopercle spine long, directed posteriorly and usually dorsally, lower directed ventrally and often slightly posteriorly; jaws medium; teeth small, re- curved, no large canines present; gill rakers medium to long, slightly dentate; first pre- anal photophore raised well above second 58 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 E E o CN n o U Z < — 1 I— < > a. o Marine Hatchetfishes • Baird 59 Zoo. AOO. 600 • 800. 1000. 1500 B * N ¥■¥■ ¥ * 10 FISH HR 20 Figure 41. Diurnal vertical distribution of A. olfersi determined by rate of capture witfi depth. A (Pacific); B = NE Atlantic. 30 Soutfiern Ocean which is even with or above third; anal pterygiophore gap with three haemal spines lacking pterygiophores; in preserva- tive pigment often quite dark; large dis- tinct pigment spots present along midline, especially evident in smaller specimens; there may be a diurnal pigment difference similar to A. hemigymnus in this species. Distribution. Horizontal distribution (Fig. 44): In the Atlantic this species is found in abundance along the African coast from about 15 °S northward into the Gulf of Guinea; it occurs in moderate numbers across the equatorial Atlantic in a belt from 5°S to 15 °N latitude; it is abundant in the Caribbean in the vicinity of the Venezuelan coast, absent from the northern Caribbean, appearing again in numbers in the western and northern Gulf of Mexico and the straits of Florida; a few small catches have been taken in the North Atlantic and along the Brazilian coast. In 60 Bulletin Museum of Comparative Zoologtj, Vol. 142, No. 1 9 1 Table 11. Regression statistics for variousI populations of a. sladeni. 14 17 Figure 42. Geographic variation in gill raker count in A. oliersi. A := NE Atlantic; B z:^ Southern Ocean. Numbers refer to sample size. the Pacific a somewhat biantitropical dis- tribution is indicated, with large populations represented in the North Pacific to about 175°W longitude, and off the California coast; another large population occurs off the coast of Chile; the species occurs north of New Zealand and south of Hawaii. A. sladeni is abundant in the northern Indian Ocean to about 15 °N and along the African coast to about 10° S; while not reported from the Bay of Bengal, it is represented by several small catches south of Java. Vertical distribution ( Fig. 45 ) : Concen- trated between 350 m and 600 m by day, with the major concentrations between 350 m and 450 m; by night concentrated be- tween 100 m and 375 m, \\'ith the major concentrations between 100 m and 300 m; no marked indication of geographic vari- ation with depth. Geographic variation. This species, like A. fiigas, has low variability in those b()d\- Regression Character A B N Pacific (42°N, 165° W) Body depth 0.38 0.52 ± .156 Dorsal blade 2.02 0.01 ± .027 Jaw lengtli -0.03 0.25 ± .076 faw width 0.48 0.13 ± .070 N = 10 £ Pacific (CaHfoiiiia) Body depth 1.25 0.53 ± .074 Dorsal blade 2.62 0.00 ± .022 Jaw lengtli 0.92 0.23 ± .027 Jaw widdi 0.62 0.10 ±.028 N = 27 Indian Ocean (05°N, 65°E) Body depth 0.80 0.56 ± .142 Dorsal blade 1.89 0.01 ± .010 Jaw length 0.22 0.24 ± .069 Jaw width -0.73 0.13 ± .043 N = 11 Caribbean Body depth 0.87 0.52 ± .163 Dorsal blade 1.29 0.02 ± .013 Jaw length -0.07 0.26 ± .081 Jaw width 0.75 0.12 ± .044 N = 9 Gulf of Guinea Body depth -0.91 0.57 ± .110 Dorsal blade 1.89 0.01 ± .014 Jaw lengtli -0.33 0.27 ± .055 Jaw width 1.14 0.11 ± .027 N = 13 SE Pacific (Chile) Body depth 0.51 0.54 ± .097 Dorsal blade 2.04 0.01 ± .012 Jaw length 1.04 0.23 ± .040 Jaw width -0.63 0.14 ± .035 N = 16 Table 12. Comparisons between mean slopes of two ch.'^racters for all populations of A. SLADEM and a. LYCHNUS FOR WHICH REGRESSION STATISTICS WERE CALCULATED. PoP. # = NUMBER OF populations; x Slope = unweighted mean slope; Total # = total number of fish measured OVER all populations. Species Character Pop. # Slope Range Total # A. sladeni body depth 6 0.54 0.52-0.57 86 A. hjchnus body depth 3 0.61 0.57-0.64 38 A. sladeni jaw widtli 6 0.12 0.10-0.14 86 A. hjchnus jaw width 3 0.15 0.14-0.16 38 Marine Hatchetfishes • Baird 61 CN o ^0 u < X u c 01 o -2 ID a. o >s O) 62 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 o J3 E o O c O o X zl Marine Hatchetfishes • Baird 63 Zoo Aoo 600 800 1000 D iSOO E P T H D • • ••• n •k • it •k • • 0 0 10 20 30 2o • • • N 2oo 400 ■k D • * ^00 600 1000 1500 A • B D C O D • E O F if. F ISH H R 20 30 Figure 45. Diurnal vertical distribution of A. sladeni determined by rate of capture with depth during day (D) and night (N). A — Pacific (California); B = Pacific (Chile); C = Gulf of Guinea; D = Gulf of Mexico and Caribbean; E = N Central Pacific; F = N Atlantic. proportions measured. Overlap is broad and sample sizes are small. Six populations were statistically defined (Table 11; Fig. 46) but only gill raker counts gave much separation. Certainly the Atlantic popu- lation is distinct from the Indian Ocean and several Pacific populations; within the latter, distinctions are not marked. The Indian Ocean, Chile, and California popu- lations show some separation, although not statistically significant. Other characters and larger sample sizes are required to better define populations in this species. Argyropelecus lychnus Gorman Figure 47 Argyropelecus lycJmus Garman, 1899: 234 (lecto- type USNM 57885, designation Schultz, 1961; tropical east Pacific, not seen; paialectot>pe MCZ 35193, seen); Ledenfeld, 1905: 170 (light organs); Berry and Perkins, 1965: 625; Grand- perrin and Rivaton, 1966: 36; Lavenberg and Ebeling, 1967: 185. 64 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 A B C D E F 16 16 20 2Z Figure 46. Geographic variation in gill raker count in A. sladeni. A ^ Gulf of Guinea; B = Caribbean; C = Indian Ocean; D ^ N Central Pacific; E =: Pacific (Chile); F r= Pacific (California). Numbers refer to sample size. Argyropclccus olfersi: Weber and DeBeaiifort, 1913: 1 (?); Clemens and Wilby, 1949: 106; Koumans, 1953: 186 (?); Morrow, 1957: 56; Koepcke, 1962: 145; Bussing, 1965: 185. Argyropclccus lycJmus lychnus Sehultz, 1961: 587 (in part); 1964: 241." Argyropeleciis sp., Kotthaus, 1967: 11 (?) (photo.). Species distinction. See A. olfersi (p. 52); differs from A. sladeni by its higher dorsal lilade, preopercle spine character- istics, presence of two rather than three haemal spines in anal pterygiophore gap, lack of distinct dark pigment spots on midline, broader body, and generally lower gill raker count (Figs. 46 and 49). Tables 12 and 13 and Figure 50 illustrate the nature and degree of difference in several of the characters mentioned above. Description. D. 9; A. 12; P. 10-11; total gill rakers 16-18; vertebrae 35-37. Medium to large species often exceeding 60 mm SL; body deep, depth at end of dorsal greater than 1.5 into SL; dorsal blade high, height about 2.5 times into its length; postabdominal spines of about equal size, anterior one slightly smaller, not smoothly curving but blunted or squared; upper preopercle spine long, di- rected posteriorly and usually ventrally; lower spine usually curved slightly ante- riorly or straight down; jaws large, teeth recurved especially in lower jaw, no large canines; gill rakers medium to short, den- tate; first preanal photophore usually lower than third; spiny scales present in adults below subcaudal photophores; the gap made by the anal pterygiophores contains two haemal spines lacking pterygiophores; ill preservative, pigment dark dorsally, diffuse on trunk with small, light pigment spots on midline. Distribution. Horizontal distribution (Fig. 44): Absent from the Atlantic; represented possibly by a single sample from the Indian Ocean (04°S, 66°E, Kottliaus, 1967). Pri- Table 13. Slope compabisons of the regression of dorsal blade height on standard length for various populations of a. lychnus (l) and a. sladeni (s). Character Population 1 — L Population 2 — S T p Dorsal blade C Pacific Chile 3.179 .005 C Pacific N Pacific 2.452 .025 C Pacific Indian 2.904 .01 Chile Chile 3.903 .001 California Chile 3.965 .001 Chile N Pacific 2.171 .045 Chile California 3.514 .001 Chile Indian 3.366 .005 California N Pacific 2.272 .035 California California 3.355 .005 California Indian 3.444 .005 Marine Hatchetfishes • Baird 65 E E o O If) o CO CO (U u X O X J) "S Q. O O) N. ■^ 3 66 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 H o N Zoo. ^00. o o • . * * • * * * 6oo. 8oo. 1000. 1500 BO C • fc 10 F ISH H R 20 30 Figure 48. Diurnal vertical distribution of A. lychnus determined by rate of capture with deptfi during the day (D) and night (N). A = Pacific (California); B = Pacific (Chile); C = Tropical E Pacific. marily restricted to the tropical Pacific; found in abundance in the eastern Pacific between 35°N and 35°S; distribution nar- rows across the equatorial Pacific as far as 160^W; a moderate catch from tlic lesser Sunda Islands indicates a possible trans- equatorial distribution in the Pacific. Vertical distribution (Fig. 48): Concen- trated between 300 m and 400 m off California by day, with the highest con- centration near 400 m; by night major concentrations occur from 200 m to 350 m, with no marked indication of geographical variation in depth. GcQfiraphic variation. Three samples from widely separated areas in the tropical east Pacific and its northern and southern boundaries gave no indication of any sig- nificant variation (Table 14; Fig. 49). Horizontal distribution data indicates an essentiallv continuous distribution in this area. Marine Hatchetfishes • Baird 67 A B C D Table 14. Regression statistics for various populations of a. lynchus. Regression \h 18 Figure 49. Geographic variation in gill raker count in A. lychnus. A = Central Pacific; B = Tropical E Pacific; C ^ Pacific (Chile); D = Pacific (California). Numbers refer to sample size. Genus Sfernoptyx Hermann, 1781 SteiiiOjityx Hermann, 1781: 8 (type species: Stc'inoptyx dicijilxana Hermann, 1781, by mono- typy). Diagnosis. Ten abdominal, three anal, three branchiostegal and five isthmus photophores; a single large dorsal pterygio- phore spine with an anterior, serrated ex- tension; first anal pterygiophore greatly enlarged, forms support for triangulate membrane above anal fin rays; premaxilla without anterior pedicels; anteriormost gill rakers reduced to toothed ridges; post- temporal and supracleithrum separate; hy- pural elements fused to form single caudal plate; haemal and neural spines greatly elongate in trunk region. Description. Photophores: PO 1; PTO 1; PRO 1; SO 1; SP 3; PAN 3; SAN 1; AN 3; SC 4. Spines: Preopercle with single ventrally oriented spine; retroarticular bears spine, preabdominal spine present; basipterygia fused to form a set of four postabdominal spines; base of first anal pterygiophore bears ventral spines; no well-developed post-temporal spines. Character SE Tacific (Chile) Body depth Dorsal blade Jaw length Jaw width E Pacific (California) Body depth Dorsal blade Jaw length Jaw width 2.38 1.56 0.84 -0.46 0.94 1.53 -0.15 -1.02 Central Pacific (10°N, 145°W) Body depth 1.42 Dorsal blade 1.79 Jaw length 0.88 Jaw width -0.77 0.57: 0.05: 0.26: 0.16 N: 0.61 0.04 0.29 0.16 N 0.64 0.05 0.28 0.14 N .120 .018 .055 .043 12 .107 .013 .058 .031 15 .156 .023 .068 .052 11 Eyes: Large, well developed, nontele- scopic. Gill rakers: Total seven to nine; well developed, \\'ith rough spiny margins; an- teriormost rakers reduced to spiny tooth- like plates extending into mouth cavity. Jaws and dentition: Jaws vertically oriented, premaxilla small, heavily toothed; maxilla heavily toothed and major upper jaw bone in gape; lower jaw heavily toothed, teeth small, sharp, triangulate; palatine teeth present; first epibranchial extended anteriorly and ventrally forming toothed arms at dorsal, posterior end of mouth. Meristics: Vertebrae 28-3 J; C. 9+10; D. 8-11; A. 14-16. Color: Bright silvery in life, dark pig- ment especially evident on dorsal surface; 68 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 30 B D 20 3 O 60 S L Figure 50. Regression of body depth (BD) on standard length (SL) In A. sladeni (A) and A. lychnus (B) off California. .silver (pickly lost in formalin preservative; sides can be very dark, often black. Internal anatomy: Air bladder and gas gland well developed and fills much of the body cavity (see Marshall, 1960); digestive system simple, consisting of a heavily pigmented, single sectioned stom- ach, capable of considerable distension, five to seven pyloric caecae of which only two to three are long and well developed, relatively small liver and short, uncoiled, thin-walled intestine. The gonads, when mature, lie against the posterior wall of the body cavity. This cavity is large, unlined with pigment, and appears capable of some expansion. Key to the Species of Sternoptyx la. Dorsal long, its lengtli greater than 1.3 times height of dorsal spine; trunk long and narrow, SL more than 3.0 times l^ody depth at end of dorsal (see regression, body depth. Table 15); body very dark, pigment forms broad band at base of caudal rays S. ohscura ( p. 69 ) . b. Dorsal short, its length less or equal to height of dorsal spine; trunk broad, SL less than 2.8 times body deptli at end of dorsal (see regression, body depth, Tal:)les 16 and 17); body pigment less uniformly dark, pigment absent or in very narrow band at base of caudal rays 2 2a. Supra-anal photophore high, its height greater than one-half the distance from ventral body margin to midline ( often raised to midhne ) ; gill raker tooth plates with prominent spines; secondary anal pterygiophores long, e.xtending posteriorly on same level as anal photophores _.._ S. pseudobscura (p. 72). b. Supra-anal photophore low, its height less than one-half distance from ventral body margin to midline; gill raker tooth plates lacking prominent spines; secondary anal pterygiophores short; not extending posteriorly on same level as anal photophores S. duiphana (p. 75 ). Marine Hatchetfishes • Baud 69 a. Sfernopfyx obscura Garman Figure 51 Sternoptyx obscura Garman, 1899: 63 (lectntype USNM 177888; designation Schultz, 1961; tropical east Pacific; not seen; paralectotype MCZ 28532; seen); Ledenfeld, 1905: 170 (light organs); Follett, 1952: 409. Sternoptyx diaphana Schultz, 1961: 587 (in part); 1964: 241 (in part); Berry and Perkins, 1965: 625 (in part). Species distinction. Differs from both S. diaphana and S. pseudohscura in its shorter dorsal spine and longer dorsal fin; longer, narrower trunk, slight extension of body Figure 51. Sfernopfyx obscuro; R/V ANTON BRUUN, Cruise 3; Station 215; SL 30 mm. 70 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 margin in front of anal photophorcs, broad pigment band on base of caudal rays, and generally dark pigment; differs from S. pseiidohsciira in its lo\\'er supra-anal photo- phore, small teeth, low gill raker tooth plates, and smaller mouth; from S. dia- phana in its extension of the ventral body margin at same level behind anal photo- phores. Tables 18 and 19, and Figures 5.3, 56, and 61 illustrate the degree of differ- ence between the three species in several of the above characters. Note especially the significant differences between sym- patric populations. Description. D. 10-11; A. 14-15; P. 10- 11; total gill rakers 7-9; vertebrae 29 (30). Small species, seldom exceeds 40 mm SL; trunk long and narrow, its length usually longer than depth; dorsal fin long, its length more than 1.3 times the length of dorsal spine; abdominal length along midline from supra-anal photophore to caudal peduncle, less than or equal to body depth at end of dorsal; postabdominal and anal pterygiophore spines long; posterior anal pterygiophores extend behind and at same level \\\\\\ anal photophore group; supra-anal photophore raised above anals one half or less the distance to midline; body margin extends slightly in front of anal photophores before curving ventrally; jaws medium; teeth small; gill raker tooth plates consist of multiple low spiny ridges; anterior dorsal surface of tongue between branchial arches smooth; few-to-no raised nodules; in preservative, pigment very dark over whole of body; pigment extends in broad band at base of caudal fin rays. Distribution. Horizontal distribution (Fig. 52): Tliis species has not been recorded from the Atlantic; it is concentrated north of 10°S latitude in the Indian Ocean, al- though small catches occur as far as 40°S; occurs in the eastern Bay of Bengal, and abundantly south of Java; occurs off the Philippines and scattered but large catches indicate in all probability a continuous distribution across the equatorial Pacific; occurs abundantly in the tropical east Table 15. Regression statistics for various populations of s. obscura. Regression Character A B Indian Ocean (5°N, 60°E) Body depth -0.35 0.31 ± .051 Abdominal length -1.16 0.40 ± .074 N = 23 Indian Ocean (3°N, 67°E) Body dcpdi 0.45 0.29 ± .133 Jaw length 2.41 0.09 ± .067 N= 10 Java (10°S, 114°E) Body depth 0.98 0.27 ± .071 Abdominal length 0.68 0.31 ± .090 N = 25 Central Pacific (11°N, 163°E) Body length -1.72 0.37 ± .067 Abdominal length 0.12 0.37 ± .074 Jaw length 0.85 0.14 ± .028 N = 20 Central Pacific (7°S, 135°W) Body depth -0.71 0.34 ± .104 Abdominal lengtli 0.71 0.35 ± .141 Jaw length 0.10 0.15 ± .057 N = 11 East Pacific (California) Body depth -0.94 0.33 ± .075 Abdominal length -1.23 0.42 ± .091 Jaw length 1.85 0.12 ± .034 N = 15 Pacific from California to the Chile-Peru border. Vertical distribution: Depth data is spotty and no depth rate plot \\'as made, however, data (Appendix B) indicates a depth range of 650 m to at least 1000 m; tropical Pacific maximum net depth figures concur in general with this range. Geographic variation. Analysis of catches from many widely scattered areas in the Pacific and Indian Ocean gave no indi- ^ cation of population variation (Table 15, | Fig. 53). This, coupled with horizontal distribution data, indicates a probable single trans-Indo-Pacific population. Marine Hatchetfishes • Boird 71 'pe is hereby designated: Neo- type MCZ 46402; 11° 06'N, 78° 2rW; 8/7/66; R/V ANTON BRUUN, Cruise 19; Station 813. Cuvier and Valenciennes, 1849: 415; Giinther, 1864: 387; Goode and Bean, 1896: 127; Al- cock, 1896: 331; Gilbert, 1905: 601; Brauer, 1906: 69 (in part); 1908: 175 (eye muscles); Holt and Byrne, 1913: 20; Weber and DeBeau- fort, 1913: 1; Jespersen, 1915: 12; Jespersen and Tiining, 1919: 220 (eye); Borochn, 1931: 68; Jespersen, 1934: 15; Roxas, 1934: 287; Buen, 1935: 52; Fowler, 1936: 1208; Beebe, 1937: 22; Parr, 1937: 49; Norman, 1937: 82; 1939: 19; Nybelin, 1948: 25; Maul, 1949a: 17; 1949b: 13 (in part); Wilimovsky, 1951; Misra, 1952: 367; Koumans, 1953: 186; Mead and Taylor, 1953: 570; Smith, 1953: 102; Haig, 1955: 321; Rass, 1955: 328; Grey, 1959: 326; Koefoed, 1961: 11; Schultz, 1961: 617 (in part); 1964: 241 (in part); Backus et al., 1965: 139 (in part); Berry and Perkins, 1965: 682 (in part); Bussing, 1965: 185; Haedrich and Nielsen, 1966: 909; Bright and Paquegnat, 1969: 34. Species distinction. See S. obscnra (p. 69) and S. pseudobscura (p. 72). Description. D. 9-11; A. 14-16; P. 10- 11; total gill rakers 7-8 (9); vertebrae 29 (30). _ Medium size species, seldom exceeds 55 mm SL; trunk very broad; its depth usually greater than its length; dorsal spine long, its length greater or equal to dorsal fin length; posterior anal pterygiophores short, little extension behind and on same level with anal photophores; supra-anal photo- phore low, not reaching more tlian one-half the distance from ventral body margin to midline, no body margin extension in front of anal photophores; jaws medium to small; teeth short and low; gill raker tooth plates with low spinate ridges; anterior dorsal ID P a o o SI o o -Q_ O O o * o A • B* CO 3 O S L 5 Q ■igure 57. Geographic variation in distance from dorsal body margin of supra-anal photophore (PO) with standard length SL) in S. pseudobscura. A =: Indian Ocean; B =Gulf of Guinea; C = Gulf of Mexico and Caribbean. 76 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Figure 58. Sternoptyx diaphana; GALATHEA; Station 494; 33 mm. surface of tongue with small nodules; post- abdominal and anal pterygiophore spines usually shorter than others in genus; in preservative pigment dark dorsally, often light and dispersed in trunk region, usually little pigment present at base of caudal rays. Neotype: measurements (mm): SL 28.4, BD 12.1, JL 05.3, CP 03.5, Ab. length 09.0; meristics: GR 7, D 10, A 15, anal photo- phores 3. Distribution. Horizontal distribution (Fig. 59): Broadly distributed in the Atlantic, caught in moderate numbers in the South Atlantic off Brazil and from 20°W to the African coast at about 35^ S; abundanth' present in the Gulf of Guinea and the tropical Atlantic; taken abundantly in the southern Caribbean, the Gulf of Mexico,! and straits of Florida; taken in the western North Atlantic; a large population occurs in the northeastern Atlantic from 25°N to 45°N latitude. In the western Indian Ocean small to moderate catches extend from 5°S to 35 °S latitude, a single catch has been observed from the eastern Indian Ocean; numerous catches indicate this species present south of Java, near Borneo, and in the Banda Sea; known also between New Guinea and the Solomon Islands, it occurs in the westem Pacific near the Philippines and along the coast of Japan, with a small I Marine Hatchetfishes • Baird 77 75 JQ E o a U o c o -c a o c O o X 78 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 AC B ••< C •• D C F ISH HR Figure 60. Diurnal vertical distribution of S. diaphana determined by rate of capture with depth during day (D) an night (N). A ^ Southern Ocean; B ^ Gulf of Guinea; C =: Gulf of Mexico and Caribbean; D =: NE Atlantic. sample taken in the North Pacific; this species occurs in small numbers off lower California and has been reported abun- dantly southeast of Hawaii; a large popu- lation extends across the South Pacific from Chile to about 160° E longitude. Vertical distribution (Fig. 60): Distrib- uted between 400 m and 1200 m; major concentrations occur between 700 m and 900 m; no diurnal movement or marked geographical depth xariation could be dei tected. Geographic variation. With the excep tion of the Pacific southern ocean popm lation, little geographic variation could b* detected; characters were few and sampl( sizes small, however (Table 17, Fig. 61); The Southern Ocean population is quit«| distinct and certainly represents an ini stance of incipient speciation. Tables 1' Marine Hatchetfishes • Baud 'able 17. Regression statistics for various populations of s. diaphana. Rf jgression Character A B laribbcan aw length -0.24 0.18 ± .066 hotophore -2.04 0.37 ± .101 N = ll ava (10°S, 114°E) lody depth -2.93 0.54 ± .107 ibdominal length 0.78 0.31 ± .056 N = 22 ndian Ocean (5°-40°S, 55° -65°E) lody depth 0.10 0.51 ± .095 Lbdoniinal length 1.18 0.31 ± .093 hotophore 0.20 0.29 ± .085 aw length 1.00 0.15 ± .088 N = 12 oiithern Ocean lody depth -0.77 0.42 ± .044 abdominal length 1.18 0.31 ± 093 aw length 1.47 0.16 ± .041 hotophore 2.23 0.11 ± .033 N = 40 outhern Ocean (Chile) lody depth -1.50 0.41 ± .113 aw length 1.46 0.14 ± .037 N=ll nd IS indicate the degree of difference letween this population and others in the pedes. In body depth it falls somewhat »et\veen most populations of S. diaphana nd S. ohscura (Table 19). Phenotypically t has supra-anal photophore characteristics resembling some populations of S. pseud- ohscura. In most characters it falls closest to other populations of S. diaphana, espe- cially in mouth and gill raker character- istics. Considering the lack of sympatry with other forms and the degree of distinctness between the species, it is presently con- sidered to represent a distinct form of S. diaphana. Genus Polyipnus Gijnther, 1887 Polyipnus Ciinther, 1887: 170 (type species: Polyipnus spinosus Giintlier, 1887, by mono- typy). Diagnosis. Ten abdominal, three supra- abdominal, and a lateral photophore; post- temporal spine(s) well developed; a fused double dorsal pterygiophore forms short spines anterior to dorsal fin rays; cleithrum projects below pectoral fin forming fanlike, spine-bearing extension posteriorly; otoliths very large with characteristic armlike ex- tension (see Kotthaus, 1967); three to four hypural elements in upper caudal lobe; lower jaw noticeably expanded dorsally. Description. Photophores: PO 1; PTO 1; BR 6; I 6; PRO 1; SO 1; SP 3; SAB 3; AB 10; L 1; PAN 5; AN 6-14; SC 4. Spines: Post-temporal extends posteriorly to form from one to three prominent spines; preopercle spined, the lateral sur- face often bearing spiny elements; retro- articular spined; ventral surface of lower jaw often serrate; cleithrum bears pre- abdominal spine; bony keel scales often bear spines ventrally; four postabdominal ABLE 18. Slope comparisons between regressions of sever,\l characters in species of StERNOPTYX. D = S. DIAPHANA; O = S. OBSCURA; P = S. PSEUDOBSCURA. Character Population 1 Population 2 T P 5ody depth Java D Central Pacific P Java D Java O Central Pacific O Southern Ocean D 2.780 2.362 2.611 .001 .025 .010 Uadominal Icngdi Java D Southern Ocean D 2.381 .023 'hotophore Caribbean P Gulf of Guinea P Florida P Caribbean D Caribbean D Caribbean D 4.977 3.240 2.767 .001 .005 .018 80 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 IS A B C □ 38 rin 22 10 6 — r- 8 Figure 61. Geographic variation in gill raker count in S. diaphana. A = Caribbean; B = Indian Ocean; C =^ Southern Ocean; D = Pacific (Chile); E = Indian Ocean — - S of Bali; F z= Java Sea. Numbers refer to sample size. spines; first anal pterygiophore may extend ventrally to form small spines. Eyes: Large, well developed, nontele- scopic, essentially laterally oriented. Gill rakers: Number 10-28; well de- veloped, long, often quite close together; first branchial arch considerably larger than succeeding arches; the inner surfaces of second and thir'd cerato- and epi- branchials bear tooth plates. Jaws and dentition: Mouth small and vertically oriented; premaxilla long, toothed, and major upper jaw bone in gape; arm of first supramaxilla elongate; dorsal margin of lower jaw greatly ex- panded which, with broadening in the meso-and metapteryoid, make the mouth cavity a long conelike basket with a sub- stantial distance between the mouth en- trance and the beginning of the branchial arches; the maxilla is toothed, but essen- tially excluded from gape; teeth small to minute, no canines, vomer and palatines bear teeth. Meristics: Vertebrae 31-36; C. 9+10; D. 10-17; A. 13-19. Color: Bright silvery in life; dark dorsal pigment band often extends ventrally and may reach lateral midline; dark pigment may form lateral striated bands on pos- terior trunk. Internal anatomy: Swim bladder and associated gland well developed, gland quite large, with grainy appearance; l)lad- der thick walled, and often heavily in- vested with fatty tissue (see Marshall, 1960); digestive system simple with bi- partate stomach, anterior section thicl< walled, the lining often raised into heavy Table 19. Comparisons between mean slopes of several characters among the species or Sternoptyx. S.O. = Southern Ocean populations of S. diaphana; Pop. # = number of popula- tions; X Slope = unweighted mean slope; Total # = total number of fish measured over ali populations. Species Character Pop. # X Slope Range Total # S. obscure body depth 6 0.32 0.27-0.37 104 S. diaphana body depth 2 0.525 0.51-0.54 34 S. diaphana ( S.O. ) body depth 2 0.415 0.41-0.42 51 S. pseudob.scura body depdi 1 0.49 0.49 13 S. diaphana photophore 4 0.31 0.29-0.37 35 S. diaphana (S.O.) photopliore 1 0.11 0.11 40 S. pseudohscura photophore 4 0.15 0.11-0.18 33 S. obscura jaw length 4 0.13 0.09-0.15 56 S. diaphana jaw length 5 0.196 0.15-0.23 41 S. diaphana (S.O.) jaw length 2 0.15 0.14-0.16 51 S. pseudohscura jaw length 5 0.15 0.15-0.16 46 Marine Hatchetfishes • Baud 81 30 SL so Figure 62. Regression of lower jaw length (JL) on standard length (SL) in S. diaphana (A) and S. pseudobscura (B) from the Tropical Atlantic. ridged folds; die posterior section diin walled and extensible, six to ten pyloric caecae, short straight intestine, and a rel- atively large well-developed liver. Gonads, when mature, lie horizontally and laterally in the body cavity; cavity lined with pig- mented membrane; nepliritic tissue is not as well developed as in other genera. Species complexes: As with Arfiyropele- ciis, there has been considerable radiation within the genus. There are three distinct species complexes, two closely related. The third complex, P. spinosiis, is quite distinct. The latter appears more primiti\e in terms of axial and caudal skeleton characteristics. The P. spinosws complex differs from the other two groups as follows: greater de- velopment of the post-temporal spine complex; otoliths (Weitzman, personal con- versation); four hypural elements in the upper caudal lobe; serrate lower jaw mar- gin; spine-bearing abdominal keel plates; the second and tliird pretenninal neural spines wedge shaped; extension of the cleithrum below the pectoral relati\ely more broadened; and the anal pterygio- phore gap is reduced. Within the P. spinosus complex there is a further dichotomy. P. nuttingi, P. oJiiohis, and P. indicus with peculiar reductions in post-temporal spine characteristics, fomi one group; P. spinosus, P. sterope, and P. tridentifer with a well-developed, post- temporal complex, fonn the other. The P. asteroides and P. htenwtus spe- cies complexes are more closely related. They differ primarily in post-temporal spine characteristics, body shape and size, photophore number and pattern, and den- tition. Both complexes have similar otoliths and resemble each other osteologically. The species P. later natiis — P. omphiis 82 BuUctin Museum of Comparative Zoology, Vol 142, No. 1 and P. iinispinus — P. aquavit us form a dichotomy within the P. latenuitus complex. The differences inckide preopercle spine length, body shape, photophore pattern, and some meristic differences (gill rakers, vertebral nnmber). P. astcroides — P. poUi — P. triphanos and P. matsu])ami — P. meteori — P. kiiciensis — P. rugg,eri form a similar dichotomy within the P. asteroides complex. The latter group is characterized by an extremely long, narrow, posterior vomerine shaft, closely allied and fitting into the parasphenoid. This shaft bears teeth anteriorly, in addi- tion to the normal lateral vomerine teeth. Other minor differences are also joresent. Key to the Species of Polyipnus la. Post-temporal spine complex, bearing one or two basal snpplementary spines; dorso-lateral edge of fused, post-temporal-supracleithrum serrate; abdominal keel scales with spiny ventral surfaces; supra-anal photophore group usually not distinctly separated from anal group P- spinosus complex 3 1). Post-temporal spine simple, bearing no basal spines; lateral edge of post-temporal-supra- cleithrum smooth; keel scales with smooth ventral surfaces (except P. imispinus); supra- anal photophores separate, usually raised well above anal group ._ __ — _ 2 a. b. 2a. Post-temporal spine long, its length greater than one-fourth the diameter of orbit; anal photophore number 10 to 13 (adults only); body long and narrow, SL greater than 1.7 times body depth; first supra-anal photophore even with or raised above the second (except P. late maim in which the first is slightly lower than die second) P. laternalus complex 8 b. Post-temporal spine short, its length less than one-fourth the diameter of orliit; anal photophore number 7 to 9 (occasionally 10); body more robust, SL less than 1.9 times body depth; first supra-anal photophore markedly lower than second - P. asteroides complex 1 1 1 ^ A a. Marine Hatchetfishes • Baird 83 3a. Post-temporal spine with two distinct basal spines; anal-subcaiidal photophore distance less than one-third the length of the subcaudal group; anal photophore number 10 to 13 4 b. Post-temporal spine with a single distinct basal spine (this reduced in P. oluohis); anal- sul)caudal distance greater than one-half of the length of the subcaudal group; anal photopbore number 6 to 9 6 4a. Anal photophore number 10; SL less than 3.6 times body depth at end of dorsal fin; caudal peduncle broad, head length less than 2.8 times narrowest peduncle depth 5 b. Anal photophore number 12 to 13; SL greater than four times body depth at end of dorsal; caudal peduncle narrow, head length greater than three times narrowest peduncle depth P. tridentifer (p. 86). 5a. Post-temporal basal spines well developed, ventralmost basal spine lengtli greater than one-half length of post-temporal spine; gill raker numlaer 24-28 P. sterope (p. 88). b. Post-temporal basal spines short, ventralmost basal spine less than one-fourth post- temporal spine length; total gill raker nmnber IS to 21 P. spinosus (p. 89). a. b. 6a. Alidominal keel scales triangulate, with one or two large ventral spines; iDost-temporal spine long, heavily spinose dorsally and laterally; first supra-anal photophore markedly lower than third; anal photophores 6 to 7 P. oluohis (p. 90). b. Abdominal keel scales rectangular, with many small ventral spines; post-temporal spine long and smootli or short and spinose dorsally only; first supra-anal photophore about even with or raised above tliird; anal photophores 8 to 9 ._ 7 a. 7a. Ventral margin of subcaudal photophores with spines (adults); anal-subcaudal photo- phore distance less than three-fourths length of subcaudal group; first supra-anal photophore higher than the last; post-temporal spine long, greater than one-half diameter of orbit P. iudiciis (p. 91). b. Ventral margin of subcaudal photophores smooth; anal-subcaudal distance greater tlian three-fourths length of subcaudal group; first supra-anal photophore lower than last; post-temporal spine short, less than one-half diameter of orbit .. . P. nuttingi (p. 92). 84 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 8a. Gill rakers 18 to 21; supra-anal photophores distinctly raised above tlie anal group; preopercle spine short, somewhat triangulate 9 b. Gill rakers 12 to 14; supra-anal photophores not markedly raised above anal group; preopercle spine long and slender - — ....10 '^^ Aap\ m=^ ^^Peep^ 9a. Sulicaudal photophores compact, length of subcaudal group less tlian narrowest caudal peduncle depth; distance from top of last supra-anal photophore to top of first anal greater than three-fourths length of preanal group P. latcrnatus (p. 92). b. Subcaudal photophores somewhat spread; length of subcaudal group equal or greater than narrowest peduncle depth; distance from top of last supra-anal photophore to top of first anal less than one-half preanal length P. omphus (p. 94). 10a. Posterior lateral margin of preopercle smooth; xentral keel scales smooth; dorsal .spine short, length less than one-fourth diameter of orbit P. aquavitiis (p. 96). b. Posterior lateral margin of preopercle serrate; ventral edge of keel scales with small spinelets; dorsal spine long, length greater than one-half diameter of orbit — . P. iini.spinus (p. 97). 11a. Teeth absent on posterior vomerine shaft; second supra-anal photophore e\en with or raised above third; supra-abdominal photophores nonsymmetrical, first photophore raised well above other two 12 b. Teeth present on posterior vomerine shaft; second supra-anal photophore lower than third; supra-abdominal photophores symmetrical, first not markedly raised above third 14 Marine Hatchetfishes Baird 85 f\ ^ ft ,^ f\ flA a. %(\ 12a. Gill rakers 16 to 18; dorsal rays 11 to 12; lower margin of dark dorsal pigment markedly raised above supra-anal photophores P. triphanos (p. 97). b. Gill rakers 20 to 24; dorsal rays 14 to 16; lower margin of dark dorsal pigment straight from dorsal spine to caudal peduncle , 13 13a. Anal pterv'giophores form circular margin below anal photophores; anal photophores number 7 to 8; SL less than 3.5 times body depth at end of dorsal P. polli (p. 98). b. Anal pterygiophore margin essentially straight; anal photophore nimiber 9 (rarely 10); SL greater than 3.7 times body deptli at end of dorsal P. asteroides (p. 99). 86 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 14a. Gill rakers 22 to 24; dark pigment bar extending to midline very narrow, its width less tlian greatest widtli of lateral photophore (see illustration p. 101) P. matsuharai ( p. 101 ) . b. Gill rakers 13 to 18; dark pigment bar absent or much wider than width of lateral photophore 15 15a. Dark pigment bar greatly reduced or absent; supra-abdominal photophores essentially in straight line (see illustration p. 102); gill rakers 18 P. ruggeri (p. 102). b. Dark pigment bar present, extending to midline; supra-abdominal photophores tri- angulate, the second markedly lower than the otlier two; gill rakers 1.3 to 17 16 16a. Gill rakers 13 to 15; light stripe behind dark pigment bar extends to mid-dorsal line; ventral border of dark dorsal pigment markedly raised above supra-anal photophores (taken from photo and description, Kotthaus, 1967) P. mctcori (p. 104). b. Gill rakers 16 to 17; light stripe behind dark pigment bar not extending to mid-dorsal line; ventral border of dark dorsal pigment not markedly raised above supra-anal photophores (see illustration p. 103) P. kiwiemis (p. 103). PoJyipnus tridentifer McCulloch Figure 64 Pohjipnus tridentifer McCulloch, 1914: 78 (lecto- type AM E.3.543; designation Schultz, 1961; Australian Bight; not seen); Schult/., 1961: 619; 1964: 247. Pohjipnus spinostis: Weber and DeBeaufort, 1913: 1; Matsubara, 1950: 192; Okada and Suzuki, 1956: 297; Suzuki, 1964: 1. Pohjipnus frazeri Fowler, 1933: 2.57; Schultz, 1961: 620. Species distinction. Differs from P. spinostis and P. sieropc in its long, narrow trunk and caudal peduncle; long, smooth post-temporal spine; more sharply angled dorsal spine; multispinose subcaudal scales; less spinose abdominal keel scales; differs from P. spinosiis by its much longer third basal post-temporal spine. Description. D. 13-14; A. 15-17; P. (12) 13-14; total gill rakers (20) 21-24; verte- brae 33-34. ' Medium size species, not often exceed- ing 60 mm SL; trunk tapering into long, narrow, caudal peduncle; its depth less or equal to length of subcaudal photophore group; post-temporal spine long, its length more than one-half the distance from its base to point of dorsal spine; second basal i post-temporal spine long; dorsal surface of post-temporal spine relatively smooth, lack- ing marked serrations; supra-abdominal photophores arranged in steplike fashion, each raised an approximate ec^ual distance, above the next; abdominal keel scales spinose, although spines very short; scales below subcaudal photopliores with several prominent spincvs; preoperclc spine di- Marine Hatchetfishes • Baird 87 I® a! .- E Q." a t/i o E o ;5 o c n ^ D- O 3 Q. 5^ a. o c t/> a 3 -cr t- 0 0) O) 0) o' ■ , (•) o c ^- () (11 3 c- 0) ^ D ^ _i- 0) Q.' 1— CO • O VI ri o (11 "^ 3 o Cl, cn a to ^ 88 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 -,-%r.::-. Figure 64. PoJyipnus tridentifer; after Matsubara, 1950. rected, at its base, posteriorly, curving distally to point ventrally or slightly an- teriorly; jaws medium; teeth minute; gill rakers long; pigment in preservative dark dorsally with narrow, dark bar extending toward midline; pigment diffuse on trunk. Distribution (Fig. 63). Restricted to the western Pacific, taken abimdantly around the Philippines, off the south China coast, off Japan, in waters north of the Strait of Malacca, and in the Great Australian Bight. Polyipnus sferope Jordan and Storks Figure 65 PoJyipnus stcropc Jordan and Starks, 1904: 581 (holotype USNM 51451; Sagami Bay, Japan; seen); Matsubara, 1941: 2; Haneda, 1952: 12 (light organs); Okada and Suzuki, 1956: 297; Suzuki, 1964: 1 (X-ray). Polyipnus spinosus: Kamohara, 1952: 17. Poli/ipnus spinosus sterope Schultz, 1961: 621; 1964: 247. Species distinction. See P. triclcntifer Figure 65. Polyipnus sterope; modified from Jordan and Starks, 1904. Marine Hatchetfishes • Baud 89 igure 66. Po/yipnus sp/nosus; modified from Gijnther, 1887. (p. (S6); differs from P. spinosus in its onger basal post-temporal spines; shorter oreopercle spine; more raised first supra- ibdominal photophore, somewhat shorter 30st-temporal spine in relation to its base- :o-dorsal spine length, and higher gill •aker count. Description. D. 13-14; A. 15-17; P. 13- L5; total gill rakers (23) 24-28; vertebrae 33-34. Medium to large species, seldom ex- ceeding 70 mm SL; trunk broadly tapering; :audal peduncle broad, its depth more :han length of subcaudal photophore group; [lost-temporal spine spinose dorsally, its length substantially less than one-half dis- tance from its base to point of dorsal spine; third basal post-temporal spine long, sec- ond basal spine prominent; dorsal spipe bigh, with flangelike anterior portion not rising sharply from dorsal surface; supra- ibdominal photophores positioned in a itep-wise arrangement, \\'ith first photo- phore raised above other two; abdominal keel scales very spinose, including those ventral to preanal photophores; subcaudal scales either smooth or with single short spine; jaws medium; gill rakers long; pre- opercle spine curves slightly anteriorly; pigment in preservative dark dorsally with very narrow bar extending toward midline; pigment diffuse on trunk. Distribution (Fig. 63). Known only from the waters around Japan, where it has been taken less abundantly than P. tridentifer. Polyipnus spinosus GUnther Figure 66 Polyipnus spinosus Giinther, 1887: 170 (holotype BMNH, East Indies; not seisn); Alcock, 1896: 331; 1899: 135; Brauer, 1906: 69 (larvae, fig.) (in part); 1908: 175 (eye muscles); Roxas, 1934: 287; Misra, 1952: 367; Koumans, 1953: 186 (?); Samuel, 1963: 101 (?). Polyipnus spinosus spinosus Schultz, 1961: 624; 1964: 247. Species distinction. See P. tridentifer (p. S6) and P. sterope (p. 88). Description. D. 13-14; A. 15-17; P. 13- 15; total gill rakers 18-21; vertebrae 33-34. Medium to small species, seldom exceed- ing 70 mm SL; trunk and caudal peduncle broad, its depth greater than or equal to length of subcaudal photophore group; post-temporal spine spinose dorsally, its basal spines reduced; post-temporal spine nearly equal to one-half the distance from its base to dorsal spine; dorsal spine similar to P. sterope; supra-abdominal photophore positioned with first photophore only slightly raised above other two; first two supra-anal photophores slightly raised from third; preopercle spine long, curving an- teriorly; abdominal and preanal keel scales 90 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Figure 67. Polyipnus o/uo/us; R/V HUGH M. SMITH, Cruise 37; Station 43; SL 33 mm. spinosc; siibcaudal scales with no spines to a single small spine; jaws medium; gill rakers long; pigment in preservative dark dorsally, with narrow dark bar extending toward midline; pigment less dark above anal photophores. Distribution (Fig. 63). Taken in num- bers off the Philippines and off the south China coast; reported from peninsular India, although these reports may repre- sent P. tridentifer. Polyipnus o/uo/us n. sp. Figure 67 Holotvpe BCFH 2562; 11° 18'N, 162° 06'E; 12/9/56; R/V HUGH M. SMITH, ciaiise 37; Station 43. Species distinction. Differs from P. indi- cus and P. nutfingi in its much broader body; post-temporal spine characteristics; triangular abdominal keel scales; supra- anal, supra-abdominal, and subeaudal photophore characteristics; and posterior extension of dorsal fin rays to end of anal photophores. Description. D. 14; A. 15; P. 13; total gill rakers 19; vertebrae 33. Known only from holotype, 33 mm SL; body very broad, narrowing abniptly to short narrow caudal peduncle; body depth 1.3 times into SL; post-temporal spine length more than one-half diameter of orbit, extends to origin of dorsal spine, very spinose dorsally and laterally; frontal ridges almost vertical, spinose; postabdomi- nal spines well developed; abdominal keel scales extend well beyond ventral body margin; these scales sharply triangulate, coming to a single or double point ven- trally; subeaudal scales smooth; first supra- abdominal photophore raised considerably above other two; subeaudal photophores raised well above anals; first supra-anal photophore noticeably lower than second Marine Hatchetfishes • Baird 91 -igure 68. Polyipnus indicus; after Schultz, 1961. which is lower than third; anal photophores in two distinct groups; jaws large; teeth small, several recurved ones in upper jaw; vomerine teeth well developed; gill rakers medium, spinose; in preservative pigment somewhat darker dorsally; pigment stri- lations present on tiimk. I Holotype: measurements (mm): SL 32.9, |BD 25.7, JL 08.3, CP 03.9; meristics: GR jl9, D 14, A 15, anal photophores 7; name: from the Hawaiian "oluolu," which means I happy. Distribution (Fig. 63). Known only from a single capture near the Marshall Islands. Polyipnus indicus Schultz Figure 68 Pohjipnus indicus Schultz, 1961: 645 (holotype BMNH; off Zanzibar; not seen; paratype USNM 179897; seen); 1964: 241. Pohjipnus nuttingi: Norman, 1939: 20. Species distinction. See P. ohioJiis (p. 90); differs from P. nuttingi by its longer, ^sharper post-temporal spines, less extended abdominal keel scales, less spinose ventral border of lower jaw, photophore character- istics,, presence of spiny subcaudal keel scales, generally lower gill raker number, and shorter post-temporal base-to-dorsal spine length compared to post-temporal spine length. Description. D. 13-14; A. 15-16 (17); P. (12) 13-14; total gill rakers 20-21 (22); vertebrae 33-34. Largest specimen less than 55 mm SL; trunk tapering to long caudal peduncle; post-temporal spine long, thin, its length greater than one-half the diameter of orbit; basal post-temporal spine short; preopercle spine long, curving anteriorly; frontal ridges minutely spinose; abdominal keel scales do not extend much below ventral body margin, these scales with multiple spines; subcaudal scales spinose; supra- abdominal photophores in steplike ar- rangement with first photophore raised substantially above second; supra-anal photophores not well separated from anals; jaws large; teeth minute; underside of 92 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 i Figure 69. Polyipnus nuttingi; modified from Gilbert, 1905. lower jaw smooth to slightly spinose; gill rakers short to medium with rough spinose internal surfaces; pigment in preservative dark dorsally and dark above anal photo- phores, dark pigment bar extends to mid- line; pigment in myomerelike striations on trunk, with definite pigment spots along posterior midline. Distribution (Fig. 63). Known from three localities in the Indian Ocean along the east African coast from the equator to 30°S; reports of P. spinosus from this area may represent P. indicus. Polyipnus nuttingi Gilbert Figure 69 Pohjipmts mittingi Gilbert, 1905: 609 (holotype USNM 51599; Hawaii; seen); Fowler, 1949: 42; Haig, 1955: 321; Schultz, 1961: 640; 1964: 247. Species distinction. See F. oluolus (p. 90) und P. indicus (p. 91). Description. D. (12) 13-14; A. 15-16; P. 13-14; total gill rakers (21) 22-24; verte- brae 3.3-34. Largest specimen less than 65 mm SL; body broad, tapering to long narrow caudal peduncle; post-temporal spine stout, rel- atively short (less than one-half eye diam- eter), slightly spinose dorsally; frontal ridges more vertically oriented than P. indicus and minutely spinose; preoperck spine short, curving anteriorly; abdominali keel scales with multiple spines; these scales extend well below ventral bod>i( margin; post-temporal spine length less than one-half the distance from its base to point of dorsal blade; subcaudal scale." smooth; supra-abdominal photophores ar- ranged in a straight line, steplike arrange- ment; the three supra-anal photophoret separated slightly but definitely from ana. photophore group; jaws large; teeth mi- nute; undersurface of lower jaw markedl) spinose; gill rakers long, spinose on interna surface; dorsal spine high; pigment ir preservative similar to P. indicus, althougl dorsal pigment bar is longer and broader Distribution (Fig. 63). Known only fron the Hawaiian Islands where it appears tc be an endemic. Polyipnus laternatus Garman Figure 70 Pohjipnus laternatus Garman, 1899: 238 ( holo type MCZ 27945; off Barbados; seen); Parr 1937: 49; Scbnltz, 1961: 639; 1964: 241. Polyipnus si)inosus: Brauer, 1906: 121 (in part) Goode and Bean, 1896: 127; Rivero, 1936: 5( Species distinction. Both P. Jaternatm and P. omphus differ from P. aquavitus anc P. unispinus by their higher meristic Marine Hatchetfishes • Baird 93 CN O ^0 CO O P if o o LU o > c Q. O 0) 94 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Figure 71. Polyipnus omphus; R/V DISCOVERY; Station 5509; SL 43 mm. counts; shorter preopercle and dorsal spines; l^roader body, photophore and pig- ment characteristics. P. laternatiis differs from P. ompluts in its broader caudal peduncle; shorter, more compact subcaudal photophores, supra-anal and supra-abdom- inal photophore characteristics, slightly shorter preopercle spine, and relatively larger eye. Description. D. 13-14 (15); A. (15) 16-17; P. 1.3-14; gill rakers (18) 19-22; vertebrae 32-33 (34). Small to medium size species, rarely ex- ceeding 55 mm SL; body relatively long and narrow, tapering into broad caudal peduncle, its width greater than width of subcaudal photophores; eye large, orbital diameter usually less than six times into SL; post-temporal spine long, thin, its total length variable (usually about one-half the diameter of orbit); dorsal spine short; preopercle spine short, broad, triangulate; abdominal keel scales smooth, not ex- tended far beyond body margin; subcaudal photophores closely allied, little space bc- tvveen each photophore; supra-anal photo- phores raised well above anals, with first supra-anal slightly lower than second; first supra-abdominal photophore raised well above other two; second supra-abdominal even with or lower than third; jaws medium to small; teeth minute; vomerine and palatine teeth small but prominent; gill rakers long, spinose on inner surface; pigment in preservative dark dorsally, dark i:)igment bar usualK' does not reach mid- line; prominent, dark spots along trunk midline; myomerelike pigment striations dorsally and vertically from midline. Distribution (Fig. 63). Restricted to the western Atlantic; abundant in the Carib- bean off Venezuela and the central Amer- ican coast, in the lesser Antilles, off Puerto Rico, Cuba, and in the straits of Florida; not reported from the Culf of Mexico oi the east coast of North America. Polyipnus omphus n. sp. Figure 71 Holotype BMNH: 11° 2rS, 48° 58'E; 8/21/64 R/V DISCOVERY; Station 5509. Pohjipntis Jatcmafiis: Kobayashi, 1963: 179; Kott haus, 1967: 22 (otoliths, photo. ). Species distinciion. See P. laternatus (p 92). Roth P. laternatus and P. omphm differ from P. acjuavitus and P. unispinm by their higher meristic counts; shorter preopercle and dorsal spines; broader body photopore and pigment characteristics. P laternatus differs from P. omphus in iti broader caudal peduncle; shorter, mo« compact subcaudal photophores; supra anal and supra-abdominal photophore Marine Hatchetfishes • Baird 95 CO o < X I— < < > 0£ a D- □ .a o a. CN tN. 96 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Figure 73. Polyipnus unispinus; after Schultz, 1938. characteristics; slightly shorter preopercle spine; and relatively larger eye. The single specimen from tlie Pacific (SIO 60-236-101) appears slightly different phenotypically from the Indian Ocean forms. These two populations should be further examined when such material is available. Description. D. 14-15; A. 16; P. 13-14; gill rakers 18-21; vertebrae (33). Largest specimen less than 50 mm SL. body narrow, tapering into narrow caudal peduncle; its greatest depth less than length of subcaudal photophore group; eye relatively small, orbital diameter greater than six times into SL; post-temporal spine long, about one-half the diameter of orbit (or greater); dorsal spine short; preopercle spine short, narrowly triangulate; abdom- inal keel scales smooth, not extending far beyond body margins; subcaudal ^Dhoto- phores spaced apart (about width of a photophore between them), distance be- tween subcaudal and anal photophores about the same as length of one of the former; supra-anal raised only slightly above anals, with first supra-anal higher than second; supra-abdominal photophores in an oblique straight line; jaws medium; teeth minute; definite vomerine teeth present; gill rakers long, spinose; in pre- servative, pigment dark dorsally with broad, dark bar reaching to or near mid- line; dark pigment spots along trunk midline with pigment striations radiating from them; dark pigment above ventral photophores. Holotype: measurements (mm): SL 40.1 BD 20.0, JL 06.1, CP 03.4; mcristics: GE 19, D 14, A 16; anal photophores 11; name:] from the Marathi word "omphus," roughlv translated as "unwanted." Distribution (Fig. 63). Extremely dis- junct range; known from a few specimeni north of Madagascar in the Indian Ocear and from a single capture in the Centra Pacific north of the Marquesas Islands Additional record: 00^00', 165°42.5'W. Polyipnus aquavitus n. sp. Figure 72 Holotype ZMUC P20969; 33° 42'S, 151° 51'E 11/13/51; R/V GALATHEA: Station 551. Species distinction. See P. laternatiis (p 92). P. iniispiniis differs from P. aquavi ills by its longer dorsal and preoperck spines, spinose preopercle and ventral kee plates, shorter subcaudal to anal photo phore distance, longer postabdominal anc Marine Hatchetfishes • Baiid 97 anal pterygiophore spines, and its narrower trunk and caudal peduncle. Description. D. (11) 12-13; A. 15-16; P. (12) 1.3-14; gill rakers 12-14; vertebrae (35). Largest specimen less than 45 mm SL; body narrow, tapering into narrow caudal peduncle; its least depth less than length of subcaudal photophores; post-temporal spine long, length greater than one-half the diameter of orbit; dorsal spine short; preopercle spine short, sharp, length less than one-half the diameter of orbit; second preopercle spine reduced; lateral surface ji preopercle smooth; abdominal keel scales smooth, not extended ventrally; mpra-anal photophores only slightly raised from anals; first supra-anal photophore 'aised above second and third; anal-sub- 3audal photophore distance one-fourth or greater than length of latter; mouth small; :eeth minute; gill rakers short to medium; n preservative, body pigment is dark over abdomen and trunk; pigment often present :n band at base of caudal rays, few dark ,Digment spots along lateral midline. Holotype: measurements (mm): SL 38.5, BD 17.7, JL 07.0, CP 03.6; meristics: GR 13, D 13, A 15, anal photophores 10; name: TOm the Danish national drink, akvavit. Disfrihution (Fig. 63). Taken abun- dantly off Sidney, Australia, and known from single captures in the Banda Sea and jetween Tasmania and New Zealand. °olyipnus unispinus Schultz Figure 73 Foltjipniis unispinus Sclmltz, 1938: 137 (holotvpc USNM 103153; Philippines; seen); 1961: 643; 1964: 247. Species distinction. See P. latematus (p. )2). Differs from P. aquavitus by its onger dorsal and preopercle spines, spinose ircopercle and ventral keel scales, shorter iubcaudal to anal photophore distance, onger postabdominal and anal pterygio- :)hore spines, and its narrower trunk and audal peduncle. Description. D. 12-13 (14); A. 1.3-15; P. 12-13; gill rakers (11) 12-14; vertebrae (.3.5-,36). Small, possibly a "dwarf" species, none yet exceeding 40 mm SL; body narrow, tapering into long narrow trunk and caudal peduncle; post-temporal spine long, almost equal to orbital diameter; dorsal spine long, high; its length about equal to one-half of orbital diameter; preopercle spine long, greater than one-half of orbital diameter; a well-developed second pre- opercle spine usually present; dorsal lateral surface of preopercle spinose; abdominal keel scales spinose ventrally; postabdom- inal and anal pterygiophore spines well developed; supra-anal photophores almost continuous with anals ; first two supra-anals raised markedly above third; distance be- tween subcaudal and anal photophores less or equal to one-fourth the length of the latter; mouth small; teeth minute; vomerine teeth prominent; gill rakers short to medium, well spaced; in preservative pig- ment slightly darker dorsally; abdomen and trunk relatively dark; Distribution (Fig. 63). Taken in small numbers off the Philippines, and repre- sented by two small samples from the Banda Sea. Polyipnus triphanos Schultz Figure 74 Polyipnus iriphanos Schultz, 1938: 140 (holotype USNM 103027; Pescador Islands; seen); 1961: 640; 1964: 247. Species distinction. See P. asteroides (p. 99); differs from P. polli by its higher anal photophore number, lower gill raker and dorsal ray counts, a straight ventral anal photophore margin, and the raised dorsal pigment border above the supra- anal photophores. Description. D. 11-12; A. 17 (18, 19); P. 1.3-14; gill rakers (15) 16-18 (19); vertebrae (33). Body broad, tapering into narrow caudal peduncle; its height slightly greater than length of subcaudal photophore group; 98 BuUctin Museum of Cuuiparative Zoology, Vol. 142, No. 1 Figure 74. Polyipnus tripbanos; after Schultz, 1938. post-temporal spine short, ncedlelike; dor- sal spine short; preopercle spine short, triangulate; abdominal keel scales extend only slightly below ventral body margin; first siipra-anal photophore markedly lower than other two; first supra-abdominal raised above others, second lower than third, jaw medium to small; teeth minute; gill rakers medium, spinose; in preservative, pigment dark dorsally; dark pigment bar reaches almost to midline followed by light stripe reaching towards mid-dorsal line; ventral margin of dark dorsal pigment markedly raised on trunk above supra-anal photophores; pigment spots present on trunk midline, striations not distinct. Distribution (Fig. 63). Known only from a few captures off the Philippines. Addi- tional Record: 05^^ 01.0'S, 127° ST'E. Polyipnus polli Schultz Figure 75 Palyijmtis polli Schultz 1961: 635 (holotype MRAC 95092; south east Atlantic; not seen; para type USXM 179878; seen); 1964: 247; Blache, 1964: 71; Backus et al, 1965: 139. Pohjipmis latcniatus: Norman, 1930: 305; Fowler, 1936: 1208; Poll, 1953: 65. Pohjipniis spinofius: Smith, 1953: 102 (?). Species distinction. See P. osteroidcs (p. 99) and P. triphanos (p. 97). Description. D. 14-15 ( 16); A. ( 15) 1&- 17; P. 13-14; gill rakers (20) 21-23; verte- brae 32-33. Medium to small species, seldom exceeds 50 mm SL; body and trunk broad, narrow- ing abruptly to small, short caudal pedun- cle; post-temporal spines short, needlelike; dorsal spine short; preopercle spine short, triangulate; abdominal keel scales smooth; not extended ventrally; first supra-anal photophore lower than other two; first supra-abdominal raised well above second which is approximately even wnth third; subcaudal photophore group short, about equal to width of dorsal pigment bar at itsi center; body margin below anal photo- phores markedly curved; anal ptery- giophores extend well beyond body margin;' jaws small; teeth minute; gill rakers medium to long, and spinose; in preserva- tive, pigment dark dorsally with pigment Marine Hatchetfishes • Baird 99 Figure 75. Polyipnus polli; after Norman, 1930. bar reaching to\\ard midline; ventral border of dorsal pigment in straight line, from lateral photophore to caudal pe- duncle; dark pigment spots on midline and between midline and border of darker dorsal pigment; pigment striations present on trunk. Distribution (Fig. 63). Restricted to the southeastern Atlantic along the west African coast from the Gulf of Guinea to 10°S latitude. Polyipnus asteroides Schultz Figure 76 Polyipnus asteroides Schultz, 1938: 138 (holotype USNM; West Indies; not seen): 1961: 640: 1964: 247; Scott, 1963: 1303. Polyipnus laternatus: Jespersen, 1934: 15. Species distinction. P. asteroides, P. tri- phanos, and P. poUi differ from P. meteori, P. matsiiharai, P. kiiciensis, and P. rw^geri by their lack of teeth on the posterior vomerine shaft, and by supra-abdominal and supra-anal photophore characteristics; P. asteroides differs from P. polli by its greater number of anal photophores, less broad trunk, longer subcaudal photophore group, relatively straight anal photophore margin, and attainment of greater size; differs from P. triphanos by its less broad body, higher gill raker and dorsal ray counts, and bodv pigment characteristics. Description. D. 14-16 (17); A. (15) 16- 17 (18); P. (12) 13-14 (15); gill rakers 20-23 (24); vertebrae 32-33. Large to giant species, often exceeds 70 mm SL; body relatively broad, tapering evenly to narrow but short caudal pe- duncle, its greatest depth greater than length of subcaudal photophores; post- temporal and dorsal spines short (less than one-fourth eye diameter); preopercle spine very short, triangulate; abdominal keel scales extend slightly below ventral body margin; first supra-anal photophore mark- edly lower than second; first supra-abdom- inal photophore raised well above other two, second and third supra-abdominals usually about same height; jaws medium to large; teeth minute; gill rakers medium to long, spinose on internal surface; in preservative, pigment dark dorsally; dark pigment bar extends toward but never 100 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 in CN O U < I u > o c a o Marine Hatchetfishes • Baird 101 Figure 77. Polyipnus matsubarai; after Schultz, 1961. reaches midline; lateral border of dark- dorsal pigment straight from dorsal spine to caudal peduncle; dark pigment spots mark lateral midHne, pigment striations present on trunk. Distribution (Fig. 63). Restricted to the western North Atlantic; abundant in the Caribbean and Gulf of Mexico from the coast of Venezuela to the straits of Florida; occurs off the outer islands of the West Indies and less abundantly along the east coast of North America; a single capture has been reported as far north as the Gulf 3f Maine. Polyipnus matsubarai Schultz Figure 77 Polyipnus matsubarai Schultz, 1961: 641 (holo- type USNM 179793; Kumanonada, Japan; seen); 1964: 247. Polyipnus japonicus Schultz, 1961: 643; 1964: 247. '^olyipnus asteroidcs: iMatsubara, 1941: 2; 1950: 192. Species distinction. See P. asteroides (p. 99); differs from P. ruggcri, P. kiwiensis, and P. meteori by its higher gill raker count, long narrow caudal peduncle, and very narrow dorsal pigment bar. Description. D. 12 (13); A. 16-17; P. (12) (13) 14-16; gill rakers 22-24; verte- brae 33. Largest specimens have not exceeded 50 mm SL; body broad, tapering into long, relatively narrow caudal peduncle; its length equal to or greater than its greatest depth; post-temporal spine rather long and needlelike, its length about one-fourth the orbital diameter; dorsal spine short; pre- opercle spine short, triangulate; abdominal keel scales smooth, with no \entral ex- tension; first supra-abdominal photophore raised above second which is equal to or slightly raised above third; supra-anal photophores in steplike arrangement, the third being highest; first three anal photo- phores even and parallel to midline; jaws 102 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 "'■ -.IC'.*.'.' • .' ■ ■ N*-* ■ // //'■'/■■/ •'/'••■'■•■'.-''//■.•■■■ .■•■'.■• .-•'^'^Whv^^ Figure 78. Po/yipnus ruggeri; R/V TUl; New Zealand; SL 47 mm. medium; teeth present on long posterior shaft of vomer lying ventral to parasphe- noid, resulting in three distinet tooth hear- ing areas on the vomer; gill rakers medium; in preservative, dorsal pigment har is ex- tremely narrow and reaches to midline; dorsal pigment horder is hroken hy light stripe hehind pigment har, reaching hroadly to mid-dorsal line; dorsal pigment border raised slightly above supra-anal photophores; small, dark pigment spots mark lateral midline. Distrihution (Fig. 63). Abundant in the waters off Japan in the North Pacific; a single capture in the mid-North Pacific represents this species. 57'S, 177° 38'E; Polyipnus ruggeri n. sp. Figure 78 Holotype DMNZ 4670; 31' 7/24/62; R/V TUI. Species distinction. Differs from the P. osteroides (p. 99) group by dentition and photophore characteristics and from P. matsuharai (p. 101) by dorsal pigment and gill raker characteristics; differs from P. meteori and P. kiwiensis by its dorsal pig- ment characteristics, higher gill raker count, and photophore patterns; further differs from P. kiwiensis by its smaller, rounder eye, longer, narrower caudal pe- duncle, and lesser distance between frontal crests ( interorbital ) . Description. D. 12; A. 16-17; P. 15; gill, rakers 18; vertebrae (33). Largest specimen less than 60 mm SL;] body broad, tapering into somewhat long and narrow caudal peduncle; its length greater than depth; post-temporal spine short, rough surfaced dorsally, less than one-fourth of the diameter of orbit; dorsal spine short, low; preopercle spine triangu- late; eye large, round, its length about equal to width; greatest distance betweer frontal crests (interorbital) less than oi equal to length of subcaudal photophort group; abdominal keel scales not extendec ventrally; supra-abdominal photophores ir essentially straight line, first may be slighth raised above third; first supra-anal photo phore noticeably lo\\'er than second, whicl is lower than third; jaws medium to large' teeth present on posterior vomerine shaft gill rakers medium, slightly spinose; in pre servative, pigment dark dorsally with ; Marine Hatchetfishes • Baird 103 ■igure 79. Polyipnus kiwiensis; R/V TUI; New Zealand; SL 60 mm. ^'ery reduced pigment bar; much reduced light stripe behind bar does not reach mid- ilorsal Hne; ventral border of dorsal pig- nent raised above supra-anal photophores; >mall dark pigment spots present on lateral nidline. Holotype: measurements (mm): SL 46.8, BD 30.3, JL 09.7, CP 05.5; mcristics: GR L8, D 12, A 17; anal photophores 9; name: lamed in honor of New Zealand's national iport, rugby. Distribution (Fig. 63). Kno\vn only from I few small captures off Wellington, New Zealand, and west of the Kermadec Islands. ^olyipnus kiwiensis n. sp. Figure 79 Motype DMNZ 4802; 36° 50'S, 176° lO'E- 9/26/62; R/V TUI. Species distinction. Differs from P. isteroides (p. 99) group by photophore characteristics and teeth on posterior omcrine shaft; from P. matsuharai by gill aker number and dorsal pigment char- icteristics (F. matsuharai, p. 101); from P. ■uggeri (p. 102) by dorsal pigment char- icteristics, eye size, gill raker number, caudal peduncle, and interorbital crests; differs from P. meteori by its higher gill raker counts, larger eye and mouth, photo- phore and dorsal pigment characteristics. Description. D. (11) 12; A. 16-17; P. 15-16; gill rakers 16-17; vertebrae (32) 33 (34). Largest specimens less than 70 mm SL; body broad, tapering rather abruptly into short caudal peduncle; its depth about equal to its length; post-temporal spine short, less than one-fourth the diameter of orbit; dorsal spine short, preopercle spine triangulate; eyes extremely large, their diameter less than seven times into SL; greatest distance between frontal crests (interorbital), greater than length of sub- caudal photophore group; abdominal keel scales not extended ventrally; first and third supra-abdominal photophores about even and raised well above second; first supra-anal photophore noticeably lower than second, which is slightly lower than third; jaws large, broad; teeth well de- veloped on posterior vomerine shaft and lo\\er yAw; gill rakers medium; in pre- servati\e broad, dark, dorsal bar reaches to midline; light stripe posterior to dorsal 104 Bulletin Museum of Coniparatwe Zoology, Vol. 142, No. 1 bar not reaching to mid-dorsal line; ventral border of dark dorsal pigment only slightly raised above supra-anal photophores; small dark pigment spots on lateral midline. Holotype: measurements (mm): SL 59.5, BD 36.4, JL 14.3, CP 09.7; meristics: GR 17, D. 12, A. 17; anal photophores 10; name: from Kiwi — a New Zealand bird; in the vernacular, a Kiwi is a native of New Zealand. Distribution (Fig. 63). Taken in moder- ate numbers near Red Mercury Island off the northeastern coast of North Island, New Zealand. Polyipnus mefeori Kotthaus Polyipniis meteori Kotthaus, 1967: 27 (liolotype lOES 20; off Seychelles, Indian Ocean; not seen ) . Species distinction. See P. asteroides (p. 99), P. matsubarai (p. 101), P. ruggeri ( p. 101 ) , and P. kiiciensis ( p. 103 ) . Description. (From description of holo- type (Kotthaus, 1967) and photograph.) D. 12; A. 16; P. 15; Still raker number 13-15. Known only from holotype ( SL 37 mm ) ; body broad, tapering to relatively long caudal peduncle (appears shorter than P. matsubarai); post-temporal spine needle- like, about equal to one-fourth the eye diameter; dorsal spine short; abdominal keel scales smooth, not extended ventrally; first and third supra-abdominal photo- phores about even and raised above second; first supra-anal markedly lower than second, which is lower than third; jaws medium; in preservative broad, dark, dorsal bar reaches to midline followed by light stripe which reaches mid-dorsal line: ventral border of dorsal pigment raised considerably above supra-anal photo- phores; dark pigment spots present on midline. Distribution (Fig. 63). Known only from a specimen taken near the Seychelle Islands in the Indian Ocean; two juvenile Fohjipmis from the east coast of Africa may represent this species. Note: Key characters checked with holotype through the courtesy of Dr. Verner Larsen, ZMUC. DISCUSSION AND CONCLUSIONS Patterns of Distribution The ecological distinctness of the family and the basic stiuctural modifications in- volved in the peculiar body form were dis- cussed above. Given this basic structural) similarity, the respccti\'e genera have di- verged morphologically and ecologically. This is apparent in the distinctive distribu- tion pattern of each genus and is indicative of the types of distributions to be found in deep-sea fishes. Fohjipnus. Although Poh/ipntis has the' basic adaptive attributes of a midwater fish, the genus — with the exception of isolated expatriates — is associated with land areas. Land-oriented distributions, have been reported in midwater fishes (Ebeling, 1962; Nafpaktitis, 1968), but these have involved mc>mbers of essentially pelagic genera. Polyipnus is a moderately speciose genus which has adapted solely to land associated environments. While continental slope areas are important, this genus occurs abundantly near oceanic islands well away from continental margins. Depth data are generally sparse, but indi- cate that Polyipnus is found from 50 m to 400 m. The extent of diurnal migration is unknown, although certain species have been reported near the surface at night off Japan (Haneda, 1952). The pseudo- pelagic" environment of this species has not been extensively sampled in most areas. Species ranges are therefore incomplete, and little is known about population struc- ture and vertical distribution. New species can be expected and additional revision will be required as collecting proceeds. The peculiar distribution of this genus may be related to land-oriented food' chains. There is an extensive amount of literature on the increased productivity associated with land areas and on the Marine Hatchetfishes • Baird 105 Dccurrence of neritic forms of zooplankton. Foh/ipnus has specialized feeding habits, and two peculiar morphological features may be involved in its adaptation to a specialized niche. These features are the jaw and branchial morphology, and the enlargement of the otic region. An addi- tional indication of biological differences From the other genera is the small number of juN'eniles collected mth the adults. Much remains to be known about the biology and ecology of this genus, as well IS its "pseudopelagic" environment. Comparison of the essentially tropical and-oriented distribution of Pohjipnus with >ther tropical shore species provides some nteresting parallels. Tropical reef and ihelf fish are diverse in the Indo-west Pacific region, with the Indo-Malayan area 'he most speciose (e.g., Ekman, 1967: 17). riie number of species declines as one proceeds from this area. Wliile present in nany of the islands of outer Polynesia 'Hawaii, Marquesas, Tuamotu archipelago). Few shore species reach the western coast of the Americas. This is attributed to the wide stretch of open water in the eastern Pacific (the zoogeographic east Pacific barrier). Contributions of Indo-west Pacific elements to the tropical Atlantic are re- duced by a similar, although not as re- strictive, central Atlantic barrier, in addi- tion to the African continent ( Briggs, 1960, 1961). The tropical shore fauna is further characterized by its "modernness." It con- sists primarily of the most advanced and latest evolved fishes, with relicts and more primitive groups less well represented. Geographic endemics are common, espe- cially near the more isolated island groups. The largest number of Pohjipnus species have been collected around the Philippine Islands. Eight of 17 kmown species occur in the tropical west Pacific. Endemics occur in New Zealand, Hawaii, and the Marshall Islands at the limits of the range in the Pacific. Three other species occur in the western Indian Ocean, thus account- ing for 14 of 17 species in the Indo-west Pacific. No species are reported from the eastern Pacific. Tlie P. spinosiis species complex is not found in the Atlantic; only three species occur there. Two are re- stricted to tropical and temperate America, and one to the west African coast. There are no trans-Panamanian species. Speci- ation tends to be geographic and endemics are numerous. Extensive sympatry be- tween species complexes is rare. Life history features apparently restrict species to land-associated waters. No open-water pelagic populations are known, and bar- riers to gene flo\^' among discontinuous populations appear considerable. Here, then, is a classic tropical shore distribution in what appears to be the most primitive genus of the family (Ebeling, 1962, indicates some of the same features in MeJamphaes) . Since such a distribution is characteristic of lately evolved groups, it is interesting to speculate on the possible recent origin of Pohjipnus. While primitive maurolicid gonostomatids are identified from the early Tertiary, Pohjipnus as pres- ently defined, is not. (It is not present in Tertiary Tethys or California deposits.) ArgyropeJecus is kno\\'n from the Oligo- cene. Its distribution is worldwide (includ- ing the Tethys fauna), as are a number of gonostomatids (admittedly a different ecology and distribution pattern). Polij- ipnus, ^^'hile primitive in axial skeleton characteristics, is nevertheless highly spe- cialized in the cranial region. These char- acters may be the major adaptive features allowing Pohjipnus access to its specialized niche, resulting in a new adaptive type w^hich possibly arose relatively recently. Ars.ijropeIecus. Argijwpelecus species are characterized by broad worldwide high seas distributions. Tlie genus is found in all tropical and temperate oceans, and is absent from polar seas. The limits of dis- tribution are bounded approximately by the 5° isothenn at 200 m. Within these broad limits, however, distribution can be quite restricted with the result that world- wide species are broken up into a series of 106 Bulletin Museum of Com par a live Zoology, Vol. 142, No. 1 disjunct populations whicli appear more or less isolated from each other. In general, species occur vertically over the same depth range wherever they are found. With the exception of A. il,iil.as, Ariiijropclecus species are partial or in- complete diurnal migrators. At night many species ascend to above 300 m, often to about 200 m from their daytime depths of 400-500 m. Catches in the upper 100 m seldom involve large numbers of in- dividuals. A. aculcaiiis is most distinct in its vertical migration, while A. gigas, the deepest living species, migrates very little. Within these broad limits (150-600 m) depth variability is high, indicating considerable microcomplexity (Appendix B). From batluscaphe obser\-ations during the day, Peres (1958) reports A. hemigymnm from 250-600 m, with large concentrations from 400-500 m. Peres' and odier bathyscaphe observations (Drs. R. Rosenblatt, R. Haed- rich, and R. Richards, personal conver- sations) indicate that Argywpelccus species do not school in the classical sense, but are somewhat isolated from one another. Catch data (Table 2.3) show the wide range in size distribution with large catches, an- other indication of nonschooling behavior. Unlike many midwater fishes, the larvae and ju\'eniles of Argyropeleciis are found in the adult environment (Table 23) (Ahls- trom, 1959). Over the range of a species distribution some gravid females and young ju\eniles were usually found. Large scale expatriation does not appear to be im- portant. Wherever a species is found in an area in numbers it seems to represent a breeding population. Argiiwpchcus is represented by seven species in three species complexes. Species are moqjhologieally distinct in most cases and, as with Poh/ipnm, broad sympatry within complexes is uncommon. Sympatry is limited to zones of mixing between allo- patric species ranges w hen it occurs within species complexes. Dwarf and giant spe- cies occur. The giant species (A. gigas) is quite restricted in distribution, limited essentially to zones of water-mass bound- aries. The dwarf species (A. Iwmigymmis), while occurring in the relatively unproduc- tive central water masses, is abundant in highly productive temperate and eastern b()undar\- current waters. Sfcnwptyx. Sfemoptyx species have broad worldwide pelagic distributions similar to Argyropelectis and with approxi- mately the same geographic limits. The juveniles are found in the adult environ- ment, although larger individuals may be found slightly deeper. There is no indi- cation of expatriation. Sternopty.x is less speeiose than ArgyropeJecus and species distinctions are much less marked. Two of the three species (S. diaphana and S. ohscura) have wide allopatric ranges, with restricted areas of overlap. S. pseudohscura and S. diaphana are broadly sympatric over much of their respective ranges. Vertically, all species are deep living (500-1500 m) and show little diurnal mi- gration. Variability in catch size ranges^ indicates that Stenwptyx probably does not school. Geographic Variation Nhiyr (1963: 333) makes the following points in a discussion of geographic varia- tion: Every population of a species differs from all other populations genetically, and when sufficiently sensitive tests are em- ployed, also biometrically. The degree oB divergence between different populations' of a species ranges from near complete identity to distinctness almost of speciest level. Various characters of a species may and usually do differ independently. The characters of a given population have atl least a partial genetic basis, and in most' cases tend to remain rather constant through the years. The absence of detectable differences! between horizontally disjunct populations is not necessarily indicative of no popu-. lation differences. In the present study, methods were not particularly sensitive, nor were many characters used. However. Marine Hatchetfishes • Baud 107 w'liere differences do exist one can delimit populations which, when coupled with dis- tributional data, should add to our under- standing of the environmental and bio- logical factors which are important in restricting species distributions. Most of the patterns of geographic vari- ation outlined by Mayr are present in Ar. pseudohscura 200 400 200 400 200 400 200 400 200 400 200 400 400 600 600 SOO 600 800 1000 800 1000 15-21 (all oceans) 10-15 ( all oceans ) 12-13 ( all oceans ) 9 (Atlantic) 7 (Pacific) 7-12 (Pacific) 6-10 (Pacific) 9-14 (all oceans) 6.5-11 (all oceans) 9-18 (all oceans) 6.5-13 (all oceans) 9-14 ( all oceans ) 6.5-11 (all oceans) 7-12 (all oceans) 6-10 (all oceans) 5-7 (Pacific); 9-11 (Indian & Atlantic) 5-4.5 (Pacific); 7.5-10 (Indian & Atlantic) 7.5-10 ( Indian & Pacific) 6-8 (Indian); 4.5-5.5 (Pacific) 4.5-5 ( Indian & Pacific ) 8-5 (all oceans) 4-5 ( all oceans ) estrictcd primarily to the eastern boundary 'urrcnts and areas of upwelling which are •haracterized by cool water between 300 n and 400 m. A. olfersi is restricted to the -vanner areas of subpolar waters character- zed by 12-13° temperatures at 200 m. A. lemiii^ijmmis is excluded only from ec^ua- orial waters, although biometric data ndicates a population structure which :;orresponds to water mass boundaries. A. iciileatus is restricted to wami central kvater masses in areas bounded approxi- mately by the 15° isotherm at 200 m. A. hjcliniis occurs only in the Pacific equa- torial water mass characterized by cool temperatures between 200 m and 400 m. S. diaphami is excluded from the equatorial water masses only. S. obscura is limited to equatorial water masses and their bound- aries while S. pseudohscura is similar to S. diaphana, although more restricted to tropical and subtropical waters. The above distribution pattern is a strong argument for the importance of fABLE 23. Catch statistics for two large samples of Argyropelecus from the North Atlantic. 5IZE class figures REFER TO THE NUxMBER OF INDIVIDUALS IN THE SAMPLE WHOSE STANDARD LENGTH ?"ALLS BETWEEN THE SIZE LIMITS; I.E., THERE WERE TWO INDIVIDUALS OF A. ACULEATUS WHOSE STANDARD LENGTHS WERE FROM 21 TO 25 MM. Total Catch Size Class (mm) Species 10 15 20 25 35 45 55 65 90 90+ 4.. hemigynmus* \. aculeatus** 240 75 9 61 37 3 43 2 80 6 10 13 29 15 8 1 * Atlantis II 13, station 1040, 0940-1125, 320-375 m. ** Chain 32, station 859, 0835-1305, 380 m. 110 Bulletin Museum of Coinpaiatice Zoology, Vol. 142, No. 1 Table 24. OccimnExcE of the species of Argy ropelecis and Sternopty.x in various wateh MASSES. X = TAKEN IN NUMBERS; S = REPORTED IN SMALL NITAIBERS USUALLY NEAR AVATER MASS BOUNDARIES; O = UNRECORDED; ? = POSSIBLE RECORD. Species 00 1 on s K 1 00 09 3 1 "a u i -a g 3 1 1 o •a 3 1 Water Mass 't •< < < ■X ^ ^ W3 (yj c/i N Atlantic Central X X X o X X X X o X S Atlantic Central X X X o X X s X o X EN Pacific Central o X X o X o o X o X WN Pacific Central o o s o o s o X o 0 ES Pacific Central s o s o o X s X o X ^^'S Pacific Central o o s o X X o 0 o 0 NE Pacific Transitional o X X X X o o s X o SE Pacific Transitional X X X X X s s X X o Pacific Equatorial o s o X o o o s X 0 Indian Eciuatorial s X X o X o o o X s Indian Central X s X ? s X o X s X Subantartic X o X 6 o o X X o 0 N Pacific Subarctic o o X o X o o X o 0 temperature in defining distribntions (e.g.. MeGowan. 1960; Xafp^aktitis, 196S). ItYs apparent, however, that absolute tempera- ture values per se are not the sole limiting factor and that eaeh water mass can be defined by a host of other physical and biological factors, all of which ma\ be important in limiting distributions. A number of recent studies indicate that water masses ha\e a biological identit\- and man\" widely diverse forms are limited to them (Bieri, 1959; Aron, 1962; Brinton. 1962; McGowan, 1963; Fager and Mc- Gowan, 1963). Additional aspects to be considered are the Indrographic features such as boundar\- areas, transitional waters and upwelling areas which pro\ ide further heterogeneity of biological importance. Pelagic hatchetfish distributions are par- ticularly illustrati\e of the biological simi- larities of areas with corresponding h\dro- graphic properties. For instance, eastern boundar\- currents which are quite similar h\drographicall\ (\A'ooster and Reid. 1963) contain the same hatchetfish species wher- ever they are found. The same can be said of central g>re areas or subpolar waters. While the physical and biological proper ties of the whole water column are im portant in the ecolog\- of a gi%en wate mass, barriers to distribution in hatchet fishes appear to be primarily a function oi the en\ironmental properties over th« depth range of the species. Furthermore the barriers become less marked witl depth so tliat discontinuities at 800-100* m occur less often than those from 200- 400 m. This is reflected in the broac distributions of the deep living Sternoptyxi with three closely related species com pared to the highl\- disjunct and mon speciose Arii.iiropclecii.s w ith its more shal low distribution. This same pattern is evil dent in other deep living forms (Davi 196:3; Ebeling, 1962; Grice, 1963; Grice an Hulsemann, 1967) where life history' features of juveniles or lar\ae do not com plicate the distribution. i Zoogeographic Regions Se\"eral features make the pelagic hatche fishes particularK" well suited to zoogeo graphical studies. Their distributions ar' broad, vet limited to waters of simila J Marine Hatchetfishes • Baiid 111 Table 25. Zoogkographic Regions. Region Species Assembly 1. E Pacific Equatorial 2. N Pacific Transitional 3. Pacific Subarctic 4. EN Pacific Central 5. WN Pacific Central 6. SE Pacific Transitional 7. Pacific Subantarctic 8. S Pacific Central 9. Indian Equatorial 0. Java-Indonesian 1. Indian Central 2. Tropical Atlantic 3. SE Atlantic Transitional 4. Venezuelan-Caribbean 5. Caribbean-Gulf Central 6. Gulf Peripheral 7. NW Atlantic Pocket 8. WN Atlantic Central 9. EN Atlantic Central :0. NE Atlantic Subarctic ',1. SW Atlantic Central ;2. W Mediterranean 13. N New Zealand Pocket A. SE Atlantic Subantarctic A. lijchnit.s, S. ohscura. A. affinis, A. hemigyinnus, A. sladeni, S. ohscura, A. lychnus (S. diaphana). A. sladeni, A. hemigtjmnus (S. diaphana). A. affinis; A. sladeni, A. hemigtjmnus, S. diaphana. A. acidealus, A. hemigymnus, S. diaphana, S. pseudohscura. A. affinis, A. sladeni, A. hemigymnus, A. hjcJinus, A. gigas, S. diaphana (A. olfersi, S. ohscura). A. olfersi, A. hemigymnus, A. gigas, S. diaphana. A. aculeatus, A. hemigymnus (S. pseudohscura). A. affinis, A. sladeni, A. hemigymnus, S. ohscura. A. affinis, A. sladeni, A. hemigymnus, S. diaphana, S. ohscura. A. aculeatus, A. hemigymnus, A. gigas, S. diaphana, S. pseudohscura. A. sladeni, S. diaphana, S. pseudohscura (A. hemigymnus). A. affinis, A. sladeni, A. gigas, A. diaphana, S. pseudohscura. A. affinis, A. sladeni, A. hemigymnus, S. diaphana, S. pseudohscura, A. aculeatus. A. aculeatus, A. hemigymnus, S. diapharm, S. pseudohscura, A. sladeni. A. affinis, A. sladeni, A. hemigymnus, A. gigas, S. diaphana. A. affinis, A. sladeni, S. diapliana, S. pseudohscura. A. aculeatus, A. hemigymnus, S. diaphana (A. gigas). A. aculeatus, A. hemigymnus, S. diapliana, S. pseudohscura (A. gigas). A. olfersi, A. hemigymnus, A. gigas, S. diaphana, S. pseudohscura. A. aculeatus, A. hemigymnus, S. diaphana, S. pseudohscura (A. sladeni). A. hemigymnus. A. sladeni. A. olfersi, A. hemigymnus, S. diaphana. iydroc;iaphic properties; they are relatively luinerous and easily eaiight; expatriation is iniited; adults and ju\eniles share the same mvironment; they are only partial migrators it best and are thus less affected by seasonal luctuations; and they occur over much of he depth range of the "mesopelagic" en\'i- onment. As we have seen above, the water masses as presently defined are too broad to ex- plain species distributions as we find them. However, the concept of water masses as bodies of water with similar hydrographic and biological properties is important, and seems to be the most significant one in explaining much of the heterogeneity in the midwater environment. The pelagic hatchetfishes are used in Figure 80 as 112 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 indicator species of waters of similar properties and their associated discontinui- ties. The results may add to greater appreciation of water masses — both con- ceptuall)' and geographically. Table 25 and Figure 80 list the zoogeographic regions and are also an attempt to indi- cate similarities between regions. The characteristic species assemblages which occur in these regions are listed under the appropriate area. No boundaries were drawn because in most cases they could not be defined. Important isotherms are in- cluded and may serve as rough boundaries. Presently defined water mass boundaries (see Sverdrup et al., 1960) in many cases mark the limits of these areas. Several attempts at defining oceanic zoogeographic regions have been made (Ebeling, 1962; Clarke, 1966) and Figure 80 represents an additional one. No at- tempt has been made to categorize these regions as primary or secondary, but cer- tainly some regions involve the whole of the mesopelagic environment, while others seem important only at shallower depths. Considerable variation exists in the sharp- ness of the boundaries and, to some extent, in the degree of species overlap. As knowl- edge of the oceans and their fauna in- creases, the nature and extent of these regions and their boundaries will become more apparent. Areas which are zoogeographic regions and have boundaries which appear throughout the "mesopelagic" environment are the tropical east Pacific, the Indian equatorial region, the northeast Atlantic, and the subantarctic, especially the Pacific portion. There is a wide subtropical belt that is continuous at deeper depths, but is broken into smaller regions above ap- proximately 600 m. The tropical east Pacific has been recognized as a major zoogeographic region, and it seems to have an endemic fauna at all levels (Brin- ton, 1962; Ebeling, 1962; Johnson and Brinton, 1963). The Indian equatorial region, while not as well known, appears to be somewhat similar to the equatorial Pacific, at least in some species of hatchet- fishes and other fishes as well (Ebeling, 1962; Gibbs and Hmwitz, 1967). The northeast Atlantic is quite different from the western Atlantic in a number of groups (Haffner, 1952; Clarke, 1966; Nafpaktitis, 1968). Additional evidence from this and other studies ( Alvarino, 1965; Gibbs, 1968) indicates that the conver- gence area, especially in the South Pacific, is a major zoogeographical region which may be quite restricted in the South At- lantic and Indian Ocean. The 5° isotherm is much closer to the central water masses at 200 m (Fig. 80), and the distance be- tween the convergences is generally less broad (Sverdrup, 1960). Regions which are distinctive for the upper 500 m are the warm central water masses of the major gyre systems, and the eastern boundary currents which are cold water areas of transition and upwelling. There are other smaller areas that are im- portant zoogeographically and are faunally similar to the major regions. These include pockets of cold water around the Gulf of Mexico, off South Africa, off the southeast United States (see Haffner on Chauliodm, 1952), in the southern Caribbean and tropical Atlantic, off Java, off New Zealand, and southeast of Hawaii. The Sternoptychidae are represented by a single species in the western Mediter- ranean, an area which seems distinct from the warmer eastern end. Tliis population is distinct from the North Atlantic one, and this distinction has been documented for other midwater fishes (Marshall, 1963). Hatchetfishes have not been taken in the Red Sea proper (Marshall, 1963) or the Gulf of California (Lavenberg and Fitch. 1966). Ecological niches and diversity. Speci- ation pattern, distribution, and population structure are three indicators of diversity, niche breadth, and heterogeneity in the mesopelagic environment. While the world- wide midwater environment is heterogene- Marine Hatchetfishes • Baird 113 CD O CN E "5 Q. n (It LO -13 CN "D (U £ XI n a) 1— _c -n n o c X o (1> O) 01 listed below from numerous institutions \\'hose kind cooperation made this study possible. C. Robins, T. Devany, J. Stieger, Institute of Marine Science, University of Miami F. Berry, W. Richards, Bureau of Com- mercial Fisheries, Miami R. Tiavenberg, Los Angeles County Museum J. Savage, J. Paxton, Hancock Foundation, University of Southern California K. Suzuki, University of Mie, Japan R. Rosenblatt, D. Hoese, Scripps Institute of Oceanography E. Ahlstrom, Bureau of Commercial Fish- eries, La Jolla J. Marr, B. Rothschild, Bureau of Com- mercial Fisheries, Honolulu J. Nielsen, Zoological Museum, Copen- hagen P. Foxton, J. Badcock, National Institute of Oceanography, England P. Greenwood, G. Palmer, British Museum (NH), England J. Moreland, Dominion Museum, New Zea- land D. Cohen, N. Gamblin, L. Schultz, U. S. National Museum, Washington B. Zaranuhee, OccanograjDhic Data C(>nter, Washington For their understanding discussions and for such gracious hospitality, I am par- ticularly grateful to T. Alie of the Uni- versity of Tokyo, and to E. Bertelsen of the Danish Carlsberg Foundation. I wish to thank the staff of the Woods Hole Oceanographic Institute, especially Richard Backus and James Craddock, who offered me every encouragement and gave freely of their time and facilities. My thanks also to Jules Crane of Cer- retos College, who kindly provided fossils from his collection; to Stanley Weitzman of the U. S. National Museum for hiS' thoughts and ideas on the subject of osteol- ogy; and to W. Bossert of Harvard Uni- versity, who introduced me to the worldi of computer science. I wish to acknowledge the staff of the Museum of Comparative Zoology, par- ticularly Myvanwy Dick; the graduate stu- dents of the fish department whose assist- ance and discussion were invaluable; B. Nafpaktitis and M. Eckardt of the Uni- versity of Southern California; Sharon Horn, and Nancy Smith (illustrations); M. HowbcM-t, who pr()\ided many of the original illustrations through the courtesy of the Woods Hole Oceanographic Insti- tute; K. S. Baird, and Penelope Lasnik fori her many hours before the c^ditorial mast. Finally, I owe a special debt of gratitude and thanks to Giles W. Mead, my major Marine Hatchetfishes • Baird 115 professor, for his guidance, encouragement, criticism, and support in making my gradu- :ite years at Harvard a full and rich ex- perience. SUMMARY 1. The Sternoptychidae are primitive stomiatoid fishes closely related to the Gonostomatidae, but different from them morphologically; most of this difference is related to the peculiar deep body shape of the former. 2. The Sternoptychidae probably arose during the early Tertiary as part of an early stomiatoid radiation. Miocene fossils of Argyropelecus could not be distinguished from their modern coun- terparts, indicating little osteological evolution in this genus since then. 3. The three genera in the family are widely divergent; each has specialized in a separate direction. 4. Polyipniis occurs only in close associ- ation with land. Its pattern of distri- bution and speciation closely parallels that of many tropical shore species. 5. Ariiijropelecus is distributed widely in all tropical and temperate seas. It is a partial migrator not often entering the upper 100 m at night. Adults and juveniles are found in about the same depth range. Argijropelecus inhabits the upper "mesopelagic" zone (100- 600 m). 6. Sternopfi/x is distributed horizontally within the same limits as Argyropele- cus. It inhabits the lower "mesopelagic" zone (500-1500 m) and does not appear to migrate diurnally. 7. ArgyropeJecus and Sternoptyx species are restricted in distribution, each spe- cies seemingly restricted to waters with similar hydrographic and biological properties. 8. Argyropeleciis is more speciose and shows more morphological variation than Sternoptyx. Species ranges in Sternoptyx are much broader, indicat- ing that barriers to distribution and heterogeneity may be more pronounced in the upper "mesopelagic" than in the lower. 9. Certain species assemblages occur in waters which are hydrographically similar. These assemblages are used to zoogeographically define distinct areas of the world's oceans. 10. Ecological niches in the Sternoptychi- dae are broad over measurable niche parameters. Allopatric species ranges are the rule and, where congeneric sympatric species occur, there is usually a considerable amount of morpho- logical or vertical distinctness. Appendix A Institutions and Cruises from Which Material Was Examined or Recorded 1. Institutions and their abbreviations. Collections of T. Abe and O. Suzuki, Tokyo, Japan. Australian Museum, Sidney, Australia. Bureau of Commercial Fisheries, Honolulu, Hawaii. Bureau of Commercial Fisheries, La Jolla, California. Bureau of Commercial Fisheries, Miami, Florida. British Museum (Natural History), London, England. Dominion Museum, Wellington, New Zealand. International Indian Ocean Expedition. Biologische Anstalt Helgoland (Meteor Indian Ocean Expedition), Hamburg, Cermany. Los Angeles County Museum, Los Angeles, California. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. ABE AM BCFH BCFL BCFM BMNH DMNZ HOE lOES LACM MCZ 116 Bulletin Museum of Comparatwe Zoology, Vol. 142, No. 1 Appendix A (Continued) Musee National d'Histoire Naturelle, Paris, France. Musee Royalc d'Afriqne Central, Tenuren. National Institute of Oceanography, Sin-rey, England. Oceanographic Data Center, Washington, D.C. Scripps Institute of Oceanography, La Jolla, California. Institute of Marine Science, University of Miami, Florida. John Hancock Foiuidation, University of Southern CaHfonu'a, Los Angeles, CaHfornia. U.S. National Museum, Washington, D.C. Woods Hole Oceanographic Institute, Woods Hole, Massachusetts. Zoological Museum, Copenhagen, Denmark. MNHNP MR AC NIO ODC SIO UMML use USNM WHO I ZMUC 2. Institutions and Cruises from Wliich Material Was Examined an d Recorded. Institution Ship and Cruise Location ABE local fishing vessels Japan AM holotypes only Australia BCFH HUGH M. SMITH 30 C Pacific HUGH M. SMITH 31 C Pacific BCFL larval and juveniles only EN Pacific BCFM GERONIMO Tropical Atlantic SILVER BAY Florida, Gulf of Mexico OREGON Gulf of Mexico BMNH DISCOVERY and others Worldwide DMNZ TUI New Zealand IIOE ANTON BRUUN 3 and 6 Indian Ocean lOES METEOR (holotype description) Indian Ocean LACM VELERO NE Pacific MCZ ATLANTIS, CAPTAIN BILL III N Atlantic GOSNOLD, BRUUN 13, Chile, Tropical Atlantic CHAIN 17-49, miscellaneous collections Mediterranean MNHNP holotypes only Atlantic MRAC holotype W African Coast NIO DISCOVERY (1955-1965) NE Atlantic, Indian Ocean ODC USNS GILLISS N Atlantic, Caribbean SIO COBB 208, 303 NE Pacific BLACK DOUGLASS 203, 303 NE Pacific HORIZON N Pacific HOLIDAY, TETHYS C & S Pacific MONSOON, BAIRD C & S Pacific UMML GERDA Florida, Gulf of Mexico PILLSBURY Caribbean, Gulf of Guinea use USNS ELTANNIN Subantarctic, Pacific USNM ALBATROSS, OREGON SILVER BAY, COMBAT Atlantic, Pacific WHOI CHAIN 60, 72, ATLANTIS II 13 Caribbean, Gulf of Mexicq ATLANTIS II 31 N & SW Atlantic ZMUC GALATHEA World Cruise THOR and miscellaneous N Adantic, collections Mediterranean Marine Hatchetfishes • Baird 117 Appendix B )epth tables of the species of Argyropelecus axd Sternoptyx. Depth data for these tables lEPRESEXT MAXIMUM NET DEPTHS ONLY. "CaTCh" REFERS TO THE NUMBER OF HAULS IN WHICH THE jumber of fish taken is shown horizontally between the figure listed and the previous figure. Depth" refers to the maximum net depths and is recorded vertically between the depth listed .ND the one listed ABOVE IT. "ZeRO" CATCH REFERS TO THE NUMBER OF NEGATIVE TOWS FOR THAT SPECIES. "+" MEANS GREATER THAN OR EQUAL TO THE VALUE LISTED. Argyropelecus affinis Night Catch Catch )epth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 36 1 0 0 0 0 0 1 100 20 1 0 0 0 0 0 1 200 5 1 0 1 1 0 0 o 200 10 0 0 0 0 0 0 0 300 13 7 1 1 0 0 0 9 300 9 1 0 0 0 0 0 1 400 0 0 0 0 0 0 0 0 400 1 1 0 0 1 0 0 2 500 4 2 0 0 0 0 1 3 500 2 2 0 1 0 0 0 3 600 0 0 0 0 0 0 0 0 600 0 0 0 0 0 0 0 0 700 0 0 0 0 0 0 0 0 700 1 2 0 0 0 0 0 2 000 0 0 0 0 0 0 0 0 1000 n O 0 0 0 0 0 0 0 000 0 O 0 1 0 0 0 4 2000 0 2 0 0 0 0 0 2 001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 jalifornia, Chile { Pacific ) Gulf of Mexico, Gulf of Guinea, Caribbean Day Catch Cat( :h )epth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 1 0 0 0 0 0 0 0 100 9 0 0 0 0 0 0 0 200 0 0 0 0 0 0 0 0 200 10 0 0 0 0 0 0 0 300 5 0 0 0 0 0 0 0 300 14 3 0 0 0 0 0 3 400 19 1 1 0 0 0 0 2 400 7 0 0 0 0 1 0 1 500 7 2 1 1 0 0 0 4 500 5 0 0 1 2 0 0 3 600 1 2 0 1 0 0 0 600 4 0 0 0 0 0 0 0 700 2 1 0 0 0 0 0 1 700 0 •"1 o 0 0 0 0 0 3 000 5 1 0 0 0 0 0 1 1000 10 0 0 0 0 0 0 0 000 6 4 0 0 0 0 0 4 2000 19 3 0 0 0 0 0 3 001 + 1 0 0 0 0 0 0 0 2001 + 7 0 0 0 0 0 0 0 ^ulf of Mexico, Gulf of Guinea, Caribbean California, Chile (Pacific) Argyropelecus gigas Night Day Catch Catch )epth 0 5 10 20 50 100 100 + N Depth 0 5 10 20 50 100 100+ N 100 30 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 200 26 0 0 0 0 0 0 0 200 7 0 0 0 0 0 0 0 300 19 0 0 0 0 0 0 0 300 10 0 0 0 0 0 0 0 400 9 0 0 0 0 0 0 0 400 34 0 1 0 0 0 0 1 500 21 1 0 0 0 0 0 1 500 25 0 1 0 0 0 0 1 600 7 0 1 0 0 1 0 2 600 12 2 0 0 0 0 0 2 700 4 1 0 0 0 0 0 1 700 8 1 0 0 0 0 0 1 000 14 0 0 0 0 0 0 0 1000 20 2 0 0 0 0 0 2 :000 4 1 0 0 0 0 0 1 2000 5 0 0 0 0 0 0 0 001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 itlantic Atlantic 118 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Appendix B ( Continued ) AriiUroi)clcciis hcmigtjtnmifi (Form A) Night <:at(.li Catch Dcplli 0 5 1 0 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 20 1 0 0 0 0 1 100 23 7 1 1 0 0 0 9 200 9 1 0 0 0 0 1 200 29 0 0 2 0 0 0 2 300 11 0 0 0 0 0 0 300 20 0 0 0 0 0 0 2 400 2 0 0 0 0 0 0 400 8 1 1 0 2 0 0 4 500 4 0 0 0 1 0 1 500 23 0 1 1 0 0 0 2 600 0 0 0 0 0 0 0 600 14 0 0 0 0 0 0 0 700 3 0 0 0 0 0 0 0 700 5 3 1 0 0 0 0 4 1000 0 0 0 0 0 0 0 1000 22 1 0 0 0 0 0 1 2000 2 0 0 0 0 0 0 0 2000 5 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 {) 0 0 Carihht' an anc 1 c;uii of Mexico North Atl antic (see Day) Catch Catch n.ptii 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N J 00 5 0 0 0 0 0 0 100 35 2 0 0 0 0 0 2 200 3 0 0 0 0 0 0 200 7 0 0 0 0 0 0 0 300 4 1 0 0 0 0 1 300 7 7 2 0 1 0 10 400 16 2 1 1 1 0 5 400 0 0 0 0 0 0 0 500 3 0 0 1 0 0 1 500 7 0 0 0 0 0 0 fiOO 2 0 0 0 0 0 0 600 0 0 0 0 0 0 0 TOO 5 2 0 0 0 0 2 700 0 0 0 0 0 0 0 1000 8 2 1 0 0 0 3 1000 0 0 0 0 0 {) 0 2000 11 2 2 0 0 0 4 2000 3 1 0 0 0 0 1 2001 + 6 1 1 0 0 0 2 2001 + 0 0 0 0 0 0 0 Southern Ocean (Pacific) Pacific ( California ) ArgtjrojK'lccu.s hcmigyniuus (Form A) Dav Depth 0 5 Catch 10 20 50 100 100+ N Depth 0 5 Catc 10 ■h 20 50 100 100 t N 100 2 0 1 0 0 0 0 1 100 1 0 0 0 0 0 0 0 200 8 0 0 0 0 1 0 1 200 0 0 0 0 0 0 0 0 300 10 0 0 1 1 0 0 2 300 2 1 0 2 0 0 0 3 400 12 ■J 1 2 2 0 2 10 400 11 3 3 0 0 0 9 500 15 7 3 4 3 3 1 21 500 4 3 1 1 1 0 0 6 600 10 2 1 3 2 1 0 9 600 2 1 0 0 1 0 0 1 700 4 0 0 0 2 1 0 3 700 3 1 0 0 0 0 0 1 1000 25 5 0 1 0 0 0 3 1000 6 1 0 0 0 0 0 0 2000 6 0 1 0 0 0 0 1 2000 10 0 0 0 0 0 0 0 2001+ 0 North Athrntic 0 0 0 (20°-40°N, 5 0 0 =-70°\V) 0 0 2001 + Culf of M 1 exico 0 c:a 0 rihlx' 0 an 0 0 0 0 Dei.th 0 5 Catch 10 20 50 100 1 00 }- X De|-,th 0 5 Catt 10 h 20 50 100 100+ N 100 0 1 0 0 0 0 1 100 8 0 0 0 0 0 0 0 200 1 1 0 0 0 0 1 200 8 0 0 0 0 0 0 0 300 3 1 0 0 0 0 1 300 11 1 0 1 0 0 0 2 400 0 3 0 0 0 0 3 400 4 0 1 0 0 0 0 1 500 0 2 0 0 0 0 2 500 3 2 1 0 0 0 0 3 600 1 1 0 0 0 1 2 600 1 0 0 0 0 0 0 0 700 0 1 0 0 0 0 1 700 1 2 0 0 0 0 0 2 1000 0 4 2 1 1 0 8 1000 0 0 0 0 0 0 0 0 2000 4 6 1 0 1 0 8 2000 3 7 0 0 0 0 0 7 2001 + 3 3 0 0 0 0 0 3 2001 + 0 1 0 0 0 0 0 1 Southern Ocean ( Pacific ) Pacific ( California ) Marine Hatchetfishes • Baird 119 Appendix B ( Continued ) Argtjropclccu.s hcniii^ynimi.s (Form B) Night Depth 0 5 Catch 10 20 50 ] 00 100+ N Depth 0 5 Catch 10 20 50 100 100+ N 100 21 0 0 0 0 0 0 0 100 27 3 1 0 1 0 0 5 200 10 0 0 0 0 0 0 0 200 21 6 2 2 0 0 0 4 300 9 2 0 0 0 0 0 2 300 17 1 0 0 2 0 0 2 400 2 0 0 0 0 0 0 0 400 7 1 2 0 0 1 0 4 500 2 0 0 0 0 0 2 500 2 1 1 1 0 0 0 3 600 0 0 0 0 0 0 0 0 600 0 2 4 0 1 0 0 5 700 3 0 0 0 0 0 0 0 700 4 0 1 0 0 0 0 1 1000 3 0 0 0 0 0 0 0 1000 14 •1 0 3 1 0 0 6 2000 2 0 0 0 0 0 0 0 2000 1 0 1 0 0 0 0 1 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Gull' of M exico, Ca libbean North Atlantic (see Day) Depth 0 5 Catch 10 20 50 100 100+ N 100 5 0 0 0 0 0 0 0 200 2 1 0 0 0 0 1 300 3 2 0 0 0 0 2 400 8 8 1 2 1 1 13 500 0 1 0 1 0 2 4 600 1 0 0 1 0 0 1 700 4 1 2 0 0 0 0 3 1000 3 4 2 1 1 0 0 8 2000 5 9 1 0 0 0 0 10 2001 + 5 0 2 1 0 0 0 3 Southern Ocean (Pacific) Argtjropc'Icciis licinigytniius (Form B' Day Depth 0 5 Catch 10 20 50 00 100+ N Depth 0 5 Catch 10 20 50 100 100+ N 100 4 0 0 0 0 0 0 0 100 1 0 0 0 0 0 0 0 200 10 0 0 0 0 0 0 0 200 2 0 0 0 0 0 0 0 300 14 0 0 0 0 0 0 0 300 1 0 0 0 0 0 1 400 22 0 0 0 0 0 1 400 3 0 0 0 0 0 0 0 500 31 1 1 0 0 0 0 500 0 0 0 0 0 0 0 0 fiOO 14 3 o 0 0 0 0 6 600 2 0 0 0 0 1 0 1 700 11 0 0 1 0 0 0 700 1 0 0 0 0 0 0 0 1000 29 1 0 0 0 0 0 1 ^ 1000 9 1 0 0 0 0 0 1 2000 7 0 0 0 0 0 0 0 2000 10 2 0 0 0 0 0 2 2001 + 0 0 0 0 0 0 0 0 2001 + 6 0 0 0 0 0 0 0 N Atlan tic (20°-40°N, 5° 70° W) Southern Ocean (Pacific) Depth 0 5 Catch 10 20 50 100 100 + \ 100 1 0 0 0 0 0 0 0 200 0 0 0 0 0 0 0 0 300 5 0 0 0 0 0 0 0 400 20 0 0 0 0 0 0 0 500 10 0 0 0 0 0 0 0 600 3 1 0 0 0 0 0 1 700 4 0 0 0 0 0 0 0 1000 7 0 0 0 0 0 0 0 2000 10 0 0 0 0 0 0 0 2001 + 1 0 0 0 0 0 0 0 Gulf of Mexico, Caribbean 120 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Cat :h Depth 0 5 10 20 50 100 100+ N 100 25 4 2 1 0 0 0 7 200 18 10 2 0 1 0 0 13 300 21 0 1 0 0 0 0 1 400 8 3 1 0 0 0 0 4 500 17 7 1 0 0 0 0 8 600 8 6 0 0 0 0 0 6 700 0 7 2 0 0 0 0 9 1000 20 3 0 0 0 0 0 3 2000 5 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 North Atl antic Appendix B ( Continued ) Argyropelecus aculcatus Night Catch Depth 0 5 10 20 50 100 100+ N 100 8 6 1 0 2 0 0 9 200 0 3 3 1 1 2 0 10 300 5 3 0 2 0 0 0 5 400 0 0 0 0 0 0 0 0 500 2 1 0 0 0 0 0 1 600 0 0 0 {) 0 0 0 0 700 2 0 0 0 0 0 0 0 1000 2 1 0 0 0 0 0 1 2000 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Catch Dav Cailf of Mexico, Caribbean Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 4 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 0 200 7 2 1 0 0 0 0 3 200 0 0 0 0 0 0 0 0 300 9 5 0 0 0 0 0 5 300 0 3 1 0 0 0 0 4 400 6 8 4 0 4 1 0 17 400 4 3 2 0 7 0 0 12 500 10 17 1 4 2 0 0 24 500 3 3 0 1 2 0 0 6 600 10 8 0 0 2 0 0 10 600 1 1 0 0 0 0 0 1 700 9 4 0 0 0 0 0 4 700 1 0 0 0 0 0 0 0 1000 24 6 0 1 0 0 0 7 1000 2 1 0 0 0 0 0 1 2000 6 0 0 1 0 0 0 1 2000 4 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 North Atlantic (20° -40°N; 5 °-70°W) Gulf of M exico , Caribbean Argyropelecus olfersi Night Dav Catch Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 34 1 0 0 0 0 0 1 100 4 0 0 0 0 0 0 0 200 23 4 2 0 0 0 0 6 200 8 0 1 0 0 0 0 1 300 21 2 1 0 0 0 0 3 300 13 1 0 0 0 0 0 1 400 22 6 0 0 0 0 0 6 400 20 2 0 0 0 0 0 2 500 21 2 1 0 0 0 0 1 500 26 1 0 0 0 0 0 1 600 7 2 0 0 0 0 0 2 600 13 0 1 0 0 0 0 1 700 4 0 0 0 0 0 0 0 700 9 0 0 0 0 0 0 0 1 000 24 1 0 0 0 0 0 1 1000 26 0 0 0 0 0 3 2000 17 1 0 0 0 0 0 1 2000 13 4 0 0 0 0 0 4 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Atlantic and S( 3 II the rn Ocean Atlantic and Southern Ocean A rgijrapclecus sladcni 1 Night Catch Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 19 1 0 1 0 0 0 2 100 35 3 0 0 0 0 0 3 200 10 0 0 0 0 0 0 0 200 6 1 2 0 0 0 0 3 300 3 5 1 1 0 0 0 7 300 7 10 2 0 0 0 0 12 400 0 1 0 0 0 0 0 1 400 13 6 1 0 1 0 0 8 500 2 0 0 0 0 0 2 500 7 4 0 0 0 0 0 4 600 0 0 0 0 0 0 0 0 600 2 0 0 0 0 0 0 0 700 2 1 0 0 0 0 0 1 700 7 0 0 0 0 0 0 0 1000 3 0 0 0 0 0 0 1 1000 11 0 0 0 0 0 0 0 2000 0 2 0 0 0 0 0 2 2000 2 2 0 0 0 0 0 2 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Gulf of Mexico, Caribbean, Gulf of Guinea Pacific ( Cah'fornia , Marine Hatchetfishes • Baird 121 Appendix B (Continued) Depth Catch 10 20 50 100 100+ N Day Depth Catch 10 20 50 100 100+ N 100 1 0 0 0 0 0 0 0 100 8 0 0 0 0 0 0 0 200 0 0 0 0 0 0 0 0 200 7 1 0 0 0 0 0 1 300 4 2 0 0 0 0 0 2 300 6 8 0 0 0 0 0 8 400 8 6 3 2 1 0 0 12 400 4 0 0 1 0 0 0 1 500 6 3 1 0 0 0 0 4 500 3 3 0 1 0 0 0 4 600 7 1 1 0 0 0 0 2 600 1 0 0 0 0 0 0 0 700 1 3 0 0 0 0 0 3 700 1 2 0 0 0 0 0 2 1000 3 4 0 0 0 0 0 4 1000 0 0 0 0 0 0 0 0 2000 7 3 0 0 0 0 0 3 2000 8 2 0 0 0 0 0 2 2001 + 0 1 0 0 0 0 0 1 2001 + 1 0 0 0 0 0 0 0 Gulf of M exico Caribbean, Gulf of Guinea Pacific (California) Argyropelecus lychnus Night Day Catch Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 42 2 0 0 0 0 0 2 100 10 0 0 0 0 0 0 0 200 10 3 0 1 0 0 0 4 200 10 0 0 0 0 0 0 0 300 17 4 2 1 0 0 0 7 300 13 3 1 0 1 0 0 5 400 23 9 0 4 0 0 0 13 400 9 0 0 0 0 0 0 0 500 7 2 2 0 0 0 0 4 500 6 2 0 0 0 0 0 2 600 2 0 0 0 0 0 0 0 600 4 0 0 0 0 0 0 0 700 7 0 0 0 0 0 0 0 700 2 2 0 0 0 0 0 2 1000 11 0 0 0 0 0 0 0 1000 10 0 0 0 0 0 0 0 2000 18 1 0 0 0 0 0 1 2000 21 1 0 0 0 0 0 1 2001 + 8 0 0 0 0 0 0 0 2001 + 7 0 0 0 0 0 0 0 California , Chile, Tropical Pacific California, Chile, T ropical Pacific Sternoptyx o hscura Night Dav Catch Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 0 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 0 200 8 2 0 0 0 0 0 2 200 2 0 0 1 0 0 0 1 300 0 0 0 0 0 0 0 0 300 0 0 0 0 0 0 0 0 400 4 1 0 0 0 0 0 I 400 1 3 0 0 0 0 0 3 500 2 1 0 0 0 0 0 1 500 0 0 0 0 0 0 0 0 600 4 2 0 0 0 0 0 2 600 0 0 0 0 0 0 0 0 700 4 1 1 0 0 0 0 2 700 0 0 0 0 0 0 0 0 1000 5 5 0 1 0 0 0 6 1000 4 1 0 0 0 0 0 1 2000 4 1 0 0 0 0 0 1 2000 7 2 1 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 ^ 2001 + 4 1 0 0 0 0 0 1 Indian 0( ;ean Indian Ocean Stciuoptyx pseudobscura Night Catch Catch Depth 0 5 10 20 50 100 100+ N Depth 0 5 10 20 50 100 100+ N 100 0 0 0 0 0 0 0 0 100 50 1 0 0 0 0 0 1 200 10 0 0 0 0 0 0 0 20O 36 0 0 0 0 0 0 0 300 16 1 0 0 0 0 0 1 300 30 0 0 0 0 0 0 0 400 5 0 0 0 0 0 0 0 400 9 0 0 0 0 0 0 0 500 3 0 0 0 0 0 0 0 500 24 1 0 0 0 0 0 1 600 6 0 0 0 0 0 0 0 600 9 0 0 0 0 0 0 0 700 6 0 0 0 0 0 0 0 700 7 0 0 0 0 0 0 0 1000 9 2 0 0 0 0 0 2 1000 12 4 1 0 0 0 0 5 2000 5 0 0 0 0 0 0 0 2000 2 1 1 0 1 0 0 3 2001 + 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Indian Ocean North Atlantic and Gulf of C .uinea 122 Bulletin Museum of Cumparative Zoology, Vol. 142, No. 1 Appendix B (Continued) Dav Depth 0 5 Catt 10 h 20 50 100 100+ N Depth 0 5 Gate 10 1 20 50 100 100 + N 100 0 0 0 0 0 0 0 0 100 4 0 0 0 0 0 0 200 ■1 o 0 0 0 0 0 0 0 200 7 0 0 0 0 0 0 300 10 2 0 0 0 0 0 2 300 15 0 0 0 0 0 0 400 4 0 0 0 0 0 0 0 400 39 0 0 0 0 0 0 500 0 0 0 0 0 0 0 0 500 35 0 0 0 0 0 0 600 0 0 0 0 0 0 0 0 600 14 1 0 0 0 0 1 700 0 0 0 0 0 0 0 0 700 12 0 0 0 0 0 0 1000 5 0 0 0 0 0 0 0 1000 16 7 2 1 0 0 10 2000 8 2 0 0 0 0 0 2 2000 12 0 0 0 0 3 2001 + 4 1 0 0 0 0 0 1 2001 + 0 1 0 0 0 0 1 Indian Ocean North Atlantie and Gulf of G ninea Slcrnuplijx (lidpliaiia Dcptli 0 5 Catt 10 h 20 50 100 100+ Day N Depth 0 5 Gate 10 h 20 50 1(10 100 t N 100 0 1 0 0 0 0 0 1 100 0 0 0 0 0 0 0 200 0 0 0 0 0 0 0 0 200 0 0 0 0 0 0 0 300 1 0 0 0 0 0 0 0 300 4 0 0 0 0 0 400 4 0 0 0 0 0 0 0 400 14 2 0 0 0 2 500 1 0 0 0 0 0 0 0 500 5 4 0 0 0 4 600 0 1 0 0 0 1 0 2 600 1 0 0 1 0 1 700 1 J 0 0 1 0 0 2 700 0 1 0 0 0 1 1000 0 0 0 0 1 0 4 1000 2 0 0 1 0 1 2000 2 1 1 1 1 0 0 4 2000 1 3 0 0 0 3 2001 + 0 0 0 0 1 0 0 1 2001 + 0 0 0 0 0 0 0 Ciilf of G lint'a GuLf of Mexico and Cari )l)eai 1 Depth 0 5 Gate 10 h 20 50 100 100+ N Depth 0 5 Gate 10 h 20 50 100 100 \- N 100 0 0 0 0 0 0 0 100 0 0 0 0 0 0 0 200 7 0 0 0 0 0 0 0 200 3 0 0 0 0 0 0 300 9 1 0 0 0 0 0 1 300 0 0 0 0 0 0 0 400 19 0 0 0 0 0 0 0 400 3 1 0 0 0 0 1 500 20 4 1 0 0 0 0 5 500 0 0 0 0 0 0 0 600 7 0 1 2 1 0 0 4 600 0 0 u 0 0 0 0 700 4 3 0 0 1 0 0 4 700 0 0 0 0 0 0 0 1000 8 4 2 2 3 0 0 11 1000 5 0 0 0 0 0 0 2000 3 1 1 0 0 0 0 2 2000 8 2 0 0 0 0 2 2001 + 0 0 0 0 0 0 0 0 2001 + 5 0 0 0 0 0 0 North Atlantic Indian Ocean Sternoptyx diaphana Depth 0 5 Catt 10 h 20 50 100 100+ Night N Depth 0 5 Gate 10 1 20 50 100 100 i \ 100 2 0 0 0 0 0 0 0 100 3 2 0 0 0 0 ■•> 200 5 0 0 0 0 0 0 0 200 3 0 0 0 0 0 0 300 3 0 0 0 0 0 0 0 300 4 0 0 0 0 0 0 400 4 1 0 0 0 0 0 1 400 12 4 0 0 0 0 4 500 4 1 0 0 0 0 0 1 500 2 1 0 0 0 0 1 600 3 1 0 0 1 0 0 2 600 1 0 1 0 0 0 1 700 1 3 0 0 1 0 0 4 700 2 2 1 0 0 0 3 1000 0 2 1 3 3 0 0 9 1000 3 1 0 1 1 0 3 2000 1 0 0 1 0 0 0 1 2000 2 4 0 0 1 0 5 2001 + 0 0 0 0 0 0 0 0 2001 + 0 4 0 0 0 0 4 NE Atlantic (20°-40°N, 0°-30°W) Southei^n Ocean (Pacific) Marine Hatchetfishes • Baird 123 Catch Depth 0 5 10 20 50 100 100 + N 100 2 1 1 0 0 0 0 2 200 0 0 0 0 0 0 0 0 300 0 1 0 0 0 0 0 1 400 0 2 0 0 0 0 0 2 500 0 2 0 0 0 0 0 2 600 0 0 0 0 0 0 0 0 700 0 0 0 0 0 1 0 1 1000 0 0 0 0 0 0 0 0 2000 0 0 0 1 1 0 0 2 2001 + 0 0 0 0 0 0 0 0 Culf ot( Guinea Catch s ter Depth 0 5 10 20 50 100 100+ N 100 0 0 0 0 0 0 0 0 200 8 2 0 0 0 0 0 2 300 0 0 0 0 0 0 0 0 400 4 1 0 0 0 0 0 1 500 3 0 0 0 0 0 0 0 600 6 0 0 0 0 0 0 0 700 5 1 0 0 0 0 0 1 1000 8 2 1 0 0 0 0 3 2000 2 0 0 0 0 0 2 2001 + 0 0 0 0 0 0 0 0 Western Indian Oce Catch Depth 0 5 10 20 50 100 100+ N 100 1 0 0 0 0 0 0 0 200 3 0 0 0 0 0 0 0 300 3 1 0 0 0 0 0 1 400 4 0 0 0 0 0 0 0 500 7 2 0 0 0 0 0 2 600 3 2 2 2 0 0 0 6 700 2 1 0 1 1 0 0 3 1000 2 3 2 4 1 0 0 10 2000 0 0 1 0 1 0 0 2 2001 + 0 0 0 0 0 0 0 0 NE Atlantic Appendix B (Continued) Depth Catch 10 20 50 100 100+ N 100 15 1 0 0 0 1 0 2 200 10 0 0 0 0 0 0 0 300 4 3 3 0 0 0 0 6 400 0 0 0 0 0 0 0 0 500 2 1 0 0 0 0 0 1 600 0 0 0 0 0 0 0 0 700 1 0 0 0 1 0 0 1 1000 0 2 0 1 0 0 0 3 2000 0 0 0 0 0 0 0 0 2001 + 0 0 0 0 0 0 0 0 Gulf of Mexico and Caribbean SternopUjx diaphana Night Day Depth Catch 10 -2 50 100 100+ N 100 30 0 0 0 0 0 0 0 200 26 0 0 0 0 0 0 0 300 19 0 0 0 0 0 0 0 400 6 1 0 0 0 0 0 1 500 18 2 0 0 0 0 0 2 600 5 ■J 0 0 1 0 0 4 700 0 2 0 1 1 0 0 4 1000 7 1 1 2 o o 0 0 7 2000 2 0 0 1 0 0 0 1 2001 + 0 0 0 0 0 0 0 0 North Atl antic (30° -45' N, 20°-70°W) Catch ■ Depth 0 5 10 20 50 100 100+ N 100 1 0 0 0 0 0 0 0 200 2 0 0 0 0 0 0 0 300 3 0 0 0 0 0 0 0 400 1 0 0 0 0 0 0 0 500 2 0 0 0 0 0 0 0 600 2 0 0 0 0 0 0 0 700 0 0 0 0 0 0 0 0 1000 3 1 0 1 1 0 0 3 2000 3 3 0 1 3 0 0 7 2001 + 0 1 0 0 0 0 0 1 Southern Ocean (Pacific' LITERATURE CITED Ahlstrom, E. H., and R. C. Counts. 1958. De- velopment and distriliution of Vinci^iicnia lucetia and related species in tlie eastern Pacific. U. S. Fish Wildlife Serv. Fish. Bull., 58: 363-416. . 1959. Vertical distril)iition of pelaj^ic fish eggs off California and Baja California. U. S. Fish Wildlife Serv. Fish. Bull., 60: 107-146. Alcock, a. 1896. Supplementary list of the marine fishes of India. J. Asiatic Soc. Bengal, 65(2): 331. . 1899. A Descriptive Catalogue of the Indian Deep-sea Fishes in the Indian Museum . . . Collected by the Royal Indian Marine Survey Ship INVESTIGATOR. Calcutta, Baptist Mission Press, p. 211. Alvarino, a. 1965. Chaetognaths. Oceanogr. Mar. Biol. Ann. Rev., 3: 115-194. Arambourg, C. 1929. Argyiopclecus logeaiti, un nouveau poisson bathypelagique du Sahelien. Bull. Soc. Geo!. France, 29: 11-15. Aron, W. 1962. The distribution of animals in the eastern North Pacific and its relationship to physical and chemical conditions. J. Fish. Res. Bd. Canada, 19: 271-314. 124 Bulletin Museum of Comparative Zoology, Vol. 142, No. 1 Backus, R. H., G. W. Mead, R. L. IIaediuch, AND A. W. Ebeling. 1965. The mesopelagic fishes collected during cruise 17 of the R 'V CHAIN, with a method for analyzing faunal transects. Bull. Mus. Conip. Zool. Harvard, 134(5): 139-157. Badcock, J. 1969. Colour variation in two meso- pelagic fishes and its correlation with ambient light conditions. Natine (London), 221: 383-385. Bahamoxde, N. 1963. Argtjropdecus en Chile. Rev. Univ. Catol. Santiago, 48: 83-86. Barnard, K. H. 1925. A monograph of tlie marine fishes of South Africa. Ft. 1. Ann. South African Mus., 21: 1-153. Bassot, J. M. 1966. On the comparative morphol- ogy of some luminous organs. In Johnson, L., and Y. Haneda (eds.), Bioluminescence in Progress. Princeton, N. J., Princeton Univ. Press. 650 pp. Beebe, W. 1927. Preliminary list of Bermuda deep-sea fish. Zoologica (New York), 22: 197-208. Berg, L. S. 1940. Classification of fishes both recent and fossil. Trav. Inst. Zool. Acad. Sci. URSS, 5: 34(>-517. Berry, F. H., and H. C. Perkins. 1965. Suney of pelagic fishes of the California current area. U. S. Fish Wildlife Serv. Fish. Bull., 65(3): 625-682. Bertelsen, E., and J. Grontved. 1949. The light organs of a bathypelagic fish A. olfersi (Cuvier) photographed by its own light. Vid. Medd. Dansk Naturh. Foren. Kjoben- havn. 111: 163-167. Bertin, L. 1940. Catalogue des types de poissons du Museum d'Histoire Naturelle. Bull. Mus. Hist. Nat. Paris, Ser. 2, 12: 244-322. BiERi, R. 1959. The distribution of the planktonic Chaetognatha in the Pacific and their re- lationship to the water masses. Limnol. Oceanogr., 4: 1-28. BiGELOw, H. B., and W. C. Schroeder. 1953. Fishes of tlie Gulf of Maine. U. S. Fish Wildlife Serv. Fish. Bull., 74: 1-577. Blache, J. 1964. Poissons bathypelagiques de la famille de Sternoptychidae (Teleostei, Clupei- formi, Stomatiodei). O.R.S.T.O.M. Ocean- ographie, 5: 71-87. Borodin, N. A. 1931. Atlantic deep-sea fishes. Bull. Mus. Comp. Zool. Harvard, 72: 55-89. Brauer, a. 1901. iJber einige von die Valdivia- Expedition gesammelte Tiefsee-Fische und ihre Augen. Sitz. Gesellsch. Befiird. Gesam. Naturw. Marburg, 1901(7): 120. . 1906. Die Tiefsee-Fische. I. Systemati- scher Tiel. Wiss. Ergeb. Deut. Tiefsee-Exped. VALDIVIA, 1898-99, 15: 69-122. . 1908. Die Tiefsee-Fische. II. Anato- mischer Teil. Wiss. Ergeb. Deut. Tiefsee- Exped. VALDIVIA, 1898-99, 15: 1-175. Briggs, J. C. 1960. Fishes of worldwide (cir- cumtropical) distribution. Copeia, (3): 171- 180. . 1961. The East Pacific Barrier and the distribution of marine shore fishes. Evolution, 15: 545-554. Bright, T. J., and W. E. Paquegnat. 1969. Deep sea hatchetfishes of the Gulf of Mexico. Quart. J. Florida Acad. Sci., 32: 26-37. Brinton, E. 1962. The distribution of Pacific euphausiids. Bull. Scripps Inst. Oceanogr., Univ. California, 8(2): 51-270. BuEN, R. DE. 1935. Fauna Ictiologica. Pt. 1. Notas y Resumenes. Inst. Espaiiol Oceanogr., 88: 1-182. Bussing, W. A. 1965. The midwater fishes of the Peru-Chile trench. Antarctic Res. Ser., 5: 185-227. Cervigon, F. 1964. Nuevas citas de Peces para Venezuela y datos sombre algunas especies poco conocidas. Novedades Cient. Contrib. Ocas. Mus. Hist. Nat. La Salle, Caracas., Ser. Zool., 31: 1-18. Clarke, F. E. 1878. On two new fishes. Trans. Proc. New Zealand Inst., 7: 91-300. Clarke, M. R. 1966. Review of the systematics and ecology of oceanic squids. Advan. Mar. Biol., 7: 91-300. Clemens, W. A., and G. V. Wilhy. 1949. Fishes of the Pacific coast of Canada. Fish. Res. Bd. Canada, Bull., 67: 1-368. Cocco, A. 1829. Su di alcuni nuovi pesci del mare di Messina. Giorn. Sci. Lett. Arti Sicilia, 26(77): 46-50. CoLLETT, R. 1903. Meddelelser om Norges Fiske i Aarene 1884-1901. Vid. Selsk. For. Chris- tiania, 9: 108. Crane, J. M., Jr. 1966. Late tertiary radiation of viperfishes (Chauliodontidae) based on a comparison of Recent and Miocene species. Los Angeles County Mus. Contri. Sci., No. 115: 29. CuviER, G. 1829. Poissons. Le Regne Animal. 2d ed., vol. 2. Paris, p. 316. , AND A. Valenciennes. 1849. Histoire Naturelle des Poissons. Paris. Vol. 22: 398- 415. Daniltshenko, p. G. 1960. Bony fishes of the Maikop beds of the Caucasus. Trudy Paleo. Inst. Moscow, 78: 1-208 (In Russian). . 1962. Fishes from the Dabakhan seriest of Georgia. Paleon. Zhurnal, 1: 111-126 (Im Russian ) . David, L. R. 1943. Miocene fishes of Southerni California. Geol. Soc. America Spec. Pap., 43: 1-733. Marine Hatchetfishes • Baird 125 David, P. M. 1963. Speciation in Chaetognatlia. Syst. Assoc. Publ., 5: 129-143. Denton, E. J., and J. A. C. Nicol. 1965. Studies on reflection of light from silvery surfaces of fishes, with special reference to the bleak Alhiimus alhiirnus. J. Mar. Biol. Assoc. U. K., 45: 711-738. DoLLFus, R. p. 1955. Premiere contribution a I'etablissement d'un fichier ichtliyologique dn Maroc Atlantique de Tanger a I'embouchure de I'oued dra. Trav. Inst. Sci. Cherifien (ser. Zool.) Rabat, 6: 1-226. i]BELiNG, A. W. 1962. Melamphaidae I. Systc- matics and zoogeography of the species in the bathypelagic fish genus Melamphaes Gunther. DANA Rept., 58: 1-164. , AND W. H. Weed, III. 1963. Melamphai- dae III. Systematics and distribution of the species in the bathypelagic fish genus, Scopelogadits Vaillant. DANA Rept., 60: 1-58. i]HRENBAUM, E. 1909. Eicr mid Larven von Fischen, 1. Nordisches Plankton, Lief. 10: 327. Hkman, S. 1967. Zoogeography of the Sea. Lon- don, Sidgwick and Jackson, Ltd. 417 pp. ''ager, W. E., and J. A. McGowAN. 1963. Zoo- plankton species groups in the North Pacific. Science, 140: 453-460. ^OLLETT, H. 1952. Annotated list, central Cali- fornia fish. Proe. California Acad. Sci., 27: 409. ''owLER, H. W. 1933. Descriptions of new fishes obtained 1907-1910, chiefly in the Philip- pine Islands and adjacent seas. Proc. Acad. Nat. Sci. Philadelphia, 85: 233-367. . 1936. The marine fishes of West Africa. Bull. American Mus. Nat. Hist., 70: 1-275. . 1949. Fishes of Oceania. Mem. Bernice P. Bishop Mus., 12: 1-540. . 1956. Fishes of the Red Sea and South- ern Arabia. Vol. 1. Jerusalem, Weizmann Press. 240 pp. ^UGLiSTER, F. C. 1960. Atlantic Ocean atlas of temperature and salinity profiles and data from the International Geophysical Year 1957-1958. Woods Hole Oceanographic In- stitution Atlas Series, 1 : 1-209. jArman, S. 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands. Mem. Mus. Comp. Zool. Harvard, 24: 1-431. jIrbs, R. H., Jr. 1968. Photoncctcs miinificus, a new species of melanostomiatid fish from the South Pacific subtropical convergence, with remarks on the convergence fauna. Los Angeles County Mus. Contri. Sci., 149: 1-6. , and B. a. Hurwitz. 1967. Systematics and zoogeography of the stomiatoid fishes Chauliochis pammelas and C. sloani, of the Indian Ocean. Copeia, (4): 798-805. Gilbert, G. H. 1905. Fishes of tlie Hawaiian Islands. Bull. U. S. Fish Gomm., 23: 575- 713. Gill, T. 1884. Note on the Stemoptychidae. Proc. U. S. Nat. Mus., 7: 349-351. GooDE, G. B., and T. H. Bean. 1896. Oceanic Ichthyology. Mem. Mus. Comp. Zool. Har- vard, 22: 1-554. Grandperrin, R., and J. RivATON. 1966. "Cori- olis": Crosiere "Alize." Individualization de plusieurs ichthyofaunes le long de I'equateur. Cab. O.R.S.T.O.M. (ser. Oceanogr.), 4(4): 36-49. Greenwood, P. L., D. E. Rosen, S. H. Weitz- MAN, AND G. S Myers. 1966. Phyletic studies of teleostean fishes with a provisional classi- fication of living forms. Bull. American Mus. Nat. Hist., 113(4): 341-455. Gregory, W. K., and G. M. Conrad. 1936. Pic- torial phylogenies of deep sea Isospondyli and Iniomi. Copeia, ( 1 ) : 21-36. Grey, M. 1959. Deep sea fishes from the Gulf of Mexico with tlie description of a new species. Fieldiana, Zool., 39: 326-339. . 1964. Family Gonostomatidae. In Fishes of tlie Western North Atlantic. Sears Found. Mar. Res., 4: 78-238. Grice, G. D. 1963. Remarks on the diversity and distribution of deep living Copepoda. Proc. Int. Congr. Zool. Washington, 16: 312-327. -, and K. Hulsemann. 1967. Bathypelagic copepods of the western Indian Ocean. Proc. U. S. Nat. Mus., 122: 1-22. GiJXTHER, A. 1864. Catalogue of the Fishes in the British Museum. London, Taylor and Francis. Vol. 5: 384-392. . 1887. Report on the scientific results of H.M.S. CHALLENGER during the years 1873-76. Rep. Sci. Res. H.M.S. CHALLEN- GER, 22: 1-335. Haedrich, R. L., and J. G. Nielsen. 1966. Fishes eaten by Alepisaiiris (Pisces, Iniomi) in the southeastern Pacific Ocean. Deep-Sea Res., 13: 909-921. Haffner, R. E. 1952. Zoogeography of the bathypelagic fish, Chatiliodus. Syst. Zool., 1: 112-133. ' Haig, J. 1955. Fishes killed by the 1950 eruption of Maima Loa. III. Stemoptychidae. Pacific Sci., 9: 321-329. Handrick, K. 1901. Zur Kenntnis des Nerven- system und der Leuchtorganc des Argyropele- cus hemigymnus. Zoologica (Stuttgart), 13 (32): 1-68. Haneda, Y. 1952. Some luminous fishes of the genera Yarella and Pohjipnus. Pacific Sci., 6: 13-21. 126 BiiUcfin Museum of Comparative Zoology, Vol. 142, No. 1 Harrissox, C. M. H. 1967. On nietliods for sampling niesopelagic fishes. Symp. Zoo!. Soc. London, 19: 71-126. Hermann-, D. J. 1781. Uber cin neues anierikan- isc'hes Fischgcschlecht, Sternoptyx diapliaua, der durchsichtige Brust-Falten-Fisch. Dcr Natuiforscher, 16: 33-34. HoLGERSEN, H. 1958. Sjeldne fisker ved soinestky- sten. Arbok Stavanger Museum, 59: 120- 124. Holt, E. W. L., and L. W. Byrne. 1913. Sixth report on tlie fishes of the Irish Atlantic slope. Fisheries, Ireland, Sci. Invest., 1912: 1-28. HuBBS, C. L. 1953. Synonymy of the liatln- pelagic fish genus Rhynchohyalus, referred to the expanded family Argentinidae. Copeia, (2): 96-97. , AND K. F. Lagler. 1947. Fishes of the Great Lakes Region. Bull. Cranbrook Inst. Sci., 26: 1-186. Jerzmanska, a. 1960. Ichthyofauna from the Jaslo shales at Solmiow (Poland). Acta "Paleont. Folonica, 5: 367-420 (English Sum- mary ) . , and S. Jucha. 1963. L'affeurment de la faune de poissons dans les schistes de Jaslo a Tubno pres de Dynow ( Karpates Polonaises ) . Ann. Soc. Geol. Pologne, 33: 159-180. Jespersen, p. 1915. Sternoptychidae. Kept. Dan- ish Oceanogr. Exped. 1908-10. Vol. II: Biology. A, 2. p. 41. . 1934. Sternoptychidae. In Joubin (ed.). Faune Ichthvologique de I'Atlantique Nord. No. 15. , AND A. V. Taning. 1919. Some Mediter- ranean and Atlantic Sternoptychidae. Vid. Medd. Naturhist. For. Kjobenhavn, 70: 220- 228. 1926. Mediterranean Sternoptychidae. Kept. Danish Oceanogr. Exped. 1908-10. A., 12: 1-59. Johnson, M. W., and E. Brinton. 1963. Bio- logical species, water masses, and currents. In Hill, M. N. (ed.). The Sea. Interscience Publ., 2: 381-414. Jordan, D. S., and B. W. Evermann. 1896. The fishes of North and Middle America. U. S. Nat. Mus. Bull., 47 (Pt. 1): 1-604. Jordan, D. S., and E. C. Starks. 1904. Japanese fishes. Bull. U. S. Fish Comm., 22: 581. Kamohara, T. 1952. Revised descriptions of the off shore bottom fishes of Prov. Tosa, Shikoku, Japan. Repts. Kochi Univ. Nat. Sci., 3: 16-24. , and T. Yamaicwva. 1965. Fishes from Amami-Ishimi and adjacent regions. Rept. Usa Mar. Biol. Stn., 12: 1-27. Kobayashi, D. 1963. First record of PoIyii>uus lalcmattts from the Pacific Ocean. Copeia, (2): 170-180. Koefoed, E. 1961. Isospondyli 2. Heterophoto- dermi, Sternoptychidae. Rept. Sci. Res. Michael Sars. N. Atlantic, 4: Pt. 2(10): 1-12. Koepcke, H. W. 1962. Lista de los peces marinos conocidos del Peru con datos de sa distri- bucion geografica. Biota, 4: 145-154. KoTTHAUS, A. 1957. Fische des Indischen Ozeans. Sonderheft aus METEOR Forschungsergeb- nisse. Reihe D, 1: 1-22. , AND G. Krefft. 1957. Fischefaunenliste der Fahrten mit F.F.S. "ANTON DOHRN" nach Island — Gronland. Ber. Deut. Wiss. Komm. Meeresforsch., N.F., 14: 169-191. KouMANS, F. D. 1953. Biological results of the Snellius Expedition. XVI. The pisces and leptocardii. Temminckia, 9: 177-210. Lavenberg, R. J., AND J. E. Fitch. 1966. An- notated list of fishes collected by midwater trawl in the Gulf of California, March-April 1964. California Fish and Game, 52: 92-110. , AND A. W. EiiELiN'G. 1967. Distribution of midwater fishes among deep water Ixisins of the Southern California shelf. Proc. Symp. Biol. California Islands, Santa Barbara, 1 : 185-201. Ledenfeld, R. V. 1905. The radiating organs of the deep sea fishes. Mem. Mus. Comp. Zool,) Harvard, 30: 170. Lowe, R. T. 1843-1860. History of the Fishes of Madeira. London, Bernard Quaritch. p. 64. Marr, J. C. 1948. Two additions to the known fauna of California. Copeia (2): 140—141. Marshall, N. B. 1960. Swimbladder structure of deep-sea fishes in relation to their system- atics and biology. DISCOVERY Rept.s., 31: 1-122. . 1963. Diversity, distribution, and speci- ation of deep-sea fishes. Svst. Assoc. Publ.. 5: 181-195. Matsubara, K. 1941. Studies on the deep sea fishes of Japan. Suisan Kenkiu-shi, Japan, 36: 1-8. . 1950. Studies on the fishes of the genus Pohjipntis found in Japan. Japan J. Ichthy., 1: 192-197. Maul, G. E. 1949a. Lista Systematica dos Peixe.'-" Assinalados nos Mares de Madeira e Indicf Alphabetico. In de Noronha, A. C, and A. A. Sarmento (eds.), Vertebrados de Madrini 2. Peixes. Funchal, Tipografia Experanca: 1-53. . 1949b. Monografia dos peLxes de Museii Municipal de Funchal. Bol. Mus. Mun Funchal, 4: 9-17. Mavr, E. 1963. Animal Species and Evolution Cambridge, Harvard Uni\ersity Press. 797 pp I Marine Hatchetfishes • Baird 127 , E. G. LiNSLEY, AND R. L. UsiNGER. 1953. Methods and Principles of Systematic Zool- ogy. New York, McGraw-Hill. 538 pp. VIcCuLLOCH, A. R. 1914. Report on some Fishes obtained by the F.I.S. ENDEAVOUR on the Coasts of Queensland, New South Wales, Victoria, Tasmania, and South and South- western Australia. Ft. 2. Zoological Results F.I.S. ENDEAVOUR 1909-10, 2: 78-87. . 1923. Fishes from Australia and Lord Howe Island. Rec. Australian Mus., 14: 118. VlcGowAX, J. A. 1960. The relationship of the distril)ution of the planktonic worm Pocohius meseres Heath to water masses of the North Pacific. Deep Sea Res., 6: 125-139. . 1963. Geographical variation in Linia- cina helicina in the North Pacific. Syst. Assoc. Publ., 5: 109-128. \Ieao, G. W., and F. H. C. Taylor. 19.53. A collection of oceanic fishes off Northeastern Japan. J. Fish. Res. Bd. Canada, 10: 570. \lisRA, K. 1952. An aid to the identification of the fishes of India, Burma, and Ceylon. II. Clupeiformes, Bathyclupeiformes, Galaxi- formes, Scopeliformes, and Ateleopiformes. Rec. Indian Mus. Calcutta, 50: .367-422. VIoRROw, J. E. 1957. Mid-depth fishes of the Yale South American Expedition. Bull. Bing- ham Oceanogr. Coll., 16: 56-71. . 1964. Suborder Stomiatoidea; families Gonostomatidae and Sternoptychidae. In Fishes of the Western North Atlantic. Sears Found. Mar. Res. 4(1): 71. VluROMTSEV, A. M. 1963. The principal hydro- logical features of the Pacific Ocean. Jerusalem, Israel Program for Scientific Trans- lation. 790 pp. Mafpaktitis, B. G. 1968. Taxonomy and dis- tribution of the lanternfishes, genera Lohi- anchia and Diaphus, in the North Atlantic. DANA Rept., 73: 1-131. VicoL, J. A. C. 1967. The luminescence of fishes. Symp. Zool. Soc. London, 19: 27-55. VoRDEN, C. R. 1961. Comparative osteology of representative salmonid fishes with particular reference to the grayling (Thyiuallus arcti- cus) and its phylogeny. J. Fish. Res. Bd. Canada, 18: 679-789. VoRMAX, J. R. 1930. Oceanic fishes and flat- fishes collected in 192.5-27. DISCO\'ERY Repts., 2: 261-370. . 19.37. Fishes. British, Australian, and New Zealand Antarctic Res. Exped. 1929-31, 1(2): 51-88. . 1939. Fishes. Sci. Repts. lohn Nburav Exped. 1933-34, 7(1): 1-116. . 1944. A draft synopsis of the orders, families, and genera of recent fishes and fish- like vertebrates. London, British Museum ( Nat. Hist. ) 649 pp. NusBAUM-HiLARowicz, J. 1923. fitudes d'anato- mie Comparee sur les Poissons . . . Pt. 2. Res. Camp. Sci. Alliert ler Monaco, 65: 1-100. Nybelin, O. 1948. Fishes collected by the Skagerak Expedition, 1946. Goteborgs K. Vet. Vitt. Samh. Handl, ser. B, 5(16): 1-95. Ogilby, J. D. 1888. Description of a new genus and species of deep sea fish from Lord Howe Island. Proc. Linn. Soc. New South Wales, 3(2): 1313. Okada, Y., AND K. Suzuki. 1956. Taxonomic considerations of tlie lantern fish Pohjii)nii.s spinosiis and related species. Pacific Sci., 10: 297-302. Parr, A. E. 1937. Concluding report on fishes with species index for articles 1-7. Bull. Bingham Oceanogr. Coll., 3: 1-79. Pauca, M. 1931. Neue Fische aus dem Oligoziin von Piatra-Neamt., Bull. Acad. Roumaine Bucarest, 14: 29-34. Pearcy, W. G. 1964. Some distributional features of mesopelagic fishes off Oregon. J. Mar. Res., 22: 83-102. Peres, J. M. 1958. Remarques generales sur un ensemble de quinze plongees effectuees avec le Bathyscaphe F.N.R.S. III. In Resultats Scientifiques des Campagnes du Bathyscaphe F.N.R.S. Ill 1954-1957. V. Ann. Inst. Oceanogr. Monaco, N. S., 35: 259-285. Poll, M. 1953. Expedition oceanographique Beige dans les eaux cotieres Africaines de I'Atlantique Sud, 1948-1949. Resultats Scien- tifique. Poissons III. Teleosteens, Malacop- terygiens. 4: 1-390. Rass, T. C. 1955. Deep water fishes of the Kurile- Kamchatka Trench. Trud. Inst. Okeanol., 12: 328-339. Rechnitzer, a. G., and J. Bohlke. 1958. Ichthtj- ococcus irrcp,ularis, a new gonostomatine fish from the Eastern Pacific. Copeia, ( 1 ) : 10-15. Regan, C. T. 1908. Report on the marine fishes collected l^y J. S. Gardiner in the Indian Ocean. Trans. Linn. Soc. Zool. London, 12: 218. . 1923. The classification of the stonii- atoid fishes. Ann. Mag. Nat. Hist., 9(11): 612-615. Rivero, L. H. 19.34. Paces nuevos para la fauna Cubana. Mem. Soc. Cubana Hist. Nat., 8: 31. . 1936. Some new, rare, and little known fishes from Cuba. Proc. Boston Soc. Nat. Hist., 41: 56. RouLE, L., and F. Angel. 1933. Poissons prove- nant des campagnes du Prince Albert ler de 128 Bullet in Museum of Comparative Zoology, Vol. 142, No. 1 Monaco. Res. Camp. Sci. Monaco, 86: 1-115. RoxAS, H. A. 1934. A review of Philippine iso- spondvlous fishes. Philippine J. Sci., 55: 287. Samuel, C. T. 1963. Notes on deep sea fishes collected by R/V CONCH off the Kerala Coast. Bull. Dept. Mar. Biol. Oceanogr. Univ. Kerala, India, 1: 101-107. Sanzo, R. 1928. Uova, larvae, e stadi giovanili di teleostei. Comit. Talassogr. Italiano, Monogr. 2: 1-60. Sauvage, L. 1891. Histoire Naturelle des Pois- sons. In Grandidier, A. (ed.), Hist. Phys. Nat. Pol. Madagascar, 16: 483. ScHROEDER, E. H. 1963. Serial Atlas of the Marine Environment. Folio 2. North Atlantic Temperatures at a Depth of 200 Meters. New York, American Geographical Society. ScHULTZ, L. P. 1937. A new species of deep sea fish, Argijropclecus antrorsospiniis, of the family Sternoptychidae. Smith. Misc. Coll., 91(27): 5-8. . 1938. Review of the fishes of the genera Polyi))nu.s and Argyropelecus (Family Ster- noptychidae) with description of three new species. Proc. U. S. Nat. Mus., 86: 135- 168. . 1961. Revision of the marine siKer hatchetfishes ( Family Sternoptychidae ) . Proc. U. S. Nat. Mus., 112: 587-649. 1964. Family Sternoptychidae. 7m Fishes of the Western North Atlantic. Sears Found. Mar. Res., 1(4): 241-273. Scott, W. B. 1965. A record of the hatchetfish Polyipnus asteroides Schnltz from the Cana- dian Atlantic region. J- Fish. Res. Bd. Canada, 22: 1303. Smith, J. L. B. 1953. The Sea Fishes of Southern Africa. South Africa, Central News Agency, Ltd. 550 pp. Suzuki, K. 1964. Two rare deep sea fishes from Kumano-Nada. Rept. Fac. Fish. Pref. Univ. Mie, 5: 1-8. Suzuki, O. 1961. Mechanical analysis on the working behavior of midwater trawl. Bull. Japan Soc. Sci. Fish., 27: 903-907. SvERDRUP, H. U., M. W. Johnson, and R. H. Fleming. 1960. The Oceans: Their Physics, Chemistry, and General Biology. 9th Ed. Englewood Cliffs, N. J., Prentice-Hall, Inc. Ta\xor, R. W. 1967. An enzyme method of clearing and staining small vertebrates. Proc. U. S. Nat. Mus., 122: 1-7. Weber, M., and L. F. de Beaufort. 1913. The Fishes of the Indo-Australiau Archipelago. Vol. 2. Leiden, E. J. Brill. 428 pp. Weitzman, S. H. 1967a. The osteology and relationships of the Astronesthidae, a family of oceanic fishes. DANA Rept., 71: 49-52. . 1967b. The origin of the stomiatoid fishes with comments on the classification of sal- moniform fishes. Copeia, ( 3 ) : 507-540. Wheeler, A. 1969. The Fishes of the British Isles and North-west Europe. London, Mac- millan. 613 pp. Whiteley, G. p. 1940. Illustrations of some Australian fishes. Australian Zool., 9: 404. Wilimovsky, N. J. 1951. The correct generic and family name of the deep sea hatchet- fishes (Stemoptyx and Sternoptychidae). Copeia, (3): 247-248. Wooster, W. S., and J. L. Reid, Jr. 1963. East- ern boundary currents. In Hill, NL N. (ed.), The Sea. New York and London, Inter- science Publishers, 1: 253-279. Zugmayer, E. 1911. Poissons provenant des campagnes du yacht Princesse Alice ( 1901- 1910). Res. Camp. Sci. Monaco, 35: 1-174. ulletin OF THE - Museum of omparative Zoology The Carabid Beetles of New Guinea. Part IV. General Considerations; Analysis and History of Fauna; Taxonomic Supplement p. J. DABLINGTON, JR. HARVARD UNIVERSITY VOLUME 142, NUMBER 2 CAMBRIDGE, MASSACHUSETTS, U.S.A. 18 OCTOBER 1971 PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Bulletin 1863- Breviora 1952- Memoirs 1864-1938 JoHNSONiA, Department of Mollusks, 1941- OccAsiONAL Papers on Mollusks, 1945- Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint, $6.50 cloth. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. $9.00 cloth. Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth. Lyman, C. P., and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian Hibernation. $3.00 paper, $4.50 cloth. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12-15. (Price list on request. ) Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae ( Mollusca : Bivalvia ) . $8.00 cloth. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. $6.75 cloth. Proceedings of the New England Zoological Club 1899-1948. ( Complete sets only. ) Publications of the Boston Society of Natural History. Authors preparing manuscripts for the Bulletin of the Museum of Comparative Zoology or Breviora should send for the current Information and Instruction Sheet, available from Mrs. Penelope Lasnik, Editor, Publications Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. © The President and Fellows of Harvard College 1971. THE CARABID BEETLES OF NEW GUINEA. PART IV. GENERAL CONSIDERATIONS; ANALYSIS AND HISTORY OF FAUNA; TAXONOMIC SUPPLEMENr p. J. DARLINGTON, JR. CONTENTS Introduction to Part IV. 133 1. Purpose; previous parts; acknowledg- ments 133 2. Sources, disposition, and adequacy of material 133 3. Preparation for work on New Guinean Carabidae; my collecting 135 4. Basic literature 136 5. Localities 138 Policies and Methods 143 6. Modern taxonomy 143 7. Types 146 8. Taxon concepts: subfamilies and tribes 147 9. Genera 147 10. Species 148 11. Subspecies 149 12. Nature of taxonomic characters 150 13. Secondary sexual and genitalic char- acters 151 14. Preservation of material 152 15. Taxonomic methods and procedures __ 153 16. Data sheets 154 Analysis and Discussion 156 17. New Guinea 156 ^The author suggests that, when the four parts of TJie Caiabid Beetles of New Guinea are bound together, the present part be divided, and the whole bound in the following order: general con- siderations, analysis, and history of the fauna from Part IV; Part I; Part II; Part III; and the taxonomic supplement from Part IV. ■ Work and publication supported by National Science Foundation Grant GB-12346. Bull. Mus. Comp, 18. The New Guinean carabid fauna: tax- onomic composition 157 19. Numbers of species 158 20. Size of individuals 161 21. Wings and wing atrophy 165 22. Explanations of wing atrophy 170 23. Summary of wing state and wing atrophy 172 24. Ecology: habitats 172 25. Ecologic composition 173 26. Altitude 175 27. Ants : 176 28. Ecologic interactions 176 29. Ecologic ranges 177 Zoogeography: Existing Geographic Patterns .. 178 30. Geographic patterns, relationships, and origins 178 31. Existing geographic relationships: problems and procedures _. 178 32. Distribution of carabid tribes from Asia to Australia 180 (Cicindelinae) _ 182 33. Ozaenini 183 34. Paussini 183 35. Scaritini 183 36. Bembidiini 184 37. Trechini 185 38. Panagaeini 186 39. Pterostichini 186 40. Agonini 188 41. Perigonini 189 42. Licinini 189 43. Chlaeniini 190 44. Oodini 190 45. Harpalini 190 46. Anaulacini 192 47. Gyclosomini 192 48. Lebiini _____ __. 193 49. Pentagonicini 196 50. Hexagoniini 197 Zool., 142(2): 129-337, October, 1971 129 130 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 51. Odacanthini -- -- 197 52. Dnptiiii 198 53. Zuphiini - 198 54. Ilelluodini 198 55. Hclluonini _ 198 56. Brachinini _ .- 198 57. Pseudomorphini 200 58. Smiiinaiy of goM),c;raphic units 200 59. Cco^iaphic units by major habitats — . 201 60. Transitions of carabid faunas from Asia to Australia - 202 61. Wallace's Line and Celebes 202 62. Moluccas 203 63. Transition from New Guinea to Aus- tralia 203 64. Summary of transitions — - — . 207 65. Faunal regions — - 210 66. Geographic patterns within New Guinea 211 Zoogeography: Dispersals and Geographic Origins 216 67. The apparent main pattern of dis- persal 216 68. Dominance, competition, and extinc- tion in dispersal 216 69. Complexity of dispersal 217 70. Place of New Guinea in the main dis- persal pattern 217 71. Directions of dispersal ____ 218 72. Direction and vagility 218 73. Carabid versus mammalian dispersals 219 74. Directions of dispersal of tribes, gen- era, and species - 220 75. Summary of direction to this point _-_. 221 76. Direction and dominance 224 77. Direction and size 224 78. Direction and wings — _ 224 79. Direction and ecology 225 80. Direction and altitude: mountain- hopping across the Malay Archipelago 225 81. Direction and age 228 82. Australian-American discontinuities . .. 229 83. Summary of directions of dispersal .^ - 230 84. Barriers; filtering at Wallace's Line, Moluccas, Cape York 232 85. Amount of dispersal now and in the past 233 86. Summary of geographic origins of New Guinean Carabidae 236 E\olution 237 87. Evolutionary perspective 237 88. Evolution of New Guinean carabid fauna as whole _ 237 89. Relative age of the New Guinean fauna 239 90. Evolution of the mountain fauna 240 91. Evolution and adaptation of separate carabid stocks on New Guinea 243 92. Agonine radiation on New Guinea — 244 93. Evolutionary trends: not toward in- crease of size 245 94. Atrophy of wings, and associated trends ___ 246 95. Loss of setae ..._ 246 96. Modification of legs and tarsi 247 97. Modification of ecology and behavior 248 98. Parallelism and convergence; develop- ment of elytral spines; color patterns .. 248 99. Mimicry _ 250 100. Mutation and dimorphism 251 Taxonomic Section 254 101. Tribal classifications 254 Taxonomic Supplement .— 255 Cicindelinae 255 Carabinae - 255 Ozaenini 255 Scaritini 255 Harpalinae 258 Bembidiini 258 Trechini _ 262 Panagaeini 262 Pterostichini 265 Agonini 272 Perigonini 322 Licinini — 322 Chlaeniini 322 Oodini 322 Harpalini 324 Lebiini 325 Pentagonicini 330 Hexagoniini 331 Odacanthini 331 Zuphiini ( Leleupidiini ) ___ 332 Helkumini 333 Brachinini 334 Bibliography 335 In lieu of index 338 ABSTRACT This is the fourth and final part of an extended work on the Carabidae (predaceous ground beetles) of New Guinea. About 24,000 specimens from New Guinea have been examined, enough to show the general characteristics of the New Guinean carabid fauna, although many details remain unknown. Collecting done in New Guinea and in adjacent eastern Australia, the collections accumulated at the Museum of Comparative Zoology, and preparations for their study are described; working collections of New Guinean Carabidae for future use are deposited in the MCZ, the British Museum (Natural History), the Bishop Museum in Honolulu, and with CSIRO ini Canberra, Australia. Localities are mapped, and those where most Carabidae have been found The carabid beetles of New Guinea • Darlington 131 (Dobodiira, Wau, Mt. Wilhelm and vicinity, and the Snow Mountains ) are briefly described. Then discussed are modern taxonomic methods in rela- tion to those of the past; the continuing usefulness of types; concepts of tribes, genera, species, and subspecies; the nature of taxonomic characters; and the relative importance of secondary sexual and genitalic characters in carabid classification. Methods and procedures of second-stage faunal taxonomy (of which this work is an example) are discussed, with emphasis on the limitations imposed by time and available material. Under "Analysis and discussion," the immense, tropical, mountainous island of New Guinea is described briefly, and its carabid fauna is de- scribed and analyzed. The taxonomic compo- sition of the fauna is summarized. The number of species of Carabidae now known from New Guinea is 667, of which 434 occur in the low- lands, below 500 m altitude, and 376 in the mountains, above 1000 m; 161 of these species occur in both the lowlands and the mountains; and 18 additional species are iniknown as to altitude. At one lowland locality ( Doliodura, Papua), 217 species have been found; at one mid-altitude locality (Wau and vicinity, North- east New Guinea, 1000-2000 m), 170 species; but numbers of species decrease sharply at higher altitudes. In size, New Guinean Carabidae are small, rarely as much as 25 m in length. The lowland fauna is bimodal in size distribution, species 2.00-2.95 and 6.00-6.95 mm long being most numerous, with a deficiency of species especially at 4.00-4.95 mm. This bimodality is a result of an apparently recent arrival in New Guinea of many small Tachtjs, which have im- posed a second mode on an otherwise unimodal fauna, but the bimodality may reflect also a vulnerability of 4 to 5 mm carabids to compe- tition with ants. The mountain carabid fauna of New Guinea is unimodal in size distribution, with tlie mode at 9.00-9.95 mm. Almost ,all lowland Carabidae in New Guinea are winged, but incidence of wing atrophy increases with altitude, reaching 95 per cent on the highest mountain tops. The causes of flightlessness at high altitudes are complex; blowing or straggling away of winged individuals is apparently unim- portant, for most of the flightless species live in dense, wind-free montane forest; limitation of irea on mountains, by requiring carabid popu- lations to be dense and stable, may be the most mportant single factor in reducing the value of light and inducing wing atrophy. Ecologically, he New Guinean carabid fauna at low altitudes •onsists of roughly one-Uiird mesophiles (ordinary I ground-living forms), one-third hydrophiles (as- ociated witli standing or running water), and >ne-third arboreal forms. Altitude affects the carabid fauna in several ways: in taxonomic com- position (Agonini become predominant), in num- ber of species (fewer at higher altitudes), in size distribution (see above), in wing state (see above), and in ecologic composition (relatively more mesophiles at higher altitudes), but size of individuals in specific stocks apparently does not decrease with altitude; altitude probably exerts its effects partly directly and partly indirectly, ])y modifying vegetation and limiting areas and habitats. Ants probably compete with and re- place especially flightless mesophile Carabidae at low altitudes in New Guinea. Under "Zoogeography," existing geographic patterns are first discussed, witli emphasis on problems and procedures. The distribution of Carabidae in the Asia-New Guinea-Australian area is described by tribes, and the relationships of the New Guinean carabid fauna are expressed in "geographic units" (separate geographic re- lationships). The totals are 173 Oriental to 120 Australian units, giving a ratio of approximately 3/2 Oriental/Australian relationships; this ratio holds (roughly) for all ecologic fractions of the New Guinean carabid fauna except in opener country in southern New Guinea, where Austra- lian elements are more numerous. Change of carabid faunas from Asia to Australia is sum- marized; change in ratio of Pterostichini/ Agonini is most striking: tliese tribes are approximately equally represented in the Oriental area (Java), but Agonini are overwhelmingly dominant in New Guinea, and Pterostichini overwhelmingly dominant in Australia; this reversal of dominance has complex historical and ecologic causes. The three principal barriers to carabid dispersal be- tween Asia and Australia seem to have been of different sorts: an old ivater gap between Borneo and Celebes ( Wallace's Line ) , a bottleneck in the Moluccas caused by the relatively small size of the islands, and ecologic barriers between New Guinea and Australia even when there was a land connection. Although New Guinean Carabidae ( and some other insects ) are more Oriental than Australian in relationship, entomologists should not put New Guinea in the Oriental Region but should accept the conventional regions, perhaps adding New Guinea to the transition area ( an extended Wallacea) between tlie Orient and Australia. Within New Guinea, some Carabidae are restricted to the western end or to the southern edge of the island (suggesting that they are recent arrivals from the Orient or from Australia), and patterns of complex differenti- ation of subspecies, species, and genera exist; but no special centers of speciation and no specially important barriers to dispersal at low altitudes within the island are indicated. Endemic genera are concentrated toward the eastern end i 132 BiiUetin Museum of Comparative Zoology, Vol. 142, No. 2 of the island; tliis may l)e a result of invasion of western New Ckiinea by Oriental stocks arri\ing from the west. Complex patterns of differenti- ation and local radiation occur on separate moun- tain ranges. The dispersals that have made the existing patterns are discussed under "Dispersals and geographic origins," with emphasis on the prob- able importance of dominance, competition, and extinction, and on complexity. Apparent direc- tions of dispersal of tribes, genera, and species of Carabidae represented in New Guinea are sum- marized. Carabid dispersal in this region is com- pared with that of mammals, and dispersal is related to dominance, size of insects, wings and flight, ecology, altitude, and the relative time of dispersal of different carabid groups. Con- clusions in any single case are tentative, but all the cases together form a pattern of multiple dispersals mainly from Asia toward Australia. Over short distances the preponderance of east- ward and southward over northward and west- ward movements has not been great, but over the route as a whole movements from Asia and the Oriental area to New Guinea and Australia ha\'e apparently been several or many times more numerous than movements from Australia and New Guinea to the Oriental area and Asia. Cara- bids have apparently been coming into New Guinea in numbers for a considerable time. There is no good evidence that arrivals have been more numerous at some times than at others; arrivals have been very numerous recently, but faunal overturns with extinctions may have obscured the evidences of earlier arrival rates. Both Oriental and Australian stocks have probably reached New Guinea at all times, but (except in the relatively dry, open areas of southern New Guinea) in- coming Oriental stocks have apparently always been more numerous than Australian ones re- gardless of size of insects (almost all were small), regardless of wing state (almost all were winged), in all main habitats, and at all altitudes ( most were lowland forms ) . Continual extinc- tions have probably been correlated with the arrivals, and the extinctions ( "withdrawals" ) too have probably tended to begin at the Oriental end of the area and proceed toward Australia. This general history accounts for the nature of the New Guinean carabid fauna as a whole and perhaps for the distrilmtion patterns of "mountain- hopping" groups and for Australian-American dis- continuities in some cases. The trend of dispersals and extinctions from Asia toward Australia fits into an apparent world-wide pattern of evolution of successive dominant groups of Carabidae in the great, climatically favorable area of the main Old-World tropics and of successive dispersals into smaller and/or less favorable areas. And the ap- parent pattern of o\erturn of the New Guinean carabid fauna itself and the pattern of dispersal of Carabidae from New Guinea to the smaller islands to the east fit the MacArthur-Wilson theory of overturn and faunal etiuilibrium on islands. The New Guinean carabid fauna thus fits into and connects both world-wide and local dispersal and equilibrium patterns in a very satisfying way. As to evolution, the New Guinean carabid fauna as a whole has evolved from an ancestral accumulation of relatively unspecialized, small, winged, vagile ancestors selected by dispersal across barriers and including relatively numerous species living in water-side habitats; carabids in water-side habitats tend to be vagile, and these habitats are less widely interrupted by climatic factors than rain forest is. The ancestral accumu- lation probably did not have a single starting time but has been an evolving continuum, added to by arrivals and subtracted from by extinctions from time to time. The existing segment of this continiumi is relatively recent; the New Guinean carabid fauna is more recent in its origins than the faunas of tropical Asia, Australia, New Cale- donia, or New Zealand; this may be because faunal overturn has been more rapid on New Guinea. From this changing continuum different Carabidae have evolved on New Guinea to different extents and in different ways; some multiplications of species and ecologic radiations are described. The principal general results have been to increase greatly the number and diversity of both ground-living and arboreal Carabidae in rain-forest, partly by ecologic shifts including shifts from water-side habitats to the forest floor and partly by nuiltiplications of species, and especially to form on the higher mountains a complex alticoline fauna which, in ecology and in superficial adaptations (including wing atrophy), is like the carabid faunas of mountains elsewhere but which consists largely of genera and species apparently derived independently from lowland ancestors on New Guinea. The evolution of the mountain fauna, evolution and adaptation of separate carabid stocks, and radiation of Agonini on New Guinea are discussed in more detail. Evolutionary trends do not include increase of size of individuals on the island but do include trends toward atrophy of wings and associated structures especially on mountains, loss of setae especially on mountains, and modification of legs and tarsi especially on mountains. Parallelism or convergence have occiured among New Guinean' Carabidae not onl>' in atrophy of wings, loss of setae, and modifications of tarsi, but also in modifications of eyes, modification of body form, development of ventral pubescence ( especially in diverse Agonini), development of elytral spines The carabid beetles of New Guinea • Darlington 133 (which have evolved in 40 apparently separate stocks of carabids in New Guinea), and in some elytral color patterns. These parallelisms and convergences seem to inxolve in different cases parallel adaptations to montane environments (wing atrophy, etc.), adaptations to specific environmental details, mechanical protection (ely- tral spines), mimicry (some color patterns), and occurrence of homologous or parallel mutations (some other color patterns). Occurrence ot dimorphism apparently resulting from mutation among New Guinean and other Carabidae is de- scribed, and the relation of mutational changes to taxonomic characters is discussed. In the "Taxonomic section," tribal classifications of Carabidae are briefly referred to (but no new classification is offered), and a Taxonomic sup- plement lists important new records and new species. Seventy-three new species are described, most in the tribe Agonini, and most from high altitudes. No new genera are described, but the Oriental Phijsodcra and Omohms (both in tribe Lebiini) are recorded from New Guinea for the first time. INTRODUCTION TO PART IV [1]' Purpose; previous parts; ac- knowledgments. This is the fourth and final part of my work on l:)eetles of the family Carabidae of the island of New Guinea. The first three parts (see Bihli- ographij at end of present part) were pri- marily taxonomic. Part I (1962) covered the Cicindelinae ( tiger beetles, which were treated relatively superficially) and the tribes of Carabidae proper from the be- ginning through the Pterostichini in the order of the Junk-Schenkling Coleopt&ro- rum Catalogus (Horn, 1926; Csiki 1927- 1933). Part II (1952), which was published before Part I, covered the tribe Agonini, which is dominant in New Guinea and in which I have a special interest; my intro- duction to Part II went beyond taxonomy to discuss the general nature and evolution of the New Guinean agonine fauna. Part III (1968) covered the remaining tribes of Carabidae, from the Perigonini through To avoid excessive insertions in page proof, I nave assigned numbers to successive items, and shall use these rather than page numbers in cross references. the Pseudomorphini. The present part (Part IV) is a general summary, analysis, and discussion of the New Guinean carabid fauna as a whole, with a taxonomic supple- ment. This part can be divided: working copies of "The Carabid Beetles of New Guinea" can be bound with the general portion of Part IV first, then Parts I, II, and III in order, and finally the taxonomic supplement, bibHography, and statement in Heu of index of Part IV. See this statement (p. 33(S) for suggestions to users. The present part actually begins with a review of material used in my work, of the history of work on New Guinean Carabi- dae, of localities (with new maps), and of my methods of work and taxonomic con- cepts. Then follow analysis and discussion of results of the work from several points of view: numbers of species, size of insects, state of wings, taxonomic composition, ecologic composition, existing geographic patterns, and origin and evolution of the fauna. Because I am a biologist (as every taxonomist should be and many are), the analysis of the fauna as a whole has been, for me, the most exciting part of my work with New Guinean Carabidae. And be- cause I am a biogeographer, the geographic patterns and geographic histories and their significance have been most exciting of all, and I have treated them in greatest detail. I am indebted for careful typing and other work done on the manuscript to Miss Wilmoth Peairs; for drawing done patiently under my direction to Mrs. Mary Catron and Mrs. Sarah Landry; and for support both of the work while in progress and of publication of it, to the National Science Foundation (Grant GB-12.346). [2] Sources, disposition, and adequacy of material. The principal sources of material used in my work on New Guinean Carabidae are listed in Part II, pp. 90-91; Part I, p. 323; and Part III, pp. 2-3. I have prepared (on cards) a consohdated list of all the museums and other institu- tions and all the persons from whom material has been received, and of the 134 BuUctin Museum of Comparative Zoology, Vol. 142, No. 2 names of collectors, but the list includes more than one himdred items and seems to me not worth the space and cost of publication. The names of pertinent mu- seums, etc., and of collectors are given under the separate species throughout my work. I need say only that I am deeply indebted to the persons there named and to the authorities of the museums and other organizations concerned. And I should add that useful material has been received, but mostly too late to be included in my work, especially from Mr. and Mrs. G. W. Cot- trell and Mr. Fred Parker. Tlie kind of work that I have been doing with New Guinean Carabidae re- quires and receives international cooper- ation on a scale which persons who are not taxonomists do not always appreciate. I haxe received specimens or information about New Guinean species not only from many sources in the United States in- cluding Hawaii, but also from Canada; England, France, Belgium, Italy, Czecho- slovakia, and Hungary; Japan and Java; Australia (several persons in different cities) and New Zealand; and New Guinea itself and the Solomons. And basic ma- terial or information that has formed part of the background of my New Guinean work has come also from persons in Sweden, Germany, Switzerland, and India. In general, borrowed material has been returned to the sources from which it was received, with duplicates kept for the Museum of Comparative Zoology (MCZ). In a few cases additional important specimens, including holotypes, have been deposited in the MCZ for safekeeping. I am especially indebted to Dr. J. J. H. Szent-Ivany for the Department of Agriculture, Port Moresby, and to Dr. R. W. Hornabrook for permitting holotypes from their ma- terial or from material in their charge to be deposited in the MCZ. This arrange- ment safeguards irreplaceable specimens and makes them more easily available to specialists, and is a real contribution to future work on the carabid beetles of New Guinea. On the other hand, duplicates of my owTi material have been widely distributed. I have tried especially to build up the best possible working collections of New Guinean Carabidae in the British Museum; the Bishop Museum in Hawaii; and with the Commonwealth Scientific and Indus- trial Research Organization (CSIRO), in Canberra, Australia. For further discussion of the place of types and of working col- lections of specimens in modern taxonomy see [6, 7]. The number of specimens of Carabidae proper (excluding Cicindelinae) actually recorded from New Guinea in Parts I-IV is about 22,500. In addition I have seen perhaps 1000 or 2000 additional specimens from New Guinea without counting them ( an exact accounting has not seemed worth the trouble), and of course I have seen many thousands more from the Oriental Region and from Australia (see Part I, pp. 325-328). Most of the specimens from New Guinea were collected during or after the war, and most of them have exact localities, often altitudes, dates, and names, of collectors. About 8000 of the New Guinean specimens were collected by my- self (see [3] ). I have at least a rough idea of the habitats of most of the species that I obtained, and some specimens collected by other persons have indications of habi- tat. Many of the recently collected speci- mens were taken at light; these probably flew, at night. However, there is often no way of kno\\'ing whether individuals taken at light came from forest or grassland or swamps, or whether tht>y lived on the ground or were arboreal. All this material is adecjuate to show the general nature of the New Guinean carabid fauna as a whole. The material probably includes most existing primarily lowland species and good samples from a few mountains, although hundreds of mountain- living species localized at different levels, on different ranges, and on different peaks I The carabid beetles of New Guinea • Darlington 135 of the same ranges surely still remain to be discovered. In all, enough species are known from enough material to allow sig- nificant statistical analysis of the fauna as a whole. But the material of most single species is not sufficient for statistical study of either individual or geographic variation. Much more collecting and much more study will have to be done to make known the variation and geographic distribution of most species even at low altitudes, and the mountain-living species are much less well known. And, although so much still remains to be done on the structure, vari- ation, and distribution of the species, much more remains to be done in the insects' biology. About all that is known of the ecology e\'en of the best kno\\'n lowland species is their gross habitat and whether or not they fly to light, and not even this much is known of most high-altitude spe- cies. And the life histories of most New Guinean Carabidae are wholly unknowm, excepting only Pseudozaena and Morion (Gressitt, 1953) and a few genera and species of "Truncatipennes" which occur also in Japan and for which Habu (1967) gives biological notes. [3] Preparation for uork on New Guin- ean Carabidae: my collecting. Previous work on Carabidae of New Guinea is briefly described in Part I, pp. 324-328. Points emphasized and worth repeating are the small amount of work done on actual New Guinean specimens in the past, and the importance of work done on Carabidae of adjacent areas, especially by T. G. Sloane on the Australian fauna and by H. E. An- drewes on the fauna of the Oriental Region including the western part of the Indo- Australian Archipelago. Sloanc's and An- drewes' collections (which I ha\'e seen and studied) and the descriptions and revisions published by these two persons are the hasis for study of the relationships and history of the New Guinean carabid fauna. My own interest in and work on Carabi- dae of the Oriental Region and Australia IS well as of New Guinea, and the col- lections accumulated at the Museum of Comparative Zoology, are briefly described too in Part I, pp. 325-328. As a result mainly of my own activities beginning in 1931-1932 (when I was a member of the Harvard Australian Expedition), the MCZ now possesses a good working collection of Carabidae not only from New Guinea but also from both major source areas ( Oriental and Australian) from which the New Guinean fauna has been mainly derived. Although my collecting is summarized in Part I (pages cited above), a few addi- tional details are worth giving here. In collecting around Dobodura I was first struck by the very slow rate at which species accumulated. The first day I found, I think, only three or four species, and few individuals. The next day I found perhaps two or three additional species. And so forth. But when I came to sort out the species and study them at the MCZ, I found that I had obtained 217 species at this one diverse but strictly lowland lo- cality! The slowness with which species accumulated was, I think, due only partly to a temporary physical handicap of mine. It was probably due partly to the fact that, although species are diverse in the tropics, many have sparse populations and (even in relatively small, ecologically homo- geneous areas) patchy distributions (Wil- son, 1958). My impression is that populations of Carabidae tend to be most sparse and most scattered on the rain-forest floor. Collect- ing in the leaf litter and loose soil on the floor of rain forest does in fact call for patience and ingenuity if the diverse fauna which lives there is to be adequately sampled. Old-fashioned sifting handles too little material to yield an adequate sample of the thinly dispersed fauna in a short time. Berlese funnels are better and are indispensible for collecting some insects, but are probably still too slow to obtain Carabidae in adequate numbers — and I had no Berlese equipment. The method that I did use, and that I 136 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 think is most effective in rapid sampling of rain-forest-floor Carabidae, is what I call washing or drowning. It involves rak- ing or scraping up large quantities of leaf litter and loose surface soil and throwing them into still water where the light is good. The debris is spread out on the water surface, thoroughly wetted and gently stirred, and perhaps stirred again at intervals. The Carabidae, of course, came to the surface of their own accord, and usually run across the floating debris toward the shore, where they are picked up by the waiting collector. Some take flight and must be caught the instant they appear. Others run down the floating debris again if they are alarmed on the surface. And some, especially some fos- sorial forms, come to the surface only after considerable delay. Sometimes the col- lecting is complicated by small birds and lizards, which wait close by and make dashes for moving insects. The collector who uses this method for the first time is likely to be disappointed in it. It requires judgment and experience in selecting the right places and in raking up debris in the right way. However, properly used, it yields good collections even from material scraped from ordinary, thin, rain-forest leaf litter. And the yield is sometimes multiplied in washing out piles of leaves that have been concentrated by some natural means, for example under the head of a tree that has fallen and from \\hich the dead leaves have dropped thicklv on the ground. Flash floods in the rain forest, which wash masses of leaves and other debris together, are best of all. provided the collector can \\ork them im- mediatelv, while the water is rising or at least before it starts to fall, before the Carabidae have had time to redisperse and before ants and other predators have had time to decimate them. A flash flood at Dobodura, which brought down the bank of a gully and dammed water back over a piece of flat ground in heavy rain forest, gave me the best collecting I had in New Guinea. In it, I found (I think) all my Oclontomasoreus humcraU.s (Part III, p. 76) at Do])odura, all my Nototarii,s papua (Part III, p. 186), my only specimen of Cola.sidio papua (present part, Tax. suppL), probably two of my three specimens of Phcropsophus catuhis (Part III, p. 328), and series of other ground-living carabids that I rarely found at other times. One additional note: at Dobodura, as elsewhere in the tropics, virtually no Carab- idae were found under stones on the ground. Under stones is, of course, where northern collectors first look for Carabidae, and failure to find them there in tropical rain forests has perhaps contributed to the idea that Carabidae are scarce insects in the tropics. But the Carabidae are there in numbers and diversity — just not under stones (see following paragraph). On the Bismarck Range and Mt. Wil- helm [5] my time was so limited that I got only a skimpy sample of the carabid fauna. I was impressed by the fact that at these altitudes (c. 2000 m and higher), Carab- idae did commonly occur under stones (cf. preceding paragraph). In fact some of the same species that in my experience; were never found under stones at low alti- tudes were found there in the Waghi Valley on th(> Bismarck Range. It therefore seems not that the requirements of tropical Carabidae are different from those of temperate ones, but that the microclimate or microhabitat under stones in the full tropics is somehow inhospitable to most Carabidae. Of my collections made outside New Guinea (Part I, pp. 327-328), the most im- portant is a comprehensive collection of wet-forest Carabidae made along the wholcj eastern edge of Australia, from northern Cape York to southern Tasmania, during 19 months in 1956-1958 (Darlington. 1960). [4] Basic literature. Work done on any fauna is carried over from generation to generation in the form of collections, publi- cations, and sometimes unpublished manu- The carabid beetles of New Guinea • Darlington 137 scripts. Collections of New Guinean Carab- idae available for future work are noted elsewhere [2]. Publications and manu- scripts essential to or resulting from work- on New Guinean Carabidae are listed in more detail in the BihJiopaphy (pp. 334- 337 ) and under many of the species treated in Parts I-IV, but the more important items may usefully be summarized here. Basic to work on Carabidae in any part of the world are the Junk-Schenkling Coleoptewrum Catalogtis (Horn, 1926; Csiki, 1927-1933) and the Zoological Record. The former (often called the "Junk Catalogue") lists all Carabidae of the world up to within a year or two of the dates given, with very few omissions and not many errors of citation, and with bibliographies and indications of distri- bution, both sometimes incomplete. This work is still available from W. Junk, Pub- lisher, 13 van Stolkweg, The Hague, Netherlands. The annual volumes of the Zoological Record list additional papers and new genera and species published from year to year, and enable students to com- pile preliminary Inbliographies and pre- liminary faunal lists. Basic works on the Carabidae of the Oriental Region and Indo-Australian Archi- pelago include H. E. Andrewes' papers (see Part I, p. 325) and especially his (1930) "Catalogue of Indian Insects, Part 18, Carabidae," which in many genera lists all species known from the Indo-Australian Archipelago including New Guinea; Lou- werens' papers, especially his (1953) re- vision of Oriental Colpodes; studies by the late Amost Jedlicka and by Akinobu Habu, especially their long papers on Oriental "Truncatipenncs" (see Bibli- ography); and work now in progress by Shun-Ichi Ueno especiallv on Trechini. Also useful to future workers should be my manuscript list of Carabidae of the Indo-Australian Archipelago including New Guinea, based on the Coleopterorum Catalogtis and the Zoological Record, but amplified and brought up to date. This manuscript is not prepared for publication, but I expect to deposit Xerox copies of it in the Department of Entomology at the British Museum, in the Bishop Museum in Honolulu, and with CSIRO at Canberra in Australia. The original manuscript will, of course, be kept at the MCZ. Basic work on Australian Carabidae has been done principally by T. G. Sloane ( see Part I, pp. 324-325), who published also two short papers ( 1907 ) on New Guinean species. Work done and being done on Australian Carabidae by B. P. Moore should also be followed by those interested in the New Guinean as well as the Austra- lian faunas. My own papers on certain groups of Australian carabids may be useful in some cases, especially since they are related to my New Guinean work; papers on my Australian collecting locali- ties ( 1960 ) and on transition of wet-forest carabid faunas from New Guinea to Tas- mania (1961) may be especially useful. My manuscript list of Australian Carabi- dae, based again on the "Junh Catalogue" and Zoological Record but amplified by search of all Sloane's papers and much other literature, may save time for later workers; a copy of it will be deposited with CSIRO at Canberra. Works on Pacific Carabidae that should be considered in studies of New Guinean species include H. E. Andrewes' (1927) paper on Carabidae of Samoa; E. C. Zim- merman's "Insects of Hawaii," especially his introductory volume (1948) and his projected volume on Hawaiian Carabidae; and other work done and being done at the Bishop Museum or on Bishon Museum material, including my paper (1970) on the Carabidae of Micronesia. As to New Guinea itself, my "The Cara- bid Beetles of New Guinea," of which the present paper is Part IV, coordinates earlier work and should be the basis for future work on New Guinean carabids, especially for third-stage taxonomic study of selected groups (see Part I, pp. 328- 330 ) . Three volumes that will be important 138 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 lOS-E 3rr ISO-E ASIA TRORC of CANCER EQUATOR '1 ..-•-'■ .t.\^^^^ ^New Britain t\ 15' S 195' E Solomon t^'' Island's TROPIC of CAPRrcORN j^ New ?>ji Coledonio FijiV ISO"^^ Figure 1. Map to show relation of New Guinea to Asia and Australia. Broken lines (at c. 100 fathoms depth) show ap- proximate limits of continental shelves and some other areas of shallow water which were probably land at times in the Pleistocene. to entomologists working on New Guinean Carabidac and other insects are in press or are planned by J. L. Gressitt and J. J. H. Szent-Ivany (joint authors). The first (now published, 1968) is a bibliography of New Guinean entomology. The second, now being planned, is a history of entomological exploration in New Guinea, with a list of localities. And the third, also being planned, is on the environment of New Guinea from an entomological standpoint. (This in- formation is from a letter from Dr. J. L. Gressitt dated December 26, 1967.) [5] Localities: Ideally, I should like to map all New Guinean localities at which Carabidae have been collected, but this has proved beyond my power. I can, how- ever, present the following new maps, pre- pared by Miss Sally Babb (now Mrs. Joseph Landry) under my direction. Figure 1 is a small-scale orthographic map de- signed to show the relation of New Guinea to other land areas from southern Asia to northern Australia. Figure 2 is a map of New Guinea as a whole showing general features of the island, some localities, and outlines of limited areas which are mapped in more detail. And Figures .'3-5 are more detailed maps of parts of New Guinea in which important carabid localities are too numerous to show on Figure 2. Many New Guinean localities are spelled in different ways by different authorities, but I cannot list alternative spellings here. Some lo- calities are put in slightly different places on different maps and by different gazet- ' teers; some margin of error should there- fore be allowed for in using my maps. And where detailed localities are too crowded to distinguish, I have shown only general The carabid beetles of New Guinea • Darlington 139 J 1 I o ■ a •* 1 ' o ' .,0 Q o i ^. Z _j - tr ;1j < A N_^ Q O S 8 UJ c- y D ^^ il^ /^ f "i ^oX i i i £ 19? 5 o ( zl -\^ y !1 r \ i \ UJ / Sq k ^ s Jz V o <^ 5 ) -^ ■ (/I o ° 3 22s II /^ ; >--! A • l^n° J. ^ y-' ^ IT ^1 /to t- ' ■' r o s ■ /7 I, o 0- ^ // Or O Or J »- ■* X^^: a: o M / ^ 5 ,' 1' ^ / ~ I « X s?t? r (Q ' « i^ ^ / 3 ej ^ 1/ i>i z / S Mp A = \r* 1 ^ ■ 4^ 1 / i? p\? ^_^ / *^ 2 !/ / \ f Q / c E ^ - V, 2 - " jJT^ F o • -J' UJ _^ ^ ( ^, yo oD o > o o * /^ V 1 / ^ ft ) 3 i > r [( z "1 ' ' . i. 3£ / o ^. J lO / "^ ;i'„ 1.^ "' ^ / r^^n^So oE ^ ^ 'Or^ Mr\ °ci , /-^ 2-, ?(r>^^ /-^"'^A, -^ O ?" .'>'* C^IO /e- <. ?^ '■■* o "'V 'tl.ss^ % T (^ _ »^-\_^ / o /o ^ / o UJ 7 15^ ) C f t/^ m V ^ O ) f < y 1 :f Hollandia). And later (in 1938-1939) ?he collected extensively in Waigeo and fapen Islands, and briefly in the Torricelli Mountains. Most of her localities will be found in the Bishop Museum list of locah- ies referred to above. A source of useful information on limited ireas in New Guinea is the series of re- ports of the Archbold Expeditions in the bulletin of the American Museum of SNOW MOUNTAIN RANGE ^^t J Scrte Volley WILHELMINA \ Comp Figure 5. Part of the Snow Mountain range. West New Guinea, showing especially the localities at which Carabidae were obtained by the Netherlands Indian-American (3rd Archbold) expedition of 1938-1939. Redrawn from Toxopeus, 1940. Natural History, including items by L. J. Brass (1941 and 1964) on the expedition to the Snow Mountains and on an expedi- tion to Mt. Wilhelm and vicinity and to the Lae-Edie Creek area (vicinity of Wau). Localities of the Archbold Expedition to the Snow Mountains have been published separately by Toxopeus (1940). See Gressitt and Szent-Ivany (1968) for references to accounts of New Guinea col- lecting by Biro, Cheesman, D'Albertis, Gressitt, MacLeay, Maindron, Szekessy 142 BiiUciin Mitscin)i of Comparotivc Z(X)logij, Vol. 142, No. 2 (concerning Biro), Szcnt-Ivany, Toxopeus, Wallace, Wilson, and others. I should at this point say something about altitudes. Exact or approximate altitudes of many localities are given on maps, or in the Bishop Museum list, or on locality labels on specimens. However, even modern altitude determinations are sometimes inaccurate, and the altitudes given on maps are not necessarily the alti- tudes at which collecting has actually been done. The risk of error is greatest in the case of material collected long ago. We are not justified in assuming that old speci- mens labeled (for example) Sattelberg came from the immediate vicinity of the town or from the same altitude. They may have been collected many miles away and many hundreds of meters above or below the altitude of the town itself. I have usually omitted old locality records in dis- cussing the distribution of New Guinea Carabidae in relation to altitude [26]. Several localities where especially im- portant collections of Carabidae have been made are \\'orth separate notice here. Dobodura and vicinity (which includes Oro Bay), Papua, are briefly described in Part I, pages 325-326, and a few more details are given in the present part [3]. It is a strictly lowland area; my collecting there probably did not extend above 200 meters altitude. But the area is otherwise diverse: habitats within easy walking dis- tance of Dobodura included heavy lowland rain forest, \'arious kinds of second growth, grassland, swamp margins, and the banks of streams including a small river, sluggish brooks, and rapid brooks in foothill topog- raphy. Important lowland habitats missing in this area were, I think, only special coastal ones, those associated with really large rivers, and those confined to the open eucalyptus country of southern New Guinea. My collection of Carabidae from Dobodura comprises 217 species [19] and more than 4,000 specimens. A second very important locality or group of localities is Wau, with Edie Creek, Mt. Mis(s)im, Mt. Kaindi, etc., in the Morobe District of North-east New Guinea. This is a diverse, mid-altitude area, indicated in Figure 3. Of this area, Gressitt (letter of December 26, 1967) says, "A description of the environment just above Wau is found on pages 182-185 of Volume 127 ( 1964) of the Bulletin of the American Museum of Natural History, by Brass. This is under the heading 'Kaindi, Morobe District.' Part of the area de- scribed in that section is what we call Edie Creek. And most of our material labeled Kaindi is from just above the area de- scribed in that section. Wau proper is just below these areas, at the foot of Kaindi, around the lower part of the Edie Creek Road. Several of our localities like Kunai Creek, Nami Creek, Delias Creek, are along the Edie Creek Road above our field station. "There is a tremendous range of vegeta- tion types from the bottom of the Bulolo Gorge just below Wau to the top of Mt, Kaindi and to the top of Mt. Missim, the higher mountain on the north side of the Valley (from which MCZ has some old material). Among the main differences ol the lower part of the valley with the ares described in Brass' paper is the fact thai Araucaria is dominant in many parts ol the valley to just about the level of oui field station. Also, palms and many othei tropical trees are in the lower forests, ex tending above the station. In Wau Vallej itself, there is a great deal of coffee grown Also, a lot of the Araucaria has been cu and pure stands of both species have beer planted after destruction of the remaining native forest. Still, there are extensivi areas under natural vegetation. And mon particularly so going higher on the tw( mountains." Brass's (1964) paper should be referrec to for further infonnation about the Wai area, especially its vegetation, anel fo photographs. Carabidae collected betweei 1000 and 2{X)0 meters altitude in this are now number 170 species, and 35 additions The carabid beetles of New Guinea • Darlington 143 pecies have been found in the vicinity ither at higher or at slightly lower alti- Lides. This is an outstandingly fine col- jction — a notable accomplishment by the iishop Museum entomologists using their ield station at Wau. My brief visit to and the collection made n the Bismarck Range and Mt. Wilhelm in )otober 1944 are briefly described in Part , pages 326-327, and Part IV ( the present >art) [3]. A sketch map of my route to it. Wilhelm is in Part I, p. 326, and ►resent Figure 3 shows the position of the lountain in relation to localities at which ther collectors have obtained Carab- iae more recently. Tlie altitude of Mt. Vilhelm, previously considered to be about 5,400 ft., is now considered to be about 4,600 ft. ( c. 4450 m ) . Specimens collected ly me in this area bear three different abels. Those from the lower, more open ountry are labeled "Chimbu Valley, Bis- narck Range, 5,000-7,500 ft.," and were aken in the densely inhabited valley, most I which has been highly modified by man, nd which is now largely grass or gardens, Ithough very limited habitats including he edges of small streams and of the Chim liver are still natural. Specimens taken in he montane forest, which changes from ain forest to moss forest with increasing Ititude. are labeled "Mt. Wilhelm, Bis- aarck Range, 7,000-10,000 ft.," and were aken on the ground in the forest, mostly mder stones and logs or beside running vater. Finally, specimens taken above the orest line, in "subalpine" habitats, are abeled "Mt. Wilhelm, Bismarck Range, hove 10,000 ft.," and were taken either in the ground vmder various kinds of cover acluding tussocks of grass, or beside small treams in tussock-grass country, or ( Macii- igomim oltipox only) in a grass tussock. or further information and photographs f the Bismarck Range and Mt. Wilhelm, ee Brass (1964) and Brookfield (1966: 79-183). Brookfield (pages cited) sum- larizes the geology as well as the egetation of the area. This range (like many of the other mountains of New Guinea, 1 think) is described as "a recent fold-structure which was uplifted in a series of stages culminating in the late Tertiary," with complexly faulted strata which include limestone. The localities in the Snow Moimtains, West New Guinea, at which L. J. Toxopeus obtained his fine collection of Carabidae, are briefly described by him (1940), with altitudes and very brief descriptions of the vegetation. I here reproduce Toxopeus' map, somewhat simplified (Fig. 5). His localities cover virtually all the important montane habitats from cultivated valleys below 2,000 m, through various types of forest at increasing altitudes, to "alpine" areas above timberline. The highest alti- tude at which collecting was done was 4,250 m, but "results were few" this high up. Evidence that "Dor(e)y" labels have been wrongly placed on many Carabidae, col- lected by Alfred Russell Wallace, that probably really came from Celebes or the Moluccas is given in Part I, pages 330-331, and Part III, page 5. Wallace did collect at Dor(e)y in West New Guinea, but he or someone else evidently labeled as from there many specimens which really came from other islands. POLICIES AND METHODS [6] Modern taxonomy. "Modem taxon- omy" means different things to different persons. To me, it means taxonomy as practiced now, and it is worth considering what the policies and methods of taxonomy now are. In general, I think modem taxon- omy, as compared with the taxonomy of one or two generations ago, is more care- fully calculated to reflect real situations in nature and more carefully designed for intelligibility and utility. Modem taxon- omy also employs new techniques and new procedures appropriate to the material and purpose of each particular piece of taxo- nomic work, but the new techniques and procedures are for the most part added to, 144 Biilhtin Museum of Comparative Zoology, Vol. 142, No. 2 not used in place of, the procedures of the past. Taxonomic work has therefore be- come increasingly complex and difficult, but also more precise and more useful. See Mayr (1969) for detailed discussion of the principles and procedures of this kind of work. And sec^ the volume on Systematic Biologij published by the Na- tional Academy of Sciences-National Re- search Council (1969) and current numbers of the journal Systematic ZooIo/.„ 1 iO-, -♦ t: o>--,. •/ /' ■ ■ ^'/.. ">j'.-. ;r X.' 1 f^t, :'^ir:-^.-- f'J.Jor- ^■0 J.vr +• (^..^ ^^' - ,■ 1. n 3r n 2-. 3-0 + •hi ^l..-A./../^, - ) it'.' 3H 1 . ifr: n / /:■,; -J. f\ ii.^!;*.;.:^" n<^. ', <,j :„ ^ A' 1 •'"v ■ . ■'■ / 1 J M,tr:. -,.-_: *.,v ../.^:., i" /!. -% / ■• . j_^i^, , ... 1 1,1 n A f. : - 7" /' .' ■'■■■ -h » "1 ('J "J Hi- 1 '^ s^ 1 '/ : + Ah le-'/- .'. f' /; ■ ' . .> 111. If 1 ^ i /J-L i //- 1 / ( * Uft! ' / ■ /-■ i-/ T^ /■/.' , \o. iv- /v.; ;'j(^/-;. ^.•^■.' + li r ,■ ■ 1 X h /t r*-^r,>„^ ... ; r/ •i" O + r ' j '•v.; J.: ■ + r./;,,. -\ ^7 n £ j-^rj--'^; 5. H.arabidae. Of the Greater Antilles, Hispaniola called also Haiti or Santo Domingo) is ,fiost nearly comparable to New Guinea, 'eing ecologically diverse and having ex- ensive mountains. New Guinea (roughly 800,000 sq. km or 300,000 sq. mi. in area) is about ten times as large as Hispaniola (roughly 80,000 sq. km or 30,000 sq. mi.). Hispaniola has a known carabid fauna of 148 species, of which 97 are lowland forms. By my rule of thumb. New Guinea should have a total of about 300 species, of which about 200 should be lowland forms. But figures given in preceding paragraphs show that New Guinea has in fact more than twice as many species as expected by this rule. The greater richness of the New Guinean fauna may be due partly to the greater ecologic richness and diversity of the island. New Guinea has very large areas of fine rain forest, several big rivers, ex- tensive swamps, etc., while the Greater Antilles have relatively little, relatively poor rain forest, no really large rivers, and few large swamps. The greater richness of New Guinea may be due in part also to the greater accessibility of the island, which was connected by land to Australia at times in the Pleistocene, and which many Carabidae have reached from the west too. The New Guinean carabid fauna is in fact continental in size and diversity. Satis- factory figures for numbers of species in continental areas of Asia and Australia are not available, so again I have to go to the other side of the world for comparisons. In 1943 (p. 41) I counted or estimated num- bers of species of Carabidae in several areas in eastern North America including New Jersey, Indiana, and North Carolina. Each of these states has more than 300 but less than 400 species of lowland Carabidae. Most of the species are widely distributed and are shared by all three states, so that the total number of lowland species in all three states together does not much exceed 400. This is considered a rich continental carabid fauna. However, the lowland fauna of New Guinea is evidently still richer — it must be remembered that even the lowland Carabidae of New 160 Bulletin Mttscttm of Comparative Zoology, Vol. 142, No. 2 Guinea are still much less well known than those of eastern North America. In 1943 (p. 41) I thought I detected a diminution of numbers of Carabidae from temperate North America into the Ameri- can tropics, although exact figures were not available. New Guinea can now be seen to have a carabid fauna rich out of all expectation for a lowland tropical area. Evidently Carabidae are very numerous in species in the lowland tropics at least in some regions. (But in proportion to the total insect fauna, Carabidae may still be less numerous in the tropics than in temperate areas.) However, they are evi- dently much more difficult to find in the tropics than in the north temperate zone, probably because they are more diverse ecologically in the tropics, and perhaps because the tropical populations tend to be sparser. (For further discussion of numbers of Carabidae in the tropics includ- ing New Guinea, and of the possible effect of competition with ants, see the present paper [27] and Fig. 11.) Mountain-living Carabidae, known above 1000 m (c. 3280 ft.) in New Guinea, now total .376 full species, of which 161 are and 215 are not known below 500 m. However, this is probably a small fraction of the total number of Carabidae existing on mountains in New Guinea. Until good collections have been made not only on different mountain ranges but also on successive peaks along single ranges, we have no basis for estimating the amount of geographic replacement of localized species that occurs from point to point in the mountains of New Guinea, and no basis for making a real estimate of total number. I can there- fore only guess, from the fragmentary collections available, that the total num- ber of mountain-living species of Carabidae in New Guinea will run to many hundreds, perhaps thousands, of species. The number of species of Carabidae oc- curring at a single locality at middle alti- tudes in New Guinea is surprisingly large. For example, 170 species have been taken between 1000 and 2000 m at and near Wau [5], in the Morobc District of North-east New Guinea. (Thirty-five additional spe- cies have been found in the same general area either below 1000 or above 2000 m.) The collections at Wau were made by persons (the Sedlaceks and others) who, although fine collectors, are not carabid specialists. Their collections are surely deficient in small ground-living species, al- though strong in arboreal ones and in light- trap material. The total number of Carab- idae existing at and near Wau betw'een 1000 and 2000 m is probably really con- siderably more than 200 species. Many of the species that occur at Wau occur also in the lowlands, but many others are either confined to the mountains or at least have not yet been found at lower altitudes; some are wide-ranging in New Guinea, others apparently confined to the general vicinity of Wau. At still higher altitudes in the mountains of New Guinea so little carabid collecting has been done that counts of species are hardly significant. However, the number of species at single localities obviously de- creases very sharply with increase of alti- tude. Incomplete samples (Table 2) show the Snow Mountains in West New Guinea with nine species above 3000 m, including only one above 4000 m, and Mt. Wilhelm in North-east New Guinea with seven species above 3000, including two above 4000 m. In summary of numbers of species: New Guinea has an unexpectedly large and diverse carabid fauna, large even by con- tinental standards. Of the total number of 667 species of Carabidae proper now known from the island, 434 have been found be- low 500 m altitude, and many of these are widely distributed on the island. At mid- altitudes (1000-2000 m) species are still numerous; some of them are widely dis-J tributed, others localized. At still higher altitudes, numbers of species at single lo- calities decrease sharply, but most of the high-altitude species are localized, and The carabid beetles of New Guinea • Darlington 161 Table 2. New Guinean Carabidae found above 3000 M (above C. 10,000 ft.) Table gives names, mean lengths, and altitudes of all Carabidae thus far recorded above 3000 m (above c. 10,000 ft.) in New Guinea. All species are (probably) ground-living mesophiles except Nebriagoniim percephahim, which is found beside mountain torrents, and Maculagonum altipox, which apparently lives in tussock grass. All species listed are flightless, with atrophied wings, except Maculagonum altipox, which has full wings. Snow Mts., West N. G. (9 species) Mectjclothorax ioxopei (4.7 mm), 4200 m Rliytiferonia nigra (19.75 mm), to 3300 m Analoma fortis (12.85 mm), to 3850 m Analoma gracilis (14 mm), 3800 m Gastragonum laevisculptum (8.3 mm), 3600 m Montagonum toxopeanum (9.0 mm), 3600 m Nebriagonum subcephalum (9.9 mm), 3300 m Chydaeus papua (9.9 mm), 3600 m Scopocles altus (3.4 mm), to 3800 m Mt. Wilhelm, N-E N.G. (7 species) Mecijclothorax sedlaceki (4.3 mm), 4250 m Maculagonum altipox (7.0 mm), above 3000 m ( winged, in tussock grass ) Nebriagonum cephalum (8.55 mm), to 4250 m (2 collections) Nebriagonum percephahim (9.9 mm), to above 3000 m ( beside running water ) Nebriagonum transitum (9.35 mm), to 3400-3500 m (2 collectors) Laevagonum subcistelum (5.95 mm), above 3000 m Chydaeus papua (9.9 mm), to 3400-3500 m (sev- eral collectors ) Mt. Albert-Edward, Papua (3 species) Analoma rosenburgi (12.6 mm), 4026 m Montagonum filiolum (10.5 mm), 3660 m Fortagonum antecessor (9.0 mm), 3660 m Mt. Giluwe, Papua ( 1 species ) Laevagonum giluwe (7.0 mm), 3750 m Mt. Amangwiwa, N-E. N. G. (1 species) Montagonum fugitum (11 mm), 3355 m much geographic replacement occurs, so that the total number of high-altitude Carabidae on the mountains of New Guinea is surely very great, although the final number can not yet even be guessed at closely. This situation is diagrammed in Figure 7. [20] Size of individuals. New Guinean Carabidae are small. Of 434 lowland spe- cies (exclusive of Cicindelinae), 388, or 89 per cent, have a mean length of 12 mm (c. V2 inch) or less, and only five species exceed a mean length of 20 mm. Of these five, three are characteristic, endemic New Guinean species : Lesticiis politiis ( 24 mm ) , Colpodes rex (21 mm), and Chlaenius pan (25 mm). The other two, still larger spe- cies are marginal or introduced: Gigadema maxilhre (32 mm) is an Australian species of which a single specimen has been found on the southern edge of New Guinea, and Catadromus tenehroides (mean 51 mm) is an Australian species of which two speci- mens were taken at military ports in New Guinea during the war (and which has been found in Java too). So, characteristic lowland Carabidae of New Guinea are all small, none more than an inch long, and most much less than that. Mountain-living Carabidae in New Guinea are small, too, none having a mean length of more than 22 mm (less than 1 inch). However, minute species (mean length less than 3 mm), although numerous in the lowlands, become relatively fewer with increasing altitude and disappear at highest altitudes. The few Carabidae known above 3000 m (Table 2) range from 3.4 to 19.75 mm mean length. Small size is a characteristic of the Carab- idae of some other East Indian islands including Celebes and the Philippines. (Sumatra, Java, and Borneo have a few larger Carabidae, notably species of Mor- molyce.) West Indian Carabidae are all small, too.i On Cuba, no carabid has a mean length of more than 25 mm (1 inch), and only four (2 species of Calosoma, a Scarites, and a Chlaenius,) exceed a mean of 20 mm; and on Hispaniola only the two Calosoma and possibly a Scarites exceed a mean of 20 mm. However, the situation is ''■ Since this was written, a very large, large- headed Scarites has been discovered in the moun- tains of eastern Puerto Rico (Hlavac, 1969). The first specimen found measured about 35 mm. It far exceeds in size any of die approximately 350 species of Carabidae previously known from the West Indies. 162 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 C 3000 M C 2000 M C 1000 M LOW LANDS Figure 7. Diagram of distribution of species in relation to altitude among New Gulnean Corabidae. Tfie diagram is not exactly quantitative, but each fiorizontal line, whether broken or not, represents c. 50 species; each unbroken vertical line, c. 10 species; and each broken vertical line, less than 10 species. Unbroken horizontal lines represent species that extend over c. the whole length of New Guinea; broken horizontal lines, species that are more localized. The diagram emphasizes that increase in altitude is accompanied both by decrease in number of species at single localities and by increasing locali- zation of species. strikingly different on Madagascar. The Carabidae of Madagascar include many very large forms especially in the tribes Scaritini and Pterostichini, some of the Madagascan scaritines being among the largest Carabidae in the world. The small size of New Guinean Carab- idae is not easy to explain. Mere existence on a large island does not necessarily favor smallness: witness the gigantic carabids on Madagascar. Nor does existence in rain forest necessarily favor smallness: witness Mecynognathus and the large Tricho- stermis in tropical rain forest in Australia ( following paragraph ) , Comparison of New Guinean species with the same or related species in Austra- lia and southern Asia indicates no general decrease of size on New Guinea [93] . New Guinean Carabidae are small because small rather than large Carabidae have reached and established themselves on the island. This suggests that difficulty of access has barred large forms, but this explanation is too simple or at least incomplete. New Guinea was connected to Australia at times in the Pleistocene. Many Australian Carabidae including many Scaritini and Pterostichini are large: one of the largest and finest Carabidae in the world, Mectj- nognathus dameli Macleay (mean length c. 50, maximum length 63 mm), is confined to a small area of poor rain forest on the tip of Cape York, just opposite New Guinea; other large Pterostichini. espe- cially species of Trichosterntis (Darlington, 1961), occur in the tropical rain forests at the base of the Cape York Peninsula; and large Carabidae occur in the drier parts of Australia too. Why did not large forms invade New Guinea when there was a land connection in the Pleistocene? Ecologic barriers evidently existed between Austra- lia and New Guinea even when land was continuous [84], but why were the bar- riers so effective against large carabids living in both rain forest and dry country?. Predation, by mammals and birds, does.i not explain the absence of large Carabidae in New Guinea. Insectivorous predators do occur there, but they occur also in Australia and Madagascar, where large The carabid beetles of New Guinea • Darlington 163 Carabidae are common. And on the West be due to increase of size, the increase Indies, where mammahan predators at having required a relatively long time least are relatively few, large Carabidae and relatively effective isolation, do not occur. The size distribution of New Guinean In spite of what has been said in the Carabidae is, unexpectedly, bimodal. A several preceding paragraphs, and al- histogram (Fig. 8) of the distribution by though the situation is evidently complex, size of all 434 lowland species shows not I think the small size of New Guinean only that most are small but also that Carabidae may be a result of a combi- separate modes occur at 2.0-2.95 and at nation of three factors: (1) difficulty of 6.0-6.95 mm, with a deficiency especially access, (2) the greater dispersibility of at 4.0-4.95 mm. A highly speculative and >mall as compared with large Carabidae, oversimplified explanation can be derived md (3) time. New Guinea has been from the possible relation of Carabidae somewhat isolated, on the west by water with ants. Very small Carabidae may be aarriers which have probably been nar- able to hide from ants, and relatively large -ower than now at times in the past but ones may be able to protect themselves, A^hich have existed for a very long time, while Carabidae in the 4.0-4.95 mm size md from Australia by fairly effective class may be especially vulnerable to com- jcologic barriers even when what is now petition with or predation by ants. If so, he island was connected with the main- this is only one aspect of a probably com- and. Small insects do disperse through plex impact of ants on Carabidae and he air more readily than large ones; the carabid faunas which is further discussed mcestors of New Guinean Carabidae were under Ants [27], lot only all small (or at least not very A simpler explanation of the bimodal arge ) but almost all of them were or may size distribution of lowland Carabidae in lave been winged when they reached the New Guinea can be derived from the sland [23, 88]. The small, winged an- predominance of the single genus Tachijs •estors of New Guinean Carabidae may ( taking the genus in a broad, old-fashioned lave crossed barriers which large Carab- sense). This is a huge genus of small dae and flightless ones did not cross. Carabidae. Although it is well represented rhis explanation does perhaps account well in some temperate areas, it is primarily •nough for the initial small size of New tropical, and includes great numbers of Guinean Carabidae. To account for the species in the tropics of all continents, iresent smaller size of Carabidae on New Most of the really small Carabidae in Guinea as compared with Madagascar, New Guinea belong to this genus: the ime must be invoked, I think. The New portion of the bimodal histogram (Fig. Guinean fauna is evidently relatively re- 8) represented by Tachys is shown by ■ent in its origins; the Madagascan fauna, hatching in the first three columns of the )robably much older. Also, Madagascar histogram. Without Tachys, the lowland s much more effectively isolated than Carabidae of New Guinea have a regular ^ew Guinea; invasions and replacements size distribution, with a single mode at )robably occur less often; and old stocks or near 6-6.95 mm. The bimodality of the )robably survive longer and have more fauna is therefore apparently due to the ime to increase in size. (See [89] for addition of a large number of species of liscussion of relative age and rate of the predominant genus Tachys to what uniover of the New Guinean fauna.) So, would otherwise be a simpler, unimodal vhile the small size of New Guinean Carab- size distribution. But this explanation does dae is not due to decrease of size, the not necessarily preclude the preceding irge size of many Madagascan forms may one. The small size of individual Tachys 164 BuUetin Museum of Comparative Zoology, Vol 142, No. 2 CO UJ o UJ a UJ OQ ID 0- I- 2- 3- 4- 5- 6- 7- 8- 9-10- 1 1- 12-13-14-15-16- 17- 18-19-2021- SIZE CLASSES (EACH N-N.95 MM) cm. 24-25- 32- 51 Figure 8. Histogram of size distribution of lowland Carabidae in New Guinea. Each species is counted once, according tci its median lengtfi. Eacfi column represents number of species of which the mean length falls between N-N.95 mm; the actual numbers of species are given above the columns. The hatched portions of the first three columns represent Tachys. See text for further explanation. may be one reason for their great success, and may enable them to Hve in the pres- ence of dominant ants more successfully than slightly larger Carabidae can do. Although several groups of Tachijs have certainly radiated to some extent in New Guinea, the number of species there is not due primarily to radiation but to the large number of separate stocks that have reached the island: my data sheets indi- cate at least 23 separate relationships be- tween different New Guincan Tachys and those of other areas, and this suggests at least 23 separate invasions of New Guinea by members of this one genus. In an attempt to get a more detailed explanation of the two size modes of low- land New Guinean Carabidae, I have made a size histogram (Fig. 9), of the 217 species that I found at Dobodura, indicat- ing the ecologic composition of the size classes. I have limited this diagram to species that I myself collected at one locality, because I know something ol their ecology. (See Ecology: habitats [24 _ for further discussion of habitats of Ne^^' Guinean Carabidae.) This histogram sug gests that lowland mesophile Carabidae do have a bimodal size distribution in New Guinea. Mesophiles compete more di- rectly with ants than hydrophiles anc arboreal Carabidae do, and would be more likely to show the effects of competition. My tentative conclusion is that th(" double-moded size distribution of lowlanc Carabidae in New Guinea is due to the presence of excessive numbers of smal species of Tachys, but that this explanatioi still allows the possibility of an underlying effect of competition with or predation b) ants. Mountain-living Carabidae in Nev, Guinea have a different size distributioi from the lowland forms. Figure 10 is ; histogram of the sizes of the 215 known strictly mountain-living species. This figure The carabid beetles of New Guinea • Darlington 165 CO lij o UJ a. CO UJ CD z 0-1- 2-3-4-5-6-7- 8-9-1011-12-13-14-15-16-17-18- SIZE CLASSES (EACH N N 95 MM) igure 9. Histogram of size distribution of Carabidae found at Dobodura. Explanotion as for Fig. 8, except hatched portions f columns represent ecologic groups: right-oblique hatching at bases of columns, hydrophiles; no hatching, mesophiles; nd left-oblique hatching at top of columns, arboreal forms. hows a single mode at a larger size — at ir near 9-9.95 mm — than the second mode •f the lowland species. Very few Tachys iccur in the mountains; their place in the listogram is represented by the hatched >art of the first column; and removal if Tachys from this histogram does not •hange it significantly. However, ants too ire relatively few at higher altitudes in 'Jew Guinea, and their fewness may have omething to do with the size distribution )f mountain-living Carabidae. In summary of the size of New Guinean Carabidae: all the characteristic ones are mall. The absence of large forms may )e due to a combination of ( 1 ) difficulty )f access, (2) the greater dispersibility of mall as compared with large Carabidae, md (3) the fact that the existing New Guinean carabid fauna is relatively recent n its origins and has not had time to ^volve large forms. Tlie lowland Carabidae of New Guinea have a bimodal size dis- tribution, with modes at 2-2.95 and 6-6.95 mm; this bimodality is due primarily to the presence of many small species of Tachys, but may also reflect an underlying relation with ants, which may compete with or prey on Carabidae especially in the 3 to 5 mm size classes. Mountain-living New Guinean Carabidae have a unimodal size distribution, with the mode at or near 9-9.05 mm; l)oth Tachys and ants are relatively few in the mountains, and their absence may partly account for the dif- ferent size distribution of mountain-living as compared with lowland Carabidae. [21] ^^^//ii,'.s and wing atrophy. I have a long-standing interest in the wings, wing atrophy, and flight of Carabidae (Darling- ton, 1936; 1943), and I have been con- stantly on the lookout for cases of wing reduction among the New Guinean species. An advantage of specimens killed in alco- 166 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 if) liJ O liJ Q. 0) U. o Ul OQ D Z CM ro lO in CM (M ro — ^ o o IV CD cvl 1 r \ 1 ^ ^ CJ OJ CVJ 1 1 0- 1- 2- 3- 4- 5- 6- 7- 8- 9- 10- II- 12-13- 14-15- 16-17-18- 19-2021-22- SIZE CLASSES (EACH N-N.95 MM) Figure 10. Histogram of size distribution of mountain-tiving Carobidoe in New Guinea. Explanation as for Fig. 8. The hatched portion of the first column represents lachys. hol, as most of mine were, is that the elytra are likely to be slightly separated and the inner wings easily visible. This is not usu- ally the case in dry-killed specimens, but if the latter are in good condition and properly mounted, it is usually possible to separate the elytra with the point of a pin and see whether or not the wings are fully developed. I have thus been able to see the wings of thousands of specimens of New Guinean Carabidae with com- paratively little difficulty, and in species in which wing reduction has been detected or suspected, I have supplemented this routine examination with more careful examination, by relaxing specimens and raising their elytra. However, I have not usually extended my examination of wings to specimens collected at light, because as a rule only winged, flying individuals can come to hght, so that light trap material is very deficient in flightless forms. In discussing state of wings, I use three symbols: +w means with wings fully de- veloped or at least long and folded at apex; ±w, wings dimorphic, full in some and reduced in other individuals; and -w, wings reduced in all individuals. At low altitudes in New Guinea most Carabidae arc +w. Many are known to fly (to Hght), and most probably do so. Of the 434 known lowland species (exclusive of Cicindelinae) only 17 (Table 3), or about 4 per cent, have been found to have any sort of wing reduction at low altitudes in New Guinea.^ (Two additional species ^ Although I have excluded tiger l)eetles (Cicindelinae) from my analysis and discussion of the New Guinean carabid fauna, one species should be mentioned in the present connection. It is Tricondyla apteia (Part I, p. 334). This is tlie only flightless tiger beetle in New Guinea, so' far as I know. Not only is it flightless itself, but it belongs to a wholly flightless Oriental genus, and has almost surely reached New Guinea with- out flying. It occurs in rain forest, but not on the forest floor. It li\'es on the trunks of trees, The carabid beetles of New Guinea • Darlington 167 Table 3. Lowland Carabidae of New Guinea in which reduction of wings has been found ( Scaritini ) Clivina dedlata (Part I, p. 372), 4-w and -w popu- lations at different localities Clivina enigatella (Part I, p. 380), +w and ±w populations at different localities ( Bembidiini ) Tachijs serrula (Part I, p. 408), +w and -w popu- lations at different localities Tachijs amhulatus (Part I, p. 425), -w at single known locality Tacht/s aviits (Part I, p. 426), ±w at Dobodura {Tachijs truncatus (Part I, p. 431; Darlington 1970: 15), apparently always +w in New Guinea but ±w in Micronesia) {Tachijs brachtjs (Part I, p. 433; present part, Tax. suppl.), -fw in New Guinea (only 4 specimens) but -w or ±w in the Moluccas and Formosa ( Taiwan ) ( Pterostichini ) Lesticus politus (Part I, p. 526), 2 specimens only seen, 1 +w and 1 -w, from different localities Platijcoehis depressiis (Part I, p. 534), ±w, geo- graphic distribution of wing forms not de- termined Loxandriis latiis (Part I, p. 551), ±w at Doborura ( Agonini ) ( No lowland species with reduced wings, although many -w in mountains ) ( Oodini ) Oodes terrestris {laevlssimiis of Part III, p. 34), apparently ±w at Dobodura, but wings only slightly reduced in "— w" individuals Oodes rossi (Part III, p. 34), single known speci- men -w ( Harpalini ) Hijphaereon timidus (Part III, p. 67), ±w (wings actually polymorphic) at Dobodura; state of wings elsewhere not determined ( Lebiini ) Nototariis papua (Part III, p. 186), -w in series from Dobodura ( Pentagonicini) Parascopodes cijaneus (Part III, p. 196), ±w at Dobodura ( Zuphiini ( Leleupidiini ) ) Colasidia papua (present part. Tax. suppl.), single known individual -w ( Brachinini ) Pheropsophus apfinomorphus (Part III, p. 237), -w Pheropsophus catulus (Part III, p. 238), -w Pheropsophus canis (Part III, p. 238), -w Total: 17 species, of which 10 ±w, 7 -w (so far as known); product of at least 14 separate wing- reductions ; little or no radiation in -w stocks that are +w at low altitudes are ±w on the Bismarck Range — see Table 5.) The patterns of occurrence of +vv and -w individuals of these 17 species are sum- marized in Table 4. These 17 species rep- resent at least 14 separate stocks, in each of which wing reduction has occurred independently. None of these stocks has radiated much at low altitudes, although speciation may have begun in the -w Tachys {amhulatus and avius), Oodes, and Pheropsophus {catulus and canis) (see again Table 3). It is noteworthy that no reduction of wings seems to have occur- red among Agonini at low altitudes in New Guinea, although the wings have atrophied in a number of stocks of this tribe on the mountains, and although the -w agonines dominate the mountain carabid fauna of the island. In being composed almost wholly of (small) winged species, the low- land carabid fauna of New Guinea is like the faunas of some lowland areas in the continental tropics and of some other tropical islands (see fifth following para- graph ) . and it is active and conspicuous on them by day. It is about 25 mm ( 1 inch ) long, slender, cylin- drical, with large eyes and long appendages. It is strikingly antlike in appearance and movements. The exceptional success of this insect may be due to its ability to compete with large preda- ceous ants. It combines antlike agility and strik- ing power ( it can bite severely ) with protective annor and perhaps better vision than ants have. It may be a biological example of the well-known principle, "if you can't beat them, join them." But it is exceptional, and its existence in the lowland rain forest of New Guinea emphasizes how few other oarabids are flightless tliere. 168 BuUctin Museum of Comparative Zoology, Vol. 142, No. 2 Table 4. Summary of distribution of +W and -W indi\tduals in 17 lowland Carabidae listed IN Table 3 ±\\' populations at single localities ( in single populations) 6 species +w and -\v populations at different localities — 3 species ±\v, hut distribution of wing forms not determined 1 species -w in all specimens seen 7 species Total 17 species Wing reduction among Carabidae is often followed by other structural changes, especially by narrowing of humeri and shortening of metepisterna. However, most of the lowland species (listed in Table 3) in which wing reduction has oc- curred either are still dimorphic (±w) or do not yet show the structural changes that tend to follow wing reduction. Of the 17 species listed in Table 3, only five begin to show the secondary structural changes (indicated above) that suggest that the insects have been flightless for a consider- able time. Tliese five are Oode.^ rossi, Nototarus papua, Cola.sidia paptia, Pherop- ■sophu.s catiihi.s, and P. cants. I have col- lected three of these species and close relatives of the other two. All five probably live in leaf litter on the floor of rain forest. This fact suggests that selective factors are relatively favorable to flightlessness on the ground in rain forest, although the small number of species that have become flight- les.s even there at low altitudes suggests that the selective advantage is limited. Special factors in favor of flightlessness on the floor of rain forest perhaps include the stability and continuity (both in space and time) of that habitat. Factors limiting flightlessness there may include patchiness of distributions [22] and liability to flood- ing. Floods often do occur on flat ground in rain forest, and Carabidae do fly to escape from them. Competition wdth ants, which are numerous on the ground in rain forest as well as in many other habitats in the lowland tropics (but which are fewer beside water and at high altitudes), may be an additional factor limiting flightless- ness. Known mountain-living species of Carab- idae in New Guinea (found above 1000 m ) total 376 species. Of these, 161 species occur also in the lowlands (below 500 m), and all of these species with wide alti- tudinal distributions are +w, except that two normally +w species of T richotichnus are locally ±w on the Bismarck Range (see Table 5). There is here a correlation, perhaps to be expected, between wide altitudinal distribution and possession of fimctional wings. Of the 215 species of New Guinean Carabidae found above 1000 m but not below 500 m, 69, or 32 per cent, exhibit wing-reduction (Table 5). Only one of these species is known to be ±w (Ga.stra- goniim teircstre. Part II, p. 226), the other 68 species being unifomily -w so far as known. In fact Gastragonum is the only strictly mountain-living genus of Carabidae in New Guinea in which the wangs are dimorphic, most of the other strictly alti- coline genera being apparently products of^ radiation of -w ancestors. (Exceptions are' Plicagoniim, riparian Potamagonum, and' probably-grass-living Maculagomim, which are uniformly +w.) The phylogenies of the mountain-living forms have not and probably can not be fully worked out, but at least 20 different stocks (probably more) ha\'e undergone wing reduction indepen- dently to produce the -w mountain forms, and secondary structural modifications and' radiation in some -w stocks indicate that wing atrophy is long-standing in many of them. At highest altitudes, above 3000 m, only 21 species of Carabidae have yet been found in New Guinea (Table 2). Of' these, 20, or 95 per cent, are -w, the only +w species on the list being Maculagomim altipox. The altitudinal distribution of +w and -w Carabidae on New Guinea is consistent The carabid beetles of New Guinea • Darlington 169 Table 5. Mountain-living Car.a.bidae of New Guinea in which reduction of wings has been FOUND Scaritini ) 'Hivina toxopci (Part I, p. 363), -w (Australian relatives both +w and — w ) Uivina kiibor ( present part, Tax. suppl. ) , -\v, (relatives +w) Bembidiini ) Jwnastis inops (Part I, p. 486), -w (genus in- cludes +w species ) Panagaeini ) Iraspedophonis gressittorum (present part, Tax. suppl. ) , -w ( Australian species -w, some Oriental -fw) Pterostichini ) iecijclothorax toxopei (Part I, p. 506), -w (Aus- tralian relatives both +w and -w) Iecijclothorax sedlaceki (present part, Tax. suppl.), -w (see preceding species) ^estkus (Part I, pp. 521ff; present part, Tax. suppl.), 5 species -w (genus includes +\v and ±\v species ) Ihytifcronia (Part I, pp. 533ff), entire genus (2 species) -w, (related Australian genus +w) 'rosopoginus (present part. Tax. suppl.), 2 species -w ( genus includes +w species ) inaloma (Part I, pp. 538ff; present part, Tax. suppl. ) , entire genus ( 4 species ) -w ( rela- tives undetermined) Agon in i ) ^otagonuru ambulator (present part. Tax. sujipl.) -w (genus otherwise entirely -|-\v) l,astragonum (Part II, pp. 222ff), 1 species ±\v, 3 -w ( genus includes -fw species ) diagonum (Part II, pp. 229ff; present part. Tax. suppl.), entire genus (6 species) -\v dontagonum (Part II, pp. 233ff; present part, Tax. suppl.), entire genus (8 species) -w Nebriagonum (Part II, pp. 235ff; present part, Tax. suppl.), entire genus (7 species) -w Laevagonum (Part II, pp. 243ff; present part, Tax. suppl. ) , entire genus ( 7 species ) -w Fortagonum (Part II, pp. 247ff; present part. Tax. suppl.), entire genus (11 species) -w ( Licinini) Microferonia baro (Part III, p. 19), -w (genus includes +\v and ±w species in Australia) ( Oodini ) Oodes wilsoni (Part III, p. 35), -w (most species of genus -\-w [A -w Coptocarpus just received from New Guinea has not \et been described] ( HariDalini ) Chydaeus (Part III, pp. 47ff; present part. Tax. suppl.), 2 species -w (genus includes -fw and ±w Oriental species) Tricliotichnus nigricatis (Part III, p. 52), ±w on Bismarck Range ( 4-w elsewhere ) Trichotichnus alius (Part III, p. 54), ±w on Bis- marck Range ( +vv elsewhere ) ( Pengatonicini ) Scopodes alius (Part III, p. 198), -w (Australian relatives include +\v and ±w species ) ( Zuphiini ( Leleupidiini ) ) Col-asida madang (present part. Tax. suppl.), -w ( known relativ es all -w, but group descended from +w stock ) Total: 71 species, of which 3 ±\\', 68 -w; product of at least 22 separate wing reductions; radiation in -w stocks in some Pterostichini and especially Agonini \ith distributions in some other, compa- able parts of the world. Some other (but lot all) tropical lowland carabid faunas ire wholly or almost wholly +w, for ex- imple in the Santa Marta region of Co- ombia, northern South America, and on 3arro Colorado Island in the Panama "anal Zone (DarHngton, 1943: 41). And n the West Indies the situation is extra- )rdinarily like that on New Guinea, ilthough the Carabidae concerned are en- irely separate in their immediate deri- vations and local evolutions. On Cuba, for example (DarHngton, 1943: 49), of 144 known lowland species, almost all are +w, and of the five (3/r ) lowland species that do show wing reduction, four are still ±w; but -w stocks appear with in- creasing altitude on the mountains, and on the summit of Pico Turquino (the highest Cuban mountain, c. 6500 ft. or 2000 m), all se\'en known species are exclusi\ely -w. I collected on Turquino myself for two weeks in 1936 and secured 459 specimens 170 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 of these seven species, every specimen -w. On Jamaica, all known lowland Carabidae (63 species) are +w, but again transition to -w fornis occurs with increasing alti- tude, and above 5000 ft. (c. 1500 m) the ten known species are 80 per cent -w, onK' 20 per cent +w (Darlington, 1943: 49)'. Wing atrophy of Carabidae in New Guinea is correlated not only with altitude but also with habitat. Carabids may be roughly classified as mesophiles (ordinary ground living species, called also geo- philes), hydrophiles (species specifically associated with open water, especially swamps and the banks of streams), and arboreal species. (See Section [24] for further discussion of this classification of habitats.) Wing atrophy has occurred fre- quently among mesophiles but rarely in the other ecologic groups in New Guinea. A few -w species are or may be hydro- philes, including 'Nehria'^omim cepholum and N. percephalum (Part II, pp. 239, 240), which live beside moimtain torrents on the Bismarck Range. But, so far as I know, not one of the many arboreal Carabidae of New Guinea is -w at any altitude. : The situation among New Guinean Carabidae suggests that mutations from a long- to a short-winged condition are con- tinually occurring at all altitudes. In the lowlands, selection apparently usually elim- inates -w individuals, and most species remain +w, although wing atrophy does sometimes occur among mesophiles living on the floor of the rain forest. On moun- tains, however, -w mutants more often survive, species often become -w, and the -w stocks persist and sometimes radiate. Tliis process probably occurs increasingly often with increasing altitude; it occurs most often among mesophiles, sometimes among hydrophiles, rarely if at all among arboreal Carabidae in New Guinea. In extreme cases this process has produced groups of interrelated -w species, which I have considered genera, confined to small areas in the mountains of New Guinea. Examples are Nehriagomtm and Laevag,o- niim at high altitudes on the Bismarck Range, etc. (Part II, pp. 235, 246; present part. Tax. siippL). [22] Ex))lantions of wing atrophy. Wings and wing atrophy of Carabidae in various environments and on mountains and islands have been discussed by me in 1936 and 1943. I want now to summarize and apply my conclusions to the situation in New Guinea. Their relationships indicate that most -w Carabidae on mountains in New Guinea are derived from ancestors that were +w when they reached the island [23, 88]. Tlie -w mountain faunas are mainly products of conversion rather than of concentration (see my 1943 paper, pp. 52-53); that is, they have been produced by atrophy processes that have occurred locally, not by accumulation of -w stocks from other regions. Certain factors should be noted that do not induce wing atrophy among mountain Carabidae or that are unimportant. Alti- tude itself (thinness of air) apparently does not induce wing atrophy or flightless- ness (see my 1943 paper, pp. 50-51). Cold apparently does not directly inhibit de- velopment of wings (ibid., p. 51). And exposure to wind — selection of -w form? by blowing away of -f w flying individuals (Darwin's factor) — seems at best relatively unimportant (ibid., p. 51), for many -w mountain-living Carabidae, including mosi of the ones on mountains in New Guinea live on the ground in dense montane foresi and are not exposed to winds. Some other factors that probably do in- crease incidence of wing atrophy anc flightlessness on mountains seem to be oi| only minor importance. One is freedoirj from flooding. In the lowlands of New Guinea, flat ground is often flooded by^ standing water, and winged Carabidat often do fly to escape the floods, whik flightless individuals are obviously morome, the result of cutting and burning of -ain forest by native farmers. What seemed :o me to be similar grassland, at least offer- ng a similar habitat to Carabidae, occurs jp to at least 2000 m in inhabited valleys Dn the Bismarck Range and elsewhere, .vhere man has cleared the mid-altitude nountain rain forest. The second, surely latural type of grassland, with the grass much lower than "kunai" and forming tus- socks, occurs above the forest line on high nountains, including the Bismarck Range. "Wetlands" can be used to include a variety of habitats where running or stand- ing water or simply wet ground support special vegetations and special Carabidae. Swamps are widespread at low altitudes in New Guinea, but are relatively few and small in the mountains where drainage is more rapid. The lowland swamps include large areas of sago palms, especially on the deltas of the big rivers. The enomious, spine-bearing leaves of these palms fall into the water and make working there difficult and dangerous. This habitat has therefore not been well collected for Carabidae, although it is evidently rich in subaquatic species. Other types of swamp with more diverse vegetations occur here and there, especially inland. And salt swamps, man- grove zones, and other special habitats occur along the sea coast and probably support special Carabidae, although very few of them have been collected. The edges of running water have also special Table 6. Ecologic composition of lowland New Guinean Carabid fauna No. in Ecologic No. of ecologic groujis species subgroups Fossorial 27 ( Scaritini ) Mesophile 133 Rain forest majority Open places minority Hydrophile 136 Wet lands 105 By streams 31 Arboreal 129 In foliage 84 ( incl. 29 Demetrida ) Trunks/logs 45 Unclassified 9 habitats ranging from fringes of dense forest to grass, reeds, and virtually sterile banks and bars, which may be stony, sandy, or muddy. Different habitats on the banks of large rivers, the edges and debris-blocks of smaller streams nmning slowly in heavy shade in rain forest, and the stony and sandy edges of rapidly run- ning brooks all have different carabid faunas. Many of these habitats are found only or chiefly in the lowlands, but rapid brooks occur at all altitudes up almost to the peaks of the highest mountains. Be- sides these larger and more obvious sub- divisions of wetlands. New Guinea presents an almost endless variety of wet spots sometimes only a few square meters in area: rain pools, overflow of streams, ac- cumulations of water in holes left by the roots of fallen trees, etc., and some com- mon carabids inhabit these places. [25] Ecologic composition. The ecologic composition of the louland carabid fauna of New Guinea (Table 6) is shown by figures compiled from my data sheets [16], from the column headed "Ecology." Be- cause my information about habitats is incomplete, I have assigned some species according to the habitats of their nearest relatives; these assignments are probably correct in most cases, but perhaps wrong 174 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 in a few.^ And in nine cases I have not wanted even to guess about the habitats of isolated species. The totals are there- fore only approximations, but they are close approximations, based on my field observations. Even when habitats are known, some species are still difficult to assign to eco- logic groups. The fossorial species might be counted as hydrophiles or (a few) as mesophiles. The line between mesophiles and hydrophiles is not sharply drawn in any case; it is crossed by various species with wide ecologic ranges. The distinction between mesophiles, rotten-log forms, and tree-trunk-arboreal forms is not sharp either. Nor is the line between mesophiles and fohage-arboreal forms; it is crossed by species like Vioh^onum viohceum, which occurs in piles of dead leaves on the ground as well as in foliage. Never- theless, these doubts and difficulties do not seriously affect the approximations given in Table 6. In Table 6, the fossorial Scaritini are separated first, although most of them are included with the hydrophiles in statistical analyses elsewhere in the present paper. Without the fossorial fonns, the lowland Carabidae of New Guinea divide almost equally into three main ecologic groups: mesophiles, hydrophiles, and arboreal forms. This is probably a fair sample of the ecologic composition of lowland cara- bid faunas in the wet tropics elsewhere, ^ Cases in which more or less closely related carabid species occur in different haliitats include the following in New Guinea. Although most Tachijs occur on the ground in wet places, T. aeneus (Part I, p. 46.3) occurs on dnj ground, and r. wallacei (Part I, p. 479) is arl)()real. Most Oodes are more or less aquatic, but O. tene.s- tris lives in leaf litter on the ground in rain forest. And, although most E^adroma li\'e in wet places, E. robusta (Part III, p. 71) li\'es on dry ground. For some examples of carabids which have made ecologic shifts within tlie limits of New Guinea, see [91]. All these cases of ecologic divergence have been allowed for in compiling Table 6, but some other, similar cases may still be undetected. although hydrophiles may be relatively more numerous in some places ( Darlington, 1943: 41). In the case of the New Guinean carabid fauna, all three main categories can be usefully subdivided. The mesophiles can be divided into rain- forest forms (the majority) and those that inhabit open places (a minority). I have not tried to give exact figures for these subgroups, because my information about the occurrence of some of the species is insufficient. The hydrophiles divide into those that occur on wetlands in general (105 species) and those that occur only by running water ( 31 species ) . The line between these two subgroups is not sharp. However, the division is important because the stream- side species are the only ones that are likely to range far up the mountain slopes. The distribution of hydrophiles depends on distribution of surface water and of special water-side habitats, and some of these habitats are independent of the type of forest cover. Some hydrophiles therefore occur in both rain-forested and opener country, and their dispersal is compar- atively little affected by discontinuities in the distribution of rain forest. Also, most of them are winged and many fly actively. They are therefore likely to be good dis- persers in areas (like the Malay Archi- pelago) where forests and opener country alternate to some extent (see [88, 91]). Finally, the arboreal forms divide into those that live in foliage (84 species, in- cluding 29 lowland Dernetrida) and those that live on tree trunks and recently fallen logs ( 45 species ) . Almost all these arboreal carabids occur in rain forest, of course. With increasing altitude, the ecologic composition of the New Guinean carabid fauna changes strikingly. At higher alti- tudes, most Carabidae are mesophiles living in (montane types of) rain forest,{ Most hydrophiles have disappeared, ex- cept a few that are strictly associated with running water. And arboreal Carabidae The carabid beetles of New Guinea • Darlington 175 are reduced in numbers too, but include relatively more species of Demetrida. I know too little about the habitats of most ?pecies at higher altitudes to give exact Figures. [26] Altitude. Altitude affects the dis- tribution of Carabidae in New Guinea in several ways. The change of genera and species from the lowlands to the highest altitudes is so great that, although ex- tensive overlapping occurs at intermediate altitudes, no species are common to the lowland and highest-mountain faunas and virtually all the genera are different. (See Table 2 for composition of the highest faunas. ) The only genus that is represented at all at highest and lowest altitudes is Scopodes, but this genus is not (in New Guinea) a regular component of the low- land fauna, and the one or two mid-alti- tude species of the genus that do descend to the lowlands at least locally are not closely related to the single very-high- altitude species. The effect of altitude is shown also by the striking reduction of numbers of species from the lowlands to the mountain tops (Fig. 7); by the dif- ferent size distributions of lowland and mountain species ( Figs. 8, 10 ) ; by the different ecologic compositions of lowland and mountain faunas [25]; and by the increased proportion of species with atro- phied wings, from about 4 per cent in the lowland fauna to about 95 per cent at highest altitudes [21]. See Szent-Ivany (1965) for further notes on the vertical distribution of some beetles in New Guinea. See Greenslade ( 1967 ) for an indication of the correlation of habitats and of insect distributions with altitude in the Solomon Islands. And see Mani ( 1968 ) for a more general discussion of high-altitude insects. Note, however, that even at highest altitudes New Guinean Carabidae do not show all the modifi- cations of alticoline forms that are found in some other parts of the world; for example, I have found no general reduction of size of individuals even at highest alti- tudes (cf. Mani, 1968: 58ff). The effect of altitude on New Guinean Carabidae is apparently exerted partly in- directly, through the effect of altitude on habitats. Some species that are common in the lowlands occur also up to at least about 2000 m where forest has been cleared and where for this or other reasons habitats, including grassland, approximate lowland habitats. For example, I found the following common lowland species of Carabidae in cleared country in the Waghi and Chimbu valleys at between about 1500 and 2100 m altitude: Tachijs fasciatus and T. aeneiis (Part I, pp. 414, 463), Notop,o- niim angusteUum, vile, and margaritum (Part II, 133, 135, 145), Chlacnius flavi- giittatus (Part III, p. 26), Egadroma quinqucpustidata and rohusta (Part III, pp. 70, 71). It seems likely that these and other winged lowland species have invaded the Waghi and Chimbu valleys from below only after the mountain valleys had been cleared. The climatic differences have not prevented the invasions, although the ecologic ranges of some of the species did apparently change with increasing altitude: some species that apparently never hide under stones at low altitudes do so in the Waghi-Chimbu area [3]. Many of the lowland species that have been found at Wau [5] at 1200 m altitude or higher have probably invaded cleared land there. In undisturbed forest and other natural habitats the changes of carabid faunas with altitude are apparently more clear-cut. My infonnation about occurrence of Carab- idae in forest at mid-altitudes is scanty. However, I do know that all species of Carabidae found in montane forest on the Bismarck Range between about 2100 and 3000 m and all species found in open tussock-grass country above about 3000 m are different from lowland species, and many of the genera are different. It is not yet possible to assess the relative importance of the direct effects of high- 176 Bulletin Museum of Comparative Zoologij, Vol. 142, No. 2 ANTS ANTS C A R A B D A GEO PHILES iHYDROPHILES 'ARBOREAL -w :=; 1 1 + W 4-W +W COOL TEMPERATE E. N. A. Ti- ll II ii \ ( \ \ V I t \ \ \ s. +w i; TROPICAL + W -I- W LOWLANDS OF NEW GUINEA I C A R A B DAE Figure 11. Diagram of (hypothetical) relation of Carabidae to ants in cool temperate eastern North America and in the tropical lowlands of New Guinea. Redrawn and modified from Darlington, 1943, p. 42, fig. 4. Diagram is intended to sug- gest that ants compete most severely with flightless (-w) ground-living (geophile) Carabidae; that the effects of competition are greater in the tropics than in the temperate zone; and that dominant ants replace most flightless ground-living Carab- idae in tropical lowland New Guinea. altitude climate (cold, continual dampness in some habitats, perhaps thinness of air) on carabid faunas. I can only say that, judging from the restriction of high- mountain species to special montane habi- tats, the indirect effects are probably more important than the direct ones. The in- direct effects are probably exerted not only by control of vegetation but also by limi- tation of habitats on mountains (absence of large swamps, large rivers, etc.), by hmitation of area on mountains (which probably affects nature of populations and, indirectly, state of wings [22]), by reduc- tion of competition with ants [27], and perhaps in other ways. [27] Ants. Ants are dominant insects, especially in the tropics, and their impact on other insects must often be tremendous. I have suggested that they may affect the size distribution of Carabidae in New Guinea [20] and that their fewness or absence at high altitudes may modify carabid faunas there [26]. Ants may also modify the ecologic composition of the lowland carabid fauna of New Guinea. This fauna, as compared with the faunas of temperate regions, is deficient in meso- philes and especially deficient in -w mesophiles, which probably compete most directly with ants. The ecologic compo- sitions of the carabid faunas of cool temper- ate eastern North America and of the tropical lowlands of New Guinea are diagrammed and compared in Figure 11. The diagram is intended to suggest that the tropica] fauna, which is larger than the temperate one in any case, would be still larger if it were not for the presence of ants, and that the latter take the place mainly of mesophiles and especially ol flightless mesophiles. [28] Ecologic interactions. The differeni ecologic factors discussed above do noi. act independently but must interact ir very complex ways on carabid faunas Major habitats and vegetations profoundh affect occvnrence of Carabidae but an The carabid beetles of New Guinea • Darlington 111 themselves dependent on climate. Altitude may act partly in relatively direct ways (by reducing temperature, increasing pre- cipitation, modifying insolation, and per- haps in other ways) and partly indirectly (by modifying habitats and vegetations). And the effect of ants varies with habitat and altitude. So, climate, habitats, altitude, and ants all interact complexly to modify the environments in which Carabidae live and to determine, directly or indirectly, how many Carabidae and what kinds of Carabidae live in different places. This statement does not do justice to the actual complexity of the environment in New Guinea. The ecologic ranges of different Carabidae on the island are surely deter- mined by the interactions of very many factors derived from the inorganic environ- ment, vegetations, and animal prey, preda- tors, parasites, and competitors. Actual details are almost w^iolly unknown and their investigation must (as I have indi- cated at the beginning of [24] ) now be left to the ccologists. [29] Ecologic ranges. I have used the phrase "ecologic range" deliberately, rather than "niche." "Niche" is used by many ecologists, and I have used the word my- self, but I think it is inaccurate and mis- leading. It suggests that the environment is full of pigeonholes with fixed bound- aries, and that different kinds of animals and plants are in fact neatly pigeonholed in nature. Experienced ecologists know that this is not so (at least I hope tHey know it!), but the concept of niche some- times confuses them just the same. For example, ecologists sometimes speak of an island as having a certain number of niches for animals, with some of the niches unfilled, as if the niches existed before the animals were there. But if niches exist for animals at all, it is the animals themselves that make them; dif- ferent kinds of animals surely in part de- termine the ecologic limits of other kinds. And the limits apparently change accord- ing to the number of animals present. When only a few^ species are present on an island, each is likely to occupy a wider segment of the environment than each will occupy when more species are present. To say that, in the first case, each species occupies several niches does not avoid the difficulty; there is no ultimate niche, no division of the environment so fine that it cannot be further divided. One might almost say that, instead of an island possess- ing a certain number of niches some of which may be unfilled, an island has no niches at all until animals come and define them. What the island does have is eco- logic dimensions or ecologic amplitude or ecologic totality, which is not divided be- forehand but which animals do divide into a few or many fractions in the course of time. It seems to me that ecologic ranges are comparable to geographic ranges. A given piece of land — say a continent — has geo- graphic dimensions, and is potentially able to support a varying number of species of plants and animals, each, of which has a geographic range. The ranges are areas occupied by populations. They do not exist until the populations occupy them, and they often change and often overlap. At- tempts have been made to divide continents into which might be called idealized (and therefore typological! ) geographic ranges corresponding to climatic zones or major biomes, but (as a zoogeographer) I know that actual species' ranges often do not fit the idealized patterns very well. Similarly, to suppose that the enviionment can be divided into niches which exist before animals occupy them is a (typological) idealization which is likely to obscure the real facts. It is better to think of each species as having an ecologic range which is a fraction or fractions of the total en- vironment. It is then easy to think of ecologic ranges as changing from time to time, or as being discontinuous, or as over- lapping each other, as they often do. The concepts of "changing niches" and "over- lapping niches" are confusing! 178 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 ZOOGEOGRAPHY: EXISTING GEOGRAPHIC PATTERNS [30] Geographic patterns, relatiomhips, and origins. Because I am a zoogeographer, especially interested in the patterns and significance of animal distribution, I shall make a geographic analysis of the New Guinean carabid fauna in much more detail than would ordinarily be attempted in a faunistic paper. In doing this, I shall begin by making a clear distinction be- tween description of existing geographic patterns and discussion of past origins. This is just the beginning of the sub- dividing of the subject that is necessary to describe and understand the zoogeog- raphy of any complex fauna. Few persons, in fact few zoogeographers, fully under- stand how complex the subject it, or how necessary it is to recognize the complexities if zoogeographic data and analyses are to be informative and significant. I doubt if faunal relationships and other zoogeo- graphic problems can yet be handled satis- factorily by simple numerical methods. These methods are too likely to give a superficial picture that hides more than it reveals. The gross taxonomic compositions of major carabid faunas from the Orient (Java) to New Guinea and tropical Aus- tralia are described and diagrammed in [64] and Figure 13. Now to be considered are the finer details of distributions and relationships that underlie the gross pat- tern. [31] Existing geographic relationships: problems and procedures. I shall begin by describing existing geographic patterns and relationships without intending to imply anything about past geographic origins (except sometimes parenthetically). The first question in this connection is, at what level are geographic patterns most signifi- cant: at the level of tribes, genera, or species, or of geographically separate stocks? The choice of levels can profoundly affect the results of analysis, and the choices are often complex. For example, Coptodera (Part III, pp. llOff), as I have treated it, is one genus; splitters would divide the New Guinean species among about five smaller genera; eight species of the genus occur in New Guinea; and these eight species represent seven geographi- cally separate stocks, each with its own relationships outside New Guinea. In mak- ing faunal comparisons, should this genus be counted as one or five or eight or seven units? This is the kind of situation that numerical zoogeography too easily hides. For another example, the genus Deme- trida (Part III, p. 140) is represented on New Guinea by 59 known species, all apparently interrelated among themselves (apparently produced on New Guinea by radiation of perhaps only one or certainly not more than a very few ancestors). Out- side New Guinea, this genus is principally Australian, but most of the Australian species are interrelated among themselves (and probably represent separate evolu- tionary radiations). There are certainly very few, perhaps only one, separate pri- mary bond(s) of relationship between thci New Guinean and Australian members of the genus. In making analyses and geo- graphic comparisons, the zoogeographer must decide whether the New Guinean Demetrida are to be treated as 59 units with Australian relationships or as one unit with Australian relationships. This de- cision makes a substantial difference in the conclusions. This genus compriseSj nearly 9 per cent of all known New Guinean carabid species. If it is decided to count all the species separately in making faunal comparisons, this decision alone in- creases the "Australian" fraction by about 9 per cent of the whole fauna. But if it i^ decided to count Demetrida in New Guinea as one geographic unit, its weight in the] sum of geographic relationships of the New Guinean carabid fauna is a fraction of 1 per cent. The decision how to count Demetrida is even more important ir I The carabid beetles of New Guinea • Darlington 179 analysis of the geographic relationships of the arboreal fraction of the carabid fauna, for Demetrida makes up nearly one- third of the arboreal Carabidae of New Guinea! My solution of the complex problem of choice of units for zoogcographic analyses and comparisons is two-fold. First, I think it is essential to describe situations in words, giving counts of genera, species, and other units, but going beyond mere presentation of figures. I shall do this for a succession of tribes of Carabidae that occur in the Asiatic-Australian area [32ff]. And second, after the situations have been described, I think arbitrary decisions have to be made and the reasons for them given. In the present case I think the decision should be to base statistical analyses pri- marily on geographic units — stocks with independent geographic relationships — be- cause they best show the degree of actual interrelationship (and also the geographic histories ) of faunas. In Coptodera and Demetrida (above) the geographic units — the number of separate bonds between the New Guinean carabid fauna and other famias — are seven and one (or very few) respectively. Decisions about geographic relationships are complicated by differences in degree of relationship. For example, Cilleniis (Part I, p. 399) is represented in both the Orient and Australia, but the New Guinean spe- cies is evidently more closely related, to Oriental than to Australian forms. I have scored it as one (Oriental) geographic unit; and in other similar cases in which single stocks have relatives in more than one region, and in which closeness of re- lationship varies, I have scored only the closest relationship in each case. When, however, relationships seem equally close with (say) Oriental and Australian forms, I have scored both (see following para- graph). Obviously these decisions are arbitrary in principle and often also in practice. Another complication occurs in cases like Mecyclothorax (Part I, p. 505; present part, Taxonomic supplement) and Scopodes altus (Part III, p. 198), in which relation- ships with Oriental and Australian forms seem about equally close, but in which the related forms are numerous in one region (in Australia, in the examples given) and relatively few in the other region. In such cases I have disregarded numbers and have scored geographic units solely ac- cording to apparent closeness of relation- ships. Mecyclothorax and Scopodes altus therefore each score one (Oriental) and one (Australian) geographic unit. Statements of existing relationships are still further complicated by great differ- ences in extent of the areas occupied by different genera, species, or other units. For example, Demetrida as a genus occurs throughout New Guinea and throughout Australia. If the New Guinean species are treated as a single geographic unit (see above), it is surely an Australian-related one. But besides the underlying Austra- lian relationship of all the New Guinean Demetrida, a second link with Australia is formed by one New Guinean species which occurs also on the extreme tip of the Cape York Peninsula (see Demetrida angidata. Part III, pp. 143, 159). Is this to be counted as a second Australian- related geographic miit in spite of the fact that the area occupied in Australia is very small? Or take the case of Syleter (Part I, p. 356), a genus which ranges from the southeastern comer of Asia to the Philippines and New Guinea, with the single New Guinean species occurring again on the extreme tip of Cape York. In terms of present distribution is this to be counted as one Oriental-related unit and also as one Australian-related unit, and if so, are the two units to be given equal weight in spite of the different extents of distribution in the Oriental Region and Australia? I think that the solution of this problem too (and of others like it) is, first, to make the situation clear, and then to make an arbitraiy de- 180 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 cision, and give the reason for it. In the present case my decision is to count as Australian-related only those geographic units that occur below the Cape York Peninsula in Australia, and to treat sepa- rately those New Guinean species that ex- tend only to Cape York ( see Table 9 [63] ) . This distinction seems to me to reveal im- portant differences in distribution patterns (and to facilitate deductions about geo- graphic histories). In the other direction, it seems to me most useful to treat as Oriental-related those geographic units that occur in Celebes or the Philippines as well as units that occur in the Oriental Region proper, but to treat separately those that occur west onl\ to the Moluccas [62; 64, footnote]. The best way to establish a basis for zoogeographic analysis of the New Guinean carabid fauna — the way that will give the most information — is, I think, to describe the relationships of New Guinean carabids tribe by tribe, emphasizing the separate geographic units as far as they can be recognized, including but not overempha- sizing counts of genera and species, and adding other significant details where pos- sible. I can do this only because I myself did much of the taxonomic work from which the details are derived. Most tribes of Carabidae are widely dis- tributed. A few small tribes are localized in various parts of the world, but no tribe is confined to or represented mainly in New Guinea. As background to understanding the relation of the New Guinean carabid famia to other famias, I shall begin by simimariz- ing the distributions of certain tribes which do not occur in New Guinea but which form part of the broader pattern of geographic relationships and of change of faunas from Asia to Australia. First will be noted tribes which (in this part of the world) occur only in Asia above or chiefly above the tropics, then tribes which occur in tropical Asia and extend toward but not to New Guinea, then a few tribes that occur in both Asia and Australia but skip New Guinea, and finally tribes that (in the part of the world under consideration) occur only in Australia. And then I shall consider the tribes that are represented in New Guinea, taking them in the order of the Coleoptewrum CataJoiius, and giving in each case all the geographically signifi- cant details that I can. The nature (size, wings, etc.) of the members of each tribe will usually be noted too, and also major habitats. (Although this survey is con- cerned only with existing distribution pat- terns, some details will be referred to again in discussion of the origins and di- rections of dispersal of the New Guinean carabid fauna. ) [32] Distribution of earahid tribes from A.sia to Australia. I shall now attempt to summarize, mainly by tribes, the distri- bution of Carabidae along a strip of the earth's surface extending almost from pole to pole, from northern Asia south and east across the Indo-Australian Archipelago (in- cluding New Guinea) to southern Aus- tralia. ( See Fig. 1 for a map of the tropical portion of this area.) Details of distri- bution in the islands and Australia are mostly from my manuscript lists (see [4]). Asia north of the tropics possesses several tribes of Carabidae wliich do not reach tropical Asia south of the Himalayas or are very poorly represented in the tropics. These northern tribes include Cychrini, Nebriini, Notiophilini, Opisthiini, Elaphrini, Loricerini, Patrobini, Amarini, and Zabrini. Two additional tribes well represented in Asia north of the tropics but not or poorly represented in tropical Asia, but present in Australia, are the Carabini and Broscini; these tribes are referred to again below. Several genera of Carabidae are outstand- ingly dominant in temperate Asia but absent or veiy poorly represented in the tropics; they include Carabus, BemI)idion (which occurs also in Australia, see be- low), and llarpalu.s. Asia north of the tropics is notable also for possessing relatively numerous Pterostichini but rel- The carabid beetles of New Guinea • Darlington 181 atively fewer Agonini and Lebiini. The mainly northern tribes and genera of Carabidae include many mesophiles and many hydrophiles but few arboreal forms. Tropical Asia and associated continental islands (Sumatra, Java, Borneo) possess a rich and diverse carabid fauna. Six small tribes are represented by single genera on the mainland of tropical Asia (most are represented in Africa too) but do not reach the islands; they are Enceladini, Melacnini, Anthiini, Disphaericini, Grani- gerini, and Idiomorphini. One additional tribe of a single genus, the Mormolycini, is confined to the Malay Peninsula, Sumatra, Java, and Borneo. Other tribes represented in tropical Asia (and more or less widely distributed elsewhere) extend for varying distances eastward across the Indo-Aus- tralian (Malay) Archipelago. The Hiletini (Camaragnathini) (one genus) have been found ( in the area in question ) only on the southeastern corner of Asia, Sumatra, and Borneo; the Omophronini (one genus), on the mainland of Asia and on Luzon in the Philippines (Darlington, 1967). The Sia- gonini (one genus) are numerous in tropical Asia and diminish eastward, the easternmost knowm species occurring on Java and on Mindoro in the Philippines (Darlington, 1967). And the Orthogoniini (one principal genus), with many tropical Asiatic species, diminish eastward, reaching the Philippines and Moluccas but probably not New Guinea. Some important genera of other tribes have similar patterns of diminution from tropical Asia eastward. For example, the dominant, widely dis- tributed genus Scarites (large fossorial Carabidae) is well represented in the tropics of Asia, has tvvo species in the Philippines, at least one on Celebes, and one on Timor, but does not reach New Guinea or Australia. The primarily Oriental Trigonotoma (rather large, ground-living, mesophile Pterostichini ) reaches the Philip- pines and Moluccas but not New Guinea or Australia. And CoIIida (medium-sized, arboreal Lebiini) occurs in tropical Asia (and other tropical regions) and eastward to the Philippines and Celebes but not New Guinea or Australia (where its place is taken by Demetrida). Asia and Australia share several tribes and important genera that are absent on most or all of the intervening islands in- cluding New Guinea. In the tribe Carabini, the genus Calosoma (large, ground-living, mesophile and xerophile Carabidae) is well represented in Asia above the tropics (and one or two species reach the tropical part of India), and two species occur in Australia, but the genus does not reach the southeastern corner of Asia and does not occur on the islands between Asia and Australia (except for the extension of one Australian species to New Caledonia and to the eastern Lesser Sunda Islands); this enormous gap in distribution is difficult to explain, for the genus does occur con- tinuously across the tropics in Africa and America. Tlie tribe Broscini (medium- sized and large, ground-living or jDartly fossorial, mesophile and xerophile Carab- idae) includes several genera in temper- ate Asia and several others in southern Australia, but few enter even the edges of the tropics, and none occurs on the islands between Asia and Australia (see Ball, 1956; and Darlington, 1965: 39-42, for discussion of the genera and zoogeog- raphy of this tribe). Pogonini (rather small, ground-living Carabidae often as- sociated with saline or alkaline habitats), too, include several genera on the main- land of Asia and several in Australia, including some in the tropics in both places, but are nearly or quite absent on the intervening islands. T\\'o tribes, Apo- tomini and Amblystomini (each consisting of one principal genus of small, ground- living, mesophile Carabidae), are well represented in Asia and extend eastward across the islands to the Philippines and Celebes or the Moluccas and occur again in Australia but probably do not occur in New Guinea; both have been recorded from New Guinea, but probably in error 182 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 (Part I, p. 398; Part III, p. 19). And the following genera of small Carabidae (all inclnding both hydrophile and mesophilc species) are dominant north of the tropics, extend into (different parts of) the west- em Indo-Australian Archipelago, and occur again in Australia, but are absent in New Guinea: Dyschirius (see under tribe Scari- tini, below): Bembidion (excluding Cil- lenus, see under tribe Bembidiini); and Trechus (see under tribe Trechini). Finally, five tribes are (in the region under consideration) confined to Austra- lia: Pamborini (one genus), to eastern Australia from the base of the Cape York Peninsula to south-central New South Wales; Migadopini, Merizodini, and Agoni- cini (with two to four genera each in Australia), to the southeastern comer of Australia and Tasmania; and Cuneipectini (one genus), to southwestern Australia. These tribes all consist primarily of ground- living mesophiles. They include both large (Pamborini, Cuneipectini) and small (Meri- zodini) forms. All existing Pamborini, Agonicini, and Cuneipectini have atrophied wings; the other tribes are winged or in- clude winged fomis. I shall now proceed from the tribes that occur in adjacent areas but that do not reach New Guinea to tribes represented on the island itself, taking them in the order of the Coleopterorum CataJo'^ius (Csiki, 1927-1933). AU the tribes rep- resented on New Guinea are primarily winged, although some include secondarily flightless forms. See again Figure 1 for a map of the area chiefly concemed in the following descriptions of distributions. [Although tiger beetles (Cicindelinae) are not included in my summary and analysis of New Guinean Caribidae, their distribution may usefully be summarized parenthetically. The tribe Collyrini (Part I, p. 334) is confined to the Oriental Region except for extensions eastward described below. The tribe consists of only two genera, both arboreal. CoUyris, of which the species are winged and (probably) live on foliage, is rich in species throughout the Oriental Region and the Philippines, with one spe- cies on Timor and one across the central part of Wallace's Line on Celebes (and supposedly reaching the Aru Islands) but none on New Guinea. Tricondyla, which is flightless and lives on tree trunks, is also well represented throughout the Oriental Region; one species is endemic on Celebes and two occur on the Philippines; and one of the Philippine species extends south and east to New Guinea (Part I, p. 334), the Solomons, and Timor, and has been found in the mid-peninsular rain forest of Cape York, Australia, although it apparently does not occur in the forest on the tip of Cape York. For further com- ments on the distribution (and history) of this species see also footnote, section [21]. The tribe Megacephalini (Part I, p. 335) consists of terrestrial, chiefly nocturnal tiger beetles. The principal genus is Mega- cephala, which is discontinuously dis- tributed in the wanner parts of the world In the Asiatic-Australian area, one species extends into northern India (from the Mediterranean region); otherwise the genus is absent in Asia and absent on the islands between Asia and Australia, except that two (winged) Australian species are doubtfullv recorded from southern New Guinea (Part I, p. 336); but about 20 species occur in Australia. The tribe Cicindclini (Part I, p. a36) includes chiefly winged, terrestrial and subarboreal, chiefly diurnal tiger beetles The tribe is worldwide in distribution. 01 smaller genera, Prothyma is represented in New Guinea by one species with probably Oriental relationships (Darlington, 1947) Caledonomorpha, with two closely relatec species, is endemic to eastern New Guinea, I do not know its geographic relationships' Distipsidera is an eastern Australian genu.' of which two species have been describee from southern New Guinea; these specie; are not directly related to each other bu The carabid beetles of New Guinea • Darlington 183 Form two separate Australian geographic the AustraHan units being of doubtful Linits; all members of the genus live on tree occurrence in New Guinea. However, the trunks, so far as I know. And Therates is arboreal Cicindelinae that live in rain in Oriental genus of which five or more forest (Tricondyla and Therates, with to- ipecies, representing at least three Oriental gether four geographic units) are all geographic units, occur in New Guinea, Oriental; while the terrestrial tiger beetles although none reaches Australia; these in- of New Guinea are at least as much Aus- ;ects live on under-story foliage in rain tralian as Oriental in present relationships, -orest. The single remaining genus of the and the Australian units include apparendy ribe ( so far as New Guinea is concerned ) older stocks as well as ( presumably more s the dominant, cosmopolitan Cicindela recent) shared species.] Tart I, p. 340), which includes many terres- [33] The pantropical OZAENINI (Part :rial as well as ( in the tropics ) subarboreal I, p. 351) includes medium-sized and )r arboreal species. New Guinea possesses small Carabidae which live on the ground 37 known species and three additional sub- usually in forest and are often associated species of this genus, representing at least with logs and rotting wood. Four winged 3ight separate geographic units (Part I, p. genera occur in the Oriental Region and 341). Two of the (older?) units are one very distinct (primitive?), flightless, \ustralian in present relationships. One endemic genus is confined to eastern )r more stocks of very small, at least partly Australia. Two of the Oriental genera, irboreal species have radiated on New Danija and Eiistia, reach Java and Borneo, Guinea; their geographic relationships are and the Philippines, but do not cross lot clear. Two Oriental stocks are rep- Wallace's Line to Celebes, but a third •esented on New Guinea by endemic spe- Oriental genus, Pseiidozaena, extends east- :ies {maino and denticoUis) but do not ward to New Guinea etc. but not Australia, ■each Australia. Cicindela discreta occurs The single species of this genus on New From Sumatra etc. to New Guinea and Guinea is the only member of the tribe lorthern Australia; C. decern fi,uttata, from there. ::elebes to New Guinea etc. but not Aus- [34] The tribe PAUSSINI (Part I, p. :ralia; and C. semicincta, in New Guinea 353) contains medium-sized and small, ;tc. and northeastern Australia and on mostly winged, chiefly ground-living, pri- Sfew Caledonia. marily myrmecophilous carabids derived In summary of the geographic relation- from Ozaenini. The tribe is pantropical, >hips of New Guinean Cicindelinae: of but is best represented in the main part Gollyrini, the island possesses one Oriental of the Old World. Ten genera occur in >pecies which occurs also on mid-Cape the Oriental Region; all of them extend York; of Megacephalini, two Australian east to Java and/or Bomeo and in some species (doubtfully) recorded from south- cases the Philippines, but none is known ?m New Guinea; and of Cicindelini, from Celebes; one species of the Oriental Prothyma with one and Therates with at genus Etiplatyrhopoliis (Part I, p. 354) least three Oriental geographic units and has been recorded from New Guinea, but Cicindela with apparently four Oriental its occurrence there is doubtful. One prin- ind five Australian units. The presence cipal genus, Arthropterus, occurs in Aus- of an endemic genus confined to eastern tralia; it is primitive and is not directly New Guinea and the occurrence of an related to any existing Oriental genus; extensive radiation of very small species of one species of it has been described from Cicindela on the island are also note- (southern?) New Guinea and may be the worthy. Tlie totals are nine Oriental and only paussid really present on the island, seven Australian geographic units, two of [35] The tribe SCARITINI (Part I, p. 184 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 355; Part IV, Tax. suppl ) is the principal, dominant tribe of terrestrial-fossorial Carab- idac. The trilie is worldwide in distri- bution although much more diverse in the tropics than in temperate regions. About 20 genera are represented in the Oriental Region and about 17 in Austraha, but only three genera have been found in New Guinea. For zoogeographic purposes, the large and small members of the tribe should be grouped separately. Oriental scaritines include about 11 genera of large and nine of small forms. Among the large fornis is Mouhotia, a fhghtless genus of three species confined to the Indochinese Peninsula; this genus is apparently related to the American Pasimachus and perhaps more distantly the Australian carenums (Biinninger, 1950). Most of the other Oriental genera of large scaritines are confined to the mainland of Asia or to Ceylon, but Oxy<^nathus reaches Java; Dlsiichus, Java and Borneo; Thli- bops, Java and the Philippines; and Scarites, limits given in [32]. The small Oriental scaritines are less well known. Most genera of them are apparently local- ized within the Oriental Region (some are represented also in Africa), but the pri- marily northern Dyschirius reaches Java and Luzon and occurs again in Australia (but not in New Guinea), and Syleter and Clivina are discussed below. Australian Scaritini include about 12 genera of large and five genera of small forms. The large forms are mostly "care- nums," which include about 11 genera and several hundred species, all flighdess, all confined to the continent of Australia and closely associated continental islands (and all probably products of radiation in Australia); a few reach Cape York, but none has yet been found on New Guinea. The carenums may be related to Mouhotia (above) in a general way, but probably not directly and not closely. The only other large scaritines in Australia are a few winged species of Geoscaptus, one of which extends to New Guinea. Small Australian scaritines include three monotypic, en- demic genera as well as a few Dyschirius (see above) and many Clivina (see be- low); these genera are all primarily winged, although the wings have atrophied in some Clivina. It will be seen that there is no direct or at least no close relationship bet\\'cen the large Scaritini of the Oriental Region and of Australia, but that a few genera (and even some species, see be- low) of smaller, winged forms are com- mon to the Orient and Australia. New Guinea itself possesses only one Australian species of Geoscaptus (the only "large" scaritine on the isUmd ) ; one species of the African-Oriental genus Syleter, be- longing to an Oriental species group, and extending to the tip of Cape York; and about 30 known species of the worldwide genus Clivina. Of the latter, zehi, wallacei, and basalis occur also in the Oriental Region and subfusa apparently has Ori- ental relationships; basalis, selhta, and jerru'^inea occur also in Australia, and zchi and inopaca reach Cape York; toxopei and vii!,il apparently have Australian rela- tionships; and a small radiation in the "australasiae group" has occurred or is oc- curring in New Guinea, the group ap- parently having Australian relationships. The relationships of some of these species are doubtful, but zebi clearly represents an Oriental stock that extends to New Guinea and Cape York. In summary: of large Scaritini, New Guinea possesses no Oriental and only one Australian species; but of small Scaritini four species that occur also in the Orienta area (as here defined) and two additional Oriental relationships, and three specie' that occur also in Australia, two more thai reach only Cape York, and three additiona Australian relationships; and some radi- ation has occurred in one of the Australian related stocks. [.36] The tribe BEMBIDIINI (Pari I, p 398) contains small, chiefly ground-living often water-loving (hydrophilc) carabids There are two principal genera widel) The carabid beetles of New Guinea • Darlington 185 distributed over the \\'orld including New Guinea, and a few smaller genera of which only one is represented in New Guinea. Bcmbidion has very many species in the north-temperate zone including temperate Asia, fewer in south-temperate areas in- cluding five in Australia (Darlington, 1962a), and still fewer in the tropics. Several temperate Asiatic stocks of the genus extend into the western part of the Malay Archipelago, reaching Celebes and/ or the Philippines (Darlington, 1959a); one species (sobriniim Boheman) may range more or less continuously from Asia to Australia along the Lesser Sunda Islands (Darlington, 1959a: 339-340); and one southern Australian species extends into the tropical northern part of Australia; but none of these reaches New Guinea. However, the island does have one Bem- bidion of a different stock, a member of the coastal (including intertidal) subgenus (or separate genus) Cillenus. This stock extends from Asia to Australia and New Zealand; the New Guinean species is re- lated to Oriental rather than Australian fomis. The absence of more-ordinary Bcmbidion in New Guinea is especially noteworthy. The other principal genus of Bembidiini, Tachys (Part I, p. 400; Part IV, Tax. suppL), is virtually cosmopolitan, but is best represented in the warmer regions of the world. Many species occur in tropical Asia and many in Australia, and some species range from Asia to Australia, but the Oriental and Australian Tachys faunas are remarkably different as wholes. New Guinea possesses 68 species, of which the geographic relationships (so far as they can be determined) are summarized in Part I, in notes under the genus and in discussions under the ten species groups. The New Guinean species include 17 that occur also in some part of the Oriental area (which for present purposes includes Celebes and the Philippines) plus three additional stocks related to Oriental forms, and eight species that occur also in Aus- tralia. (Several species that range from Asia to Australia are included in both these totals. ) The third bembidiine genus in New Guinea, Limnasiis, is widely distributed in the Old World and reaches the West Indies and Hawaii. It is represented in New Guinea by two winged, eyed species, \\'hich range respectively from southern Asia to Australia and from southern Asia to New Guinea, and by one endemic, flightless, blind species known only from the Bismarck Range. (No aniline Bem- bidiini have yet been found in New Guinea, but some probably occur there. They are minute (often c. 1 mm long), blind carabs which usually occur in soil and which are usually obtained only by special collecting methods.) In summarv. New Guinean Bembidiini include 19 species ^^'hich occur also in the Oriental area and four additional stocks with Oriental relationships, and nine spe- cies which occur also in Australia. [37] The TRECHINI (Part I, p. 487) are small, hydrophile and geophile cara- bids. In this tribe. New Guinea is notable principally for what it lacks. Tlie tribe as a whole is worldwide. Terrestrial, meso- phile members of the tribe are numerous in the north-temperate zone and numerous also south of the tropics in southeastern Australia ( and in southern South America ) ; many of those in both Asia and Australia are now flightless, but winged species occur too in both places; the ancestral fomi(s) presumably dispersed by flight. A few species of Trechus have been found at high altitudes in northern Luzon in the Philippines (Darlington, 1959a), but other- wise mesophile trechines are entirely unknown in the Malay Archipelago, includ- ing New Guinea, at any altitude. Their absence in New Guinea is remarkable. However, hydrophile, stream-side trechines are sparingly represented in New Guinea. Of the widely distributed Old-World ( and West Indian) genus PcriJeptus, one Ori- ental species reaches western New Guinea; 186 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 and Perileptodea is confined to New Guinea, except that one of the New Guinean species occurs also in the Solo- mons. It is noteworthy that PeriJeptiis (above), although it occurs from Asia to western New Guinea and again in Aus- tralia, is apparently absent in the main part of New Guinea, where its place may be taken bv Perileptodcs. [38] The tribe PANAGAEINI (Part I, p. 492; Part IV, Tax. suppl.) includes chiefly medium-sized, terrestrial, mesophile forms, widely scattered over the world. Geographically significant Asiatic genera not represented on New Guinea include winged Fana<^acus, which occurs around the north-temperate zone (and in Central America and the West Indies) including temperate Asia and Japan but not in tropical Asia; flightless Bracht/onychus, confined to the tropical southeastern corner of Asia; and Avinged Euschizomerus, an African-Oriental genus that reaches Java, Borneo, and the Philippines. Five genera have been found in New Guinea, all the New Guinean species being winged except the Craspedophorus. Of Trichisio, the one New Guinean species is supposedly en- demic but closely related to both Oriental and Australian species. Of the tropical Oriental (and African) genus Dischi^ssiis, one widely distributed Oriental species reaches New Guinea. Of the African- Oriental Microcosmodes (Microschemtis), one species (doubtfully assigned to the genus) occurs in New Guinea and Austra- lia. The Oriental genus Peronomcrus is represented in New Guinea by one Oriental and one endemic species, the latter ap- parently representing a second Oriental stock. And Craspedophorus, although well represented in both the Oriental Region (and Africa) and Australia, is known in (eastern) New Guinea by only one en- demic species (a single individual) prob- ably related to Australian rather tlian Oriental forms. (Although most species of Craspedophorus including probably all the Australian ones are now flightless, at least one of the Oriental (Philippine) species i; still winged.) In summary. New Guineai panagaeines include two Oriental specie; and two additional species with Orienta relationships, and one Australian specie: and two additional species with Australiai relationships. [39] The tribe PTEROSTICHINI (Par I, p. 497; Part IV, Tax. siippl.) includes i large number of carabids, diverse in size some winged and some not, many of then mesophiles, some hydrophiles, but ven few arboreal. Tlie tribe is worldwide bu is both taxonomically and geographically diverse, with different genera distributee in many different patterns. The Australia! genera of the tribe have been reclassifiec by Moore (1965), who previously (1963 transferred some supposed pterostichine: to the tribe (or subfamily) Psydrini. O the latter (primitive?) group, 18 genen are now known in Australia (Moore, 1963) Nomitis occurs in North America, Europe and locally in Africa; and Psydrtis is ii North America. The only psydrine genu; that reaches New Guinea is Mecyclothorax which (for statistical purposes) I hav( continued to count as a pterostichine. In the north temperate zone, mesophih pterostichines are dominant carabids; tem perate Asiatic genera that do not ente: the tropics are too numerous to list. Oi the other hand, only a few, small generj are confined to the tropical Asiatic main land. A few additional Oriental genen reach Wallace's Line but not Celebes Moriouidiiis (two species) is known onh from Tonkin ( North Vietnam ) and Borneo Diceromenis (one species), from India t( to Java and Timor; Dicaelindus (few spe cies), from Ceylon and Burma to Java anc Borneo; and Metabacetiis (few species) from Burma to Java, Borneo, and th( Philippines. (The pterostichine genera bes represented in tropical Asia and the west ern Malay Archipelago are actually Lesti ciis, Trigonotoma, and CaeJostomus (meso philes), Ahacetus (mostly hydrophiles) and Morion (subarboreal), all referred t( The carabid beetles of New Guinea • Darlington 187 gain below.) The mesophile genus Trigo- lotoma, which includes some winged spe- ies and which is primarily Oriental, occurs ast to Celebes and the Moluccas but not 0 New Guinea. In Australia, mesophile pterostichines are gain dominant, but the genera are almost 11 different from those of Asia. Australian ;enera that do not reach New Guinea are 00 numerous to list. Genera of Pterostichini that do occur in ■Jew Guinea may, for zoogeographic pur- loses, be grouped according to habitats, ^ost are winged; some -w species occur 1 primarily winged genera, and some small ndemic genera are now wholly -w, but 11 are or may be derived from ancestors hat were winged when they reached New Guinea. Ten genera of New Guinean Pterosti- hini are (or probably are) mesophiles. Tiey include two endemic, mountain-living ;enera, of which Rhytiferonia (with two pecies) may have Australian relationships, nd Analoma (Paraloma) (with four spe- ies). Oriental relationships. A third en- lemic genus, HapJofcroniu, with one pecies, may be related to (derived from) Australian ) Loxandrus ( for which see inder hydrophiles, below). Monotypic h-achijdius ranges from southeastern Asia 0 New Guinea etc. but not Australia- In he African and Oriental genus Caelosto- mis, picipes ranges from Asia to Australia perhaps carried by man); the other four 'Jew Guinean species are endemic but epresent at least one additional Oriental geographic unit; and one of the endemic pecies reaches Cape York. Meci/clotliomx Part I, pp. 498, 505; present part [SO], ^ax. siippl. ) , is mainly Australian ( and iawaiian) but two species occur at very ligh altitudes in New Guinea, and related pecies, on mountains in Java; I score this IS one Australian and one Oriental geo- graphic relationship. Catadromus is a nainly Australian genus of very large :;arabids of which two individuals of one Australian species have been taken in New Guinea; this species occurs also on Java; I think it may be carried by man; however, in terms of present distribution, it repre- sents both one Australian and one Oriental geographic unit. Prosopogmus is mainly Australian; three endemic species on New Guinea comprise one Australian relation- ship; and one species is on the Moluccas. Lesticus is a primarily Oriental genus with eight New Guinean species; one of them occurs in Australia (and is the only Lesti- cus there); the other seven are endemic but their relationships are Oriental, and I count them as one Oriental unit of relation- ship. And the Oriental genus Cosmodiscus includes two New Guinean species which have independent relationships with Ori- ental species, and one of the New Guinean species occurs also in Australia. In sum- mary: the mesophile Pterostichini of New Guinea apparently include three species shared with the Orient plus six additional Oriental relationships; four species shared with Australia plus four additional Austra- lian relationships; and one New Guinean species that reaches Cape York. Six genera of Pterostichini on New Guinea are primarily hydrophiles, or prob- ably so. Two of them, endemic Homa- loncsiota and Nchriojeronia (with together three species), probably live beside run- ning water; they are related to each other and to Loxandrus (below); I count them as one Australian unit of relationship. Of genera associated with standing water, the African-Oriental-Australian genus Aha- cetus is represented on New Guinea by two principal Oriental species, one of which reaches Australia; the primarily Australian Platycoelus, by four endemic species, probably representing at least two Australian relationships (one species reaches the Moluccas); Australian-Amer- ican Loxandrus (Part I, pp. 498, 549; present part [82] ), by four endemic species probabK' representing at least two Austra- lian and one Oriental ( Celebes ) geographic relationships; and Tiferonia, by one en- demic species, the only other known species 188 Bulletin Museum of Comparafive Zoology, Vol. 142, No. 2 of the genus being Oriental (Philippine), of which one Oriental species reaches New In summary: h\dr()philc Pterostichini in Guinea. Other agonine genera that are New Guinea include two Oriental species well represented in temperate Asia but are and two additional Oriental relationships; absent or few in the tropics include Cala- one species shared with Australia and thus, Sphodnis, etc., and Syniichus (Lind- probabK' at least five additional Australian roth, 1956). relationships; and the Australian-related Most of the agonines of tropical Asia I.oxandrus and its apparent derivatives, and the western part of the Malay Archi- llomalouesiola and Nehrioferunia and per- pelago are now placed in the catch-all haps T if crania, have apparently radiated genus CoJpodes (Louwerens, 1953); their on New Guinea, while the Oriental-related classification is difficult and their geo- hydrophile pterostichines have scarcely graphic relationships are largely unknown; done so. the few distinct, small. Oriental agonine The single subarboreal (bark-and-log- genera include Onycholahis and Dirotiis, inhabiting) genus of Pterostichini in New represented from the Malay Peninsula to Guinea is pantropical Morion. The tvvo Java and Borneo, with the foiTner genus New Guinean species belong to a primarily reaching also the Philippines. Oriental rather than Australian group of The genus Homothes (including Aeolo- the genus; one of the species is endemic dcrmus) is discontinuous, with one species and the other extends to Australia. In distributed from the Malay Peninsula to terms of present distribution, the New Celebes and the Philippines, none in New Guinean members of the genus score one Guinea, but several in Australia. Other- Oriental relationship and one species wise, the extremely limited Australian shared with Australia. agonine fauna includes only the endemic. The sum of existing geographic relation- monotypic, flightless Odonta'j^onum (Dar- ships of all New Guinean Pterostichini lington, 1956) in North Queensland, and (three preceding paragraphs) is five spe- a small number of species all of which I cies shared with the Orient and nine ad- tentatively assign to Notagonutn or Col- > ditional Oriental relationships, six species podcs. i shared with Australia and nine additional Known New Guinean Agonini (Part II; Australian relationships, and one additional present part. Tax. stippl.) now total 21 species extending to Cape York. genera and 160 species. They are medium- [40] The distribution of the tribe sized Carabidae, diverse ecologically. All AGONINI is discussed in the present part those found at low altitudes in New Guinea [92]. The tribe is worldwide, better repre- are winged; many of the mountain-living sented in the tropics than in the temperate groups now have atrophied wings (but all zones, and especially dominant in New may be derived from ancestors that were Guinea (Fig. 13 [64]), but deficient in \\'inged when they reached New Guinea). Australia. They may be grouped according to habitat, In the north-temperate zone, agonines i^s follows, arc moderately numerous and include the Mesophiles among the New Guinean i dominant, primarily winged, hydrophile Agonini include the Oriental genus Arhy- and mesophile genus Ai:,onuJ7}, with nu- tinu.s, with three endemic New Guinean merous subgenera; the geographic limits of species, representing at least two Oriental this genus are doubtful (because of doubt stocks, and Tarsagonum, with one endemic about the assignment of some tropical species, the genus otherwise known only species), but the only subgenus that ranges from Borneo; these two genera occur at far into the Oriental tropics is probably low altitudes. Also mesophiles are certain Sericoda ( pres(>nt part [80], Tax. suppL), species of the "genera of convenience" The carabid beetles of New Guinea • Darlington 189 Notagonum and Colpodes (of which the geographic relationships, of the mesophile species, have not been determined in detail) and some or all the species of nine endemic genera, most but not all of which are momitain-living, and most but not all of which have atrophied wings; this whole assemblage is Oriental in general relationships, but I cannot decipher the separate geographic units. Hydrophiles among the New Guinean Agonini include at lower altitudes the single New Guinean Lorostemma, which is closely related to an Oriental species; some species of Notaganum; and the one species (several subspecies) of the endemic genus Lithagomim; and at higher altitudes some additional Notagonuni; the endemic genus Potamagonum; and some species of the endemic genus Nebrkigomim. Excepting the Lorostemma (which has been derived independently from the Orient), these agonines may all be products of the com- plex radiation of Agonini that has occurred on New Guinea. I cannot state their geo- graphic ties in detail, except to repeat that they are all basically Oriental in relation- ships, and to add that two of the species of Notagonum occur also in Australia. These two are N. submetalliciwi (present part. Tax. siippl.) (which is distributed as if it has reached New Guinea from Australia) and N. dentellum (present part. Tax. siipp].) (which has apparently reached North Queensland from New Guinea). Arboreal agonines in New Guinea in- clude the African-Oriental genus Euplenes, with two endemic species probably repre- senting two Oriental geographic units; the Oriental genus Dicranoncm, of which the one species that occurs in New Guinea ranges also from southern Asia to Australia; the subgenus Sericoda of the primarily northern genus Agomwi, of \\hich one Oriental species reaches New Guinea (present part, Ta.x. siippL)-, Violagomnn, a mainly insular genus (Darlington, MS), with the one New Guinean species oc- curring also in Australia; and some species of the "genus of convenience" Colpodes, of which three New Guinean species occur also in the Oriental area and one reaches Australia. In summary of the geographic relation- ships of New Guinean Agonini: the meso- philes include four specific Oriental units of relationship; the hydrophiles, two spe- cies shared with Australia; and the arboreal forms, five Oriental species, two additional specific Oriental relationships, and three species shared with Australia. But it should be repeated again, and emphasized, that the whole diverse assemblage of Agonini on New Guinea is Oriental in general relationships, although only a few of the separate geographic units can be dis- tinguished. [41] The tribe PERIGONINI (Part III, p. 5) contains small, winged, mesophile carabids which live mainly in rotting logs and leaf htter. Only the worldwide genus Perigona {.semu Jato) is represented in New Guinea. It includes numerous species in the Oriental Region, 14 in New Guinea, and only five in Australia (Darhngton', 1964). Of the New Guinean species, five occur also in the Oriental area (which includes the Philippines) and three more have separate Oriental relationships, and one species occurs also in Australia. [42] The tribe LICININI (Part III, p. 14; present part, Tax. suppJ.) includes both medium-sized and small, mostly winged Carabidae, of which some are hydrophiles and some mesophiles. The tribe is nearly worldwide in distribution. A few genera, including Licinus, occur in north-temperate areas but do not reach tropical Asia. DipJochila {Remhiis) is widel\- distributed in the north and in Africa and the Orient; several species reach Java, Borneo, and the Philippines, but none farther east. Several genera are confined to or occur mainly in Australia, which has more licinines than any other part of the world. Five genera are represented on New Guinea: the widely distributed genus Badister, by one species which occms also 190 BuUctin Museum of Comparative Zoology, Vol. 142, No. 2 in the Orient and Australia; primarily Aus- tralian PJnjsolacstJius, by one species which occurs also in the Orient; monotypic Omestes, by O. torta, which is also in the Oriental area (Celebes, Philippines); primarily Australian Dicrochile, by one Australian and three endemic species rep- resenting at least one additional Australian relationship; and primarily Australian Microf crania (Part III, p. 18; present part, [80] ) by one species on the Bismarck Range in New Guinea, and one on moun- tains in Java. The sum of geographic units seems to me to be three species shared with the Orient (chiefly the Philippines) plus one additional Oriental relationship, and two species shared with Australia plus two additional Australian relationships. How- ever, regardless of these unit scores, all the New Guinean licinines except the Badister belong to groups that are characteristically Australian rather than Oriental. (See Ball, 1959, for further discussion of the classifi- cation and distribution of some members of this tribe.) [43] The tribe CHLAENIINI (Part III, p. 20; present part [9] ) includes medium- sized (less often small or large), primarily winged, hydrophile and mesophile Carab- idae. Tlie tribe is most diverse in Africa and Eurasia. A few small genera (each with one or very few species) are confined to temperate Eurasia or the Asiatic tropics. Very small chlaeniines of the African- Oriental genus CaUifitomimii.'i reach Java, Borneo, and the Philippines (and Timor), and of the Oriental genus Chlaeminus, Java and the Philippines. However, only the worldwide genus Chkienius (.sensii lato) crosses the central part of Wallace's Line to Celebes, New Guinea, and Austra- lia, and species decrease in number in this direction, Java having about 30, New Guinea 12, and Australia 10. Tlie New Guinean species include the following geographic units (see under the species in Part III, pp. 23ff for details): six species shared with the Oriental area plus three additional Oriental relationships, and four species shared with Austi^alia plus two additional Australian relationships. Several of the species range from the Orient to Australia or have close relatives in both, and score as both Oriental and Australian units. [44] The tribe OODINI (Part III, p. 30; present part, Tax. suppl.) contains mostly medium-sized carabids most of which are subaquatic, although a few have left the water and become terrestrial. Most are winged, but some terrestrial fonns have atrophied wings. Oodines occur in all the warmer parts of the world, but are rel- atively few in temperate areas. In the Oriental Region, Systolocrauius (also in Africa) and Holcocoleus are confined to the mainland, and Simous- (see under Oodes hevissimus. Part III, p. 34) ranges from India to Java and Borneo. Australia possesses numerous species of the endemic genus Coptocarpus, which reaches the Cape York peninsula but not New Guinea^ Only two genera are represented on New Guinea: Amitrichis, which is Oriental- Australian (and American, see [82]), and Codes, which is nearly worldwide. New Guinean species include the following geographic units (see under the species in Part III, pp. 32ff): four species that occur also in the Oriental area (as here defined ) and two species that occur also in Australia. The geographic relationships of the other species are undetermined. i [45] The tribe HARPALINI (Part III, p. 38; present part. Tax. suppl.) contains a large proportion of the common, medium- sized Carabidae that live on the ground especially in open country in all parts of the world, and the tribe includes also some common smaller forms that live especially in wet places. Most are winged. The dis- tribution of the tribe in the Asiatic- Austra-! lian area including New Guinea is best^ summarized by subtribes (see Part III, p ^ A Coptocarpus has now been found in Nev' Guinea (see Taxonotnic supplement, footnot< under Oodini ) . The carabid beetles of New Guinea • Darlington 191 39), although this classification is an over-simplification. One additional harpa- line not belonging to any of the following subtribes should be mentioned: one species of the Old- World-tropical genus Pachytra- cltehis (subtribc Pachytrachelina; Csiki, 1932: 1082) ranges from India etc. to Australia perhaps along the Lesser Sunda Islands, although the genus does not occur in New Guinea. Harpalini of subtribe Anisodactylina are chiefly medium-sized mesophiles. Tlie prin- cipal genus in the north-temperate zone is Anisodactylus (sensii lato), which is Hol- arctic; it enters the edge of the tropics in southeastern Asia but does not reach the tropical islands; however, two related, monotypic, primarily insular genera, Rluj- sopiis and Harpalomimetes, do reach Java and/or Borneo (Wallace's Line). Also in this subtribe is Chijdaeus (Part III, p. 47; present part. Tax. suppL), which occurs at high altitudes on the Himalayas and on mountains in Formosa, Sumatra, Java, the Philippines, and New Guinea, but does not reach Australia. It has apparently "mountain hopped" some 4,000 miles ( more than 5,000 km), by steps, from the moun- tains of Asia across the Malay Archipelago (see [80]). Three other genera of Ani- sodactylina are best represented in Austra- lia but reach New Guinea and extend westward into and across the Malay Archi- pelago, chiefly in relatively open country including open eucalyptus woodland, iil- though some species occur in rain forest too. Of these primarily Australian genera, Gnathaphanus has five species in New Guinea all of which occur also in Australia and two of which reach only the southern edge of New Guinea, the others reaching (respectively) West New Guinea, the Malay Peninsula, and India; DiapJwromerus, two endemic species in New Guinea closely related to (different) Australian species, one of the New Guinean species reaching also the Moluccas; and IlypJiorpox, one New Guinean species which apparently occurs also in northeastern Australia and extends west to Java and Sumatra. The geographic units of New Guinean Ani- sodactylina are therefore three species that occur also in the Orient plus one additional Oriental relationship, and six species that occur also in Australia plus two additional Australian relationships. (However, ex- cepting Chydaeus, the Anisodactylina that reach New Guinea are all primarily Aus- tralian, and they show an exceptionally clear pattern of spread westward across the islands.) Harpalini of subtribe Pelmatellina are represented in New Guinea by only three small, endemic, water-loving species of the diverse Australian genus Lecanomerus (Part III, p. 45); they may all be derived from one ancestor and I count them as one Australian geographic relationship. Harpalini of subtribe Harpalina are (in the area under discussion) primarily Asi- atic-Oriental and do not occur in Australia at all, except that one or two genera reach just the northern edge of the continent. Harpahis itself is a dominant Holarctic genus which occurs south to Java ( one spe- cies) but does not cross Wallace's Line. Oxycentrus reaches at least Celebes; Dio- ryche, at least Celebes and Timor; but these few details do not do justice to the numbers and complexity of limits of the Oriental Harpalina. Members of this sub- tribe are dominant on the ground in New Guinea, especially in rain forest, but the New Guinean Harpalina represent merely the fringe of the much richer Asiatic- Oriental fauna. The principal genus of the subtribe that does reach New Guinea is Trichotichmis (Part III, p. 48), with 14 species on the island showing at least three separate Oriental relationships {^^Carhanus," "Lampetes" and the others). Harpaloxenus, with five species on New Guinea, is ap- parently closely related to Trichotichntis and counts as at least one additional Ori- ental relationship. Lyter is a new, endemic, monotypic genus confined to New Guinea (and perhaps derived from Trichotichnus on the island). Coleolissus has two en- 192 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 demic species on New Guinea representing perhaps only one Oriental stock; one of the species of Colcolissii.'i is represented on Cape York. Finally, Platymctopus is represented on New Guinea by one Ori- ental (Philippine) species that apparently reaches only western New Guinea. In summary: the geographic units of New Guinean Harpalina are one species shared with the Orient (Philippines) and at least eight additional Oriental relationships, and one New Guinean species reaching Cape York. (This is a remarkably clear pattern of multiple dispersal of a dominant Asiatic- Oriental group south and east across the Malay Archipelago.) Harpalini of the subtribe Acupalpina are relatively small forms most of which live in wet places on the ground. Their generic classification is not very satisfac- tory; all the genera represented in New Guinea are widely distributed at least in the Old World, but their zoogeographic limits (except as they concern New Guinea) need not be discussed here. Of Egadwma, New Guinea possesses four species, of which three apparently occur also in both the Orient and Australia and the fourth is closely related to an Oriental species (present part. Tax. suppL). Anoplo- genius is represented in New Guinea by one species that probably ranges from the Orient (Sumatra) to Australia; Stenolo- phu.s', by t\vo species both probably present in the Oriental area and in Australia. Finally, the six New Guinean Acupalpus include apparently two species that occur also in the Orient plus two additional Oriental relationships, and one species that occurs also in Australia plus two additional Australian relationships (see details given in Notes- under the species, Part III, pp. 73ff). The sum of geographic units of New Guinean Acupalpina is therefore eight species plus three additional relationships Oriental, and seven species plus two re- lationships Australian. (But the pattern as a whole is clearly one of multiple dis- persal of Acupalpina from the Orient to New Guinea and Australia.) The sum of geographic units of New Guinean Harpalini of all subtribes is 12 species shared with the Orient plus 12 additional Oriental relationships, 13 species shared with Australia plus five additional Australian relationships, and one additional species reaching Cape York. [46] The small, pantropical tribe ANAU- LACINI (Part III, p. 76) contains small, winged, mesophile carabs which usually live in leaf litter on the ground in rain forest. New Guinea possesses four genera: Anauhicus is represented by one species, which is widely distributed also in both the Orient and Australia; Caphora, by one Oriental species that has been foimd also on the tip of Cape York; Acphnidius, by one Oriental species that does not reach Australia; and Odontomasorcus, which is endemic and of unknown relationships, by one species and one additional subspecies. The geographic units are therefore three species that occur also in the Orient, one also in Australia, and one additional occur- rence on Cape York. The few additional members of the tribe that occur in the Orient are geographically insignificant; no additional ones occur in Australia. [47] The CYCLOSOMINI (Part III, p. 78) is another small tribe of rather small, winged Carabidae widely distributed in! the warmer parts of the world. Of the fewi genera represented in tropical Asia, Tetra- gonodenis (which is widely distributed in' the Old World and the Americas) and Cijclosomus (which is African and Ori- ental) reach Java and the Philippines but not Celebes. These genera are terrestrial In Australia the tribe is represented only by numerous species of Sawthrocrepis Most of them live on shaggy tree trunks but one group of small species is foliage' arboreal, and this group is represented b> endemic species on New Guinea, Java, the Philippines, and some Lesser Sundi Islands. The New Guinean species is ap I The carabid beetles of New Guinea • Darlington 193 parently closely related to both Australian and Oriental (Javan, etc.) forms. [48] The tribe LEBIINI (Part III, p. 80; present part, Tax. suppl. ) is, among Carab- idae, equalled in number of species on New Guinea only by the Agonini. Ho\\'- ever, the Lebiini are far more numerous and diverse at low altitudes on the island and far more complex in their discernible geographic relationships. Most tropical lebiines are arboreal, and all the 160 New Guinean species except Nototarus papua are winged. In discussing the New Guinean forms in detail (below) I shall group them by major habitats. First, how- ever, I shall state briefly the distribution of genera that occur in neighboring regions but do not reach New Guinea. The Ori- ental forms are especially numerous and significant, but I cannot group them by habitats, because I do not know the habi- tats of many of them. Tlie north-temperate zone possesses com- paratively few Lebiini. Of the genera that do occur in temperate Asia, the most im- portant is Holarctic, terrestrial Cijmindls, a genus well represented in temperate Asia but which scarcely enters the Asiatic tropics. Tropical Asia possesses relatively more Lebiini, but only half a dozen or so small genera of the tribe are actually confined to the tropical Asiatic mainland. A much larger number of genera occur on the Malay Archipelago. Some of them are represented on the mainland of tropical Asia too or even in Japan, and some reach New Guinea (as detailed below), but about 15 do not reach New Guinea. Of these 15 genera that are represented in the western part of the Malay Archipelago but that do not reach New^ Guinea, 12 reach Java and/or Borneo (and sometimes also the Philippines and/or some of the Lesser Sunda Islands) but (so far as known) do not cross the central part of Wallace's Line to Celebes. These are mostly small genera of one or very few species; they include AUocota (three spe- cies), distributed from Burma to Java, Borneo, and the Philippines; Lehidia (four species), from eastern Asia and Japan to Java; and Dromius, with many species in the northern hemisphere, fewer in tropical Asia, and one species (probably unde- scribed) on Sumatra and two on Java. Two other, important genera of Lebiini do cross Wallace's Line but do not reach New Guinea: Lioptera, with about eight species, is distributed from southeastern Asia and Japan to Borneo, Celebes, and the Philippines; and Callida, well repre- sented in all the warmer parts of the world except the Australian region but including tropical Asia, reaches Java, Borneo, Cele- bes, and the Philippines, and also some of the Lesser Sundas. Australia has comparatively few Lebiini, and many of those that do occur in Aus- tralia are the same as or related to New Guinean species and are noted in my enumeration (below) of the geographic relationships of New Guinean genera. Additional significant lebiines in Austra- lia include FhiJophJoeus, with many species occmiing through most or all of the continent, chiefly on the shaggy trunks of eucalypts and other trees. Agonochila and Demetrido are numerous in Australia (and have apparently radiated separately there) chiefly on tree trrmks, but are represented on New Guinea too, of course. And FhJoeocarahus and Trigonothops (these t\\'o genera I think mainly on tree trunks) and Nototoriis and Anomotoms (I think chiefly on the ground ) are well represented in Australia too, although present also in New Guinea. Other genera of lebiines in Australia seem geographically unimportant. New^ Guinea has representatives of 32 genera of Lebiini, and the geographic re- lationships even of some single genera are complex. In discussing their relationships, I shall divide them into ecologic groups comparable to those into which I have divided some preceding tribes, although the basis of division is not quite the same. Among New Guinean Lebiini few genera 194 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 are terrestrial, none is hydrophilous, but many are arboreal; and the arboreal forms are divided, those that live on tree trunks and those that live in foliage being grouped separately. And several genera which do not fit into this ecologic classification or of which the habitats are unknown are noted last of all. Terrestrial (mesophile) or probably ter- restrial Lebiini in New Guinea include one species of the Asiatic-Australian genus Lachnoderma, the New Guinean species occurring also in Australia; one Oriental species of the Oriental genus Sintirus; one endemic species of Peliocypas, this being the easternmost representative of a diverse Oriental group; one Oriental-Australian species of Syntomiis; two species of Micro- lestes, one with Oriental and one with apparent Australian relationships; two en- demic species of Apri.stus, the easternmost representatives of this African-Oriental (and North American) genus, representing perhaps a single Oriental geographic unit; one endemic species of the Australian genus Nototaru.s; eight endemic species of the Asiatic-Australian genus Anomotarus, the geographic relationships of one of the species being Oriental and the others undetermined (except that one extends to the Moluccas); and one Oriental species of the Oriental genus Omohrus. In sum- mary, the terrestrial or probably terrestrial Lebiini of New Guinea include three spe- cies that occur also in the Orient plus four additional Oriental relationships, and two species that occur also in Australia plus two additional Australian relationships. New Guinean Lebiini that live mainly on tree trunks and on fallen logs include one endemic species of the Oriental genus Stenotelus; one Oriental species of Miscelus that has been found also in mid-Cape York, and two endemic species of the same genus representing at least one additional Oriental relationship; nine species of the mainly New Guinean genus Miniithodes, one of the new Guinean species having a relative in the Orient (Celebes) and one a relative in Australia; 14 species of the widely distributed tropical genus Catas- copus related as detailed below; one en- demic species of Pericahts, the easternmost (except for another endemic species on New Britain) of a species-rich Oriental genus; eight species of the widely dis- tributed genus Coptodera, detailed below; one endemic species of the Oriental genus Mochtherus; and two species of the pri- marily New Guinean genus Stricklandia, one of the New Guinean species reaching the Moluccas, with a closely related species in Australia. In more detail, the New Guinean Catascopus (Part III, pp. lOlff, see especially Notes under the genus) in- clude elegans, which ranges from tropical Asia to northern Australia; smaragdulus, from tropical Asia to New Guinea and mid-peninsular Cape York; facialis, from tropical Asia to Western New Guinea;, laevig^atus, which occurs also in the Moluc- cas and has a close relative in Australia; and aniensis, which occurs also in mid- peninsular Cape York. The geographic units of the genus listed in the preceding sentence are three species shared with the Orient; and one species shared with Aus- tralia, one additional Australian relation- ship, and two additional extensions to Cape York. (Nevertheless, Catascopus is most diverse in the Orient; several additional stocks have probably reached New Guinea in the past, although I cannot now dis- tinguish them, and all the few Australian species seem to represent more or less recent invasions from New Guinea.) Coptodera (Part III, p. llOff) includes endemic i!,rossa, with no recognized close relatives; cijaneUu, lincolata, eluta, and oxyptera, all of which occur also in some part of the Oriental area; ornatipennis, which occurs also in the Moluccas and has a relative in Celebes (Oriental area); and pajmella and tcau, \\'hich together form' one Australian-related unit: total units in this genus: four species that occur also in the Oriental area plus one additional Ori- ental relationship, and one Australian re- The carabid beetles of New Guinea • Darlington 195 lationship. Geographic units for all the tree-trunk-living New Guinean Lebiini are eight also-Oriental species plus six Oriental relationships, and one also-Australian spe- cies plus four Australian relationships, and three additional species reaching Cape York. Arboreal Lebiini that probably live mainly in foliage on New Guinea include the following: three endemic species of the Oriental genus Aristolehia perhaps representing one Oriental stock, one of the species occurring also on the tip of Cape York; seven species of the worldwide genus Lebia, their geographic relationships detailed below; one endemic species (the easternmost of the genus) of the Oriental genus Physodera; one endemic species of the mainly Oriental genus Holcodenis, the New Guinean species having close relatives in both the Orient and Australia; mono- typic Oxydontus, the one species occurring also in the Orient; 13 species of the mainly Oriental DoUchoctis, related as described below; monotypic Celaenephes, the one species occurring also in the Orient and Australia; three species of the mainly African-Oriental Parena, one of the species occurring also in the Orient, one also in Australia, and one in the Orient and Aus- tralia; and 59 known endemic species of the otherwise mainly Australian Demetrida (all perhaps derived from one Australian- related stock) with one of the New Guinean species found also on the tip, of Cape York. In more detail, the New Guinean Lehia include karenia, which is also Oriental; melanonota (present part, Tax. siippl. ) which is Oriental and Aus- tralian; papucUa, with a close relative in Australia; and additional species of which the relationships are undeteiTnined or are with other New Guinean species. And of DoUchoctis, striata ranges from southern Asia to New Guinea and Australia; rnicro- dera, from Sumatra to New Guinea; and the remaining 11 New Guinean species of the genus, all members of or perhaps de- rived from the aculeata group, include one species (actdeata) that occurs also in the Oriental area (Celebes) and reaches mid- peninsular Cape York, and at least one additional Oriental relationship. The sum of geographic units of the foliage-living New Guinean Lebiini is nine species that occur also in the Orient plus four Oriental relationships, and five species that occur also in Australia plus three Australian re- lationships and three additional occur- rences on Cape York. Besides the Lebiini listed in preceding paragraphs, four lebiine genera of which I do not know the habitats occur in New Guinea. These genera are probably arboreal, but I do not know whether (in New Guinea) they live on tree trunks or in foliage. Agonochila is a primarily Aus- tralian genus with seven small endemic species in New Guinea representing per- haps only one Australian-related stock; most members of this genus in Australia live on tree trunks, but a few small tropical Australian species inhabit foliage, and the New Guinean ones may do so too. Minii- phloeus is a monotypic genus confined to New Guinea; its relationships and habitat are unknown. Phloeocarahus is primarily Australian but is represented on New Guinea by one species that occurs also in Australia and one that is endemic and represents a separate Australian-related unit. And Trigonothops is primarily Aus- tralian but includes one New Guinean species closely related to an Australian species. The geographic units of these four genera total one species shared \\'ith Aus- tralia plus tliree Australian relationships. Finally, as far as Lebiini are concerned, two genera are represented in New Guinea by single species that are carried by man and that cannot be placed in the preceding habitat classification. They are Anchista and Endynomena, each represented in New Guinea by one species that has been found also in the Orient but not in Australia. (Plochioniis pallens, if it turns up in New Guinea, will be a third man-distributed lebiine in New Guinea.) 196 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 Table 7. Summary of geographic relationships of New Guineax Lebiini Habitat Species also Or. Additional Or. units Species also Au. Additional An. units Additional to C. York Ground 3 4 2 2 — Trunks 8 6 1 4 3 Foliage 9 4 5 3 3 Prob. arb. — — 1 3 - Man-carried 2 - — - - Total 22 14 9 12 6 Table 7 sums up the geographic relation- ships of New Guinean Lebiini. The distribution and relationships of Lebiini from Asia to Austraha may be summarized as follows. The tribe is rela- tively poorly represented (and chiefly terrestrial) in temperate Asia. It is better represented (and more arboreal) in trop- ical Asia, with some small genera confined to the tropical Asiatic mainland and others, including larger genera, extending onto the Malay Archipelago for various distances. The lebiines of the Western (Oriental) part of the archipelago are numerous and include about a dozen (mostly small) genera that reach Java and/or Borneo but do not cross Wallace's Line to Celebes, and also two important genera that do extend to Celebes but not New Guinea. The Lebiini of New Guinea are numerous too, and include 22 species that occur also in the Oriental area plus at least 14 Ori- ental relationships, nine species that occur also in Australia plus at least 12 Australian relationships and six additional species that reach Cape York. Some New Guinean genera are represented also in Australia, but additional Australian genera are few, the important ones being mostly tree-trunk- living or terrestrial. This whole main pattern of distribution from Asia to Australia reflects the fact that Lebiini are primarily arboreal Caiabidae, far more numerous in the complex arboreal habitats of tropical rain forest than they are in thinner and less complex temperate woodlands. The localization of a number of small genera toward the Oriental end of the Malay Archipelago and the extensive radi- ation in several genera in New Guinea ( see below) suggests that, although some species have been able to disperse from southern Asia to Australia, some other lebiines have dispersed less effectively than might have been expected of such active, winged Carabidae. Their dispersal may have been limited either by limitation of their power of dispersal or by discontinuities in the distribution of the rain forests in which most of them live. It is noteworthy that the lebiines that have radiated in New Guinea have diverse geographic relation- ships: the foliage-living Demetrido (59 species on New Guinea derived from one or a few ancestors! ) is Australian-related; the foliage-living DoJichoctis and tree- trunk Catascopiis ( in both of which moder- ate radiations have occurred in New Guinea) are primarily Oriental in relation- ships; and the foliage-living Dicraspeda *" (six New Guinean species) and tree-trunk Mimithodes (nine New Guinean species) are primarily New Guinean. The only ground-living lebiine genus in which radi- ation seems to have occurred on New Guinea is Anomotarus (eight New Guinean species, seven of them perhaps products of local radiation); the primary geographic relationship of the New Guinean members of this genus is undetermined. [49] The small tribe PENTAGONICINI" (Part III, p. 191) includes only four genera, of which one is confined to New Zealand, the other three being represented in New Guinea. Pentagonica occurs in all The carabid beetles of New Guinea • Darlington 197 the warmer regions of the world and in- ckides six species on New Guinea: two of them apparently range from southern Asia to Australia; two more, from southern Asia to New Guinea and Cape York; and two are endemic. The one known species of Paroscopodes occurs in eastern New Guinea and northern Australia. And Scopodes is divisible into two stocks: one stock is mainly Australian but includes one species at very high altitudes on the Snow Mountains of New Guinea and one on high mountains in Java; and the other stock is confined to New Guinea and includes seven species which are perhaps all prod- ucts of one (Australian-related) radiation. Pentagonicines are small Carabidae. The New Guinean species of Penfa^onica are arboreal, in foliage; of Parascopodes, ground- or grass-living; and of Scopodes, ground- or log-living, [50] The small, African-Oriental tribe HEXAGONIINI (Part III, p. 202; present part, Tax. siippl.) includes two genera in the Orient: Dinopelma (about 11 species) is apparently confined to the western part of the Malay Archipelago, with several species on Java, Borneo, and the Philip- pines but none on Celebes; Hexa^onia (which occurs also in Africa) is rather diverse in the Orient; two Oriental stocks are represented in New Guinea by single endemic species; and one of these stocks has one endemic species in Australia, too. The members of this tribe that I have collected are rather small, winged Carabi- dae which live under the leaf sheaths of tall grasses. [51] Tlie tribe ODACANTHINI (Part III, p. 203), which is worldwide in dis- tribution, includes small and medium- sized, winged carabids of which some are terrestrial and some arboreal, some of the subarboreal forms being also hydrophilous. Few occur in temperate Eurasia. The Orient possesses many, diverse species of the worldwide genus CoUiurls ( see below ) and representatives of a few smaller genera (some named below). New Guinea has eight genera, 19 species, rather diverse in ecology and in geographic relationships (see below). And Australia has several small endemic genera as well as representa- ti\es of several of the genera that occur on New Guinea. New Guinean Odacanthini are as fol- lows. The genus CoJUuris (see above) in- cludes four species in New Guinea, of which one occurs also in the Orient, one is endemic but related to an Australian spe- cies (it and the preceding probably live in grass especially in wet places), and two are endemic but probably represent one Oriental stock, and one of these species occurs also on the tip of Cape York (these species live on or near the ground in wet places). Of the Oriental-Australian genus Casnoidea, only two species surely occur in New Guinea: one is endemic, and one occurs also in Australia (the members of this genus live in grass and reeds over water). The one knowoi species of Basisti- cus occurs in northeastern Australia and southern New Guinea (it lives on the ground in more or less open country). Clarencia is an Australian genus with two species on New Guinea: one occurs also in Australia, the other is endemic but Aus- tralian-related (they live on or near the gromid in wet places). Dicraspeda is pri- marily New Guinean, with six rather di- verse species; three geographic stocks can be distinguished; hrunnea ranges from Java and the Philippines to New Guinea and northeastern Australia; longiloba, dnhia, and hispinosa are endemic and perhaps represent one stock which is confined to New Guinea except that longiloho reaches New Britain and dnhia Cape York; and quadrispinosa and violacea are related to each other (and probably derived from the same stock as the preceding) and are confined to New Guinea except that one or both reach the Moluccas, New Britain, and/or the Solomons. (All species of this genus li\e in under-story foliage of rain forest.) One Oriental species of the Afri- can-Oriental genus Lachnothorax reaches 198 Bulletin Musetim of Comparative Zoology, Vol. 142, No. 2 New Guinea but not Australia (I think it lives beside running water). One New Guinean speeies is tentatively assigned to the Australian genus Eiidalia but may also have Oriental relationships (this is prob- ably water-loving too). And Dohodiim is a monotypic genus confined to New Guinea but perhaps derived from the Australian Eudalia (it lives among stones beside turbulent brooks). The sum of geographic units of the New Guinean Odacanthini is three species that occur also in the Oriental area plus one Oriental relationship, and four species that occur also in Australia plus foiur Australian relationships, and two additional species reaching Cape York. [52] The DRYPTINI (Part III, p. 216) is a small tribe of medium-sized, usually winged Carabidae which usually live on the ground or in grass, I think. Of this tribe, the widely distributed, terrestrial genus Galeritiila (Reichardt, 1967) crosses Wallace's Line to Celebes; the Old-World (and Brazilian) genus Dnjpta is repre- sented in New Guinea by two endemic species probably related to different Ori- ental forms, by one Australian species which reaches only the southern edge of New Guinea, and by one additional en- demic species; and the Old-World genus Desera includes one New Guinean species which is endemic (except that it reaches also New Britain and New Ireland) but closely related to species in both the Orient and Australia. Besides these, the tribe contains only a few small genera none of which occurs in the area under consider- ation except that Pseiidaptinus (Thalpius), a genus of small terrestrial dryptines widely distributed in the Americas, has also one species in Australia. [53] Of the small tribe ZUPHIINI ( Part III, p. 218; present part. Tax. siippl.) which includes small, mesophile and hvdro- phile carabids, New Guinea possesses one endemic species of the worldwide genus Zuphium and one Australian species of the same genus that reaches only southern New Guinea; four species of the African- Oriental genus of Planetes, one being also Oriental and one also Australian; and two endemic species of Colmidia representing one Oriental stock. The few other (small) genera of the tribe include Oriental A<:,astiis, which reaches Java, Borneo, and the Philippines, and Acrogemi.s, confined to Australia. [54] Of the small, Oriental-Australian tribe HELLUODINI (Part III, p. 222), which includes only one principal genus of medium-sized, winged, mesophile carabids. New Guinea possesses nine species of Fo'^ono antensis ^ Dolidwctis striata -- ^ Demetrida angulata _.__ — x Pentagonica pallipes ^ II blanda x -- x II erichsoni x CoUitiris par x Clarcncia quadriguttata ^ Dicraspeda brunnea x x. x II dubia — - X collecting on Cape York, and on previously published records. Further collecting would probably fill some gaps, but never- theless carabid distributions evidently are irregular in these rain forests. Of the 24 New Guinean species tabulated, six have been found only on the tip of the peninsula, but only six of the other species have been found there, leaving 12 species that occur in more-southeni rain-forest tracts but are apparently absent on the tip of Cape York. The tip-of-peninsular rain forest is small and of comparatively poor quality, and the carabid fauna probably really is much more limited than are the faunas of the larger and better rani forests farther south. Sixteen of the 18 species that do occur farther south have been found in the mid- peninsular rain forest. And only ten have been found in the base-of-peninsular forests (but see below). The detailed pattern shown by Table 9 should be supplemented by two general statements. First, a few of the species tabulated, including Viohfionum viola- ceum, extend still farther south in Austra- lia. And second, a considerable number of endemic species derived from New Guinean groups exist in the base-of- peninsular rain forest, and in some cases still farther south in Australia. (This whole situation, of somewhat irregular occurrence of New Guinean species in the isolated rain forests of Cape York, and existence of additional derived species farther south, suggests continual dispersal of rain-forest j species from New Guinea to Australia. The insects probably disperse with diffi- culty from forest tract to forest tract. Some species probably survive only temporarily in some tracts. Survival is probably cor- 1 related with area: extinctions probably occur most often in the smallest tract, which is the tip-of-peninsular one; while survival time in the largest tract, at the base of the peninsula, has often been long The carabid beetles of New Guinea • Darlington 207 enough for differentiation of species. All this is consistent with a history of con- tinuing dispersal from New Guinea into Australia, across ecologic filter-barriers, even when there was a broad land con- nection. Such dispersal across ecologic barriers to a series of islands of rain forest is comparable to dispersal across water gaps to the islands of an archipelago and should be susceptible to analysis by meth- ods developed by MacArthur and Wilson (1967).) To complete this general account of transition of carabid faunas in the Asiatic- Australian transect, I should add that within Australia, between the tropical rain forests of North Queensland and the south- temperate rain forests of southern Austra- lia and Tasmania, there is not only an almost complete change of species and genera but also a second partial change of dominant tribes, from Pterostichini as principal dominant mesophiles to (in the far south) dominance shared by Broscini, Trechini, Licinini, and even "antarctic" Migadopini and Merizodini, as well as some Pterostichini. This change too is described in more detail in my 1961 paper. [64] Summary of tramition,s. In sum- mary of transitions of carabid faunas from north-temperate Asia to south-temperate Australia, there is first a profound change of dominant tribes and genera from the north-temperate zone to the tropics in Asia; then a major transition of tropical faunas from the Orient to Australia, with the most obvious changes at Wallace's Line, between Celebes and New Guinea, and (even in rain forest) between New Guinea and tropical AustraUa; and finally another profound change of dominant tribes and genera from tropical to south-temperate Australia. The carabid faunas in the north and south temperate zones, at opposite ends of this series of transitions, are re- markably similar in certain ways, for ex- ample, in presence of Broscini, of flightless "Trechiis," and of Bemhidion. These groups must somehow have crossed the tropics in the past. However, they do not occur in New Guinea now, and further consider- ation of them would be out of place here. I have discussed them in more detail else- where (1965). The gross changes in taxonomic com- position of carabid faunas within the trop- ics, from Java to New Guinea to tropical Austraha (North Queensland), are shown by histograms in Figure 13.^ The histo- gram of the Javan fauna is based on a list extracted from my MS list of Indo- Australian Carabidae [4]. That for the New Guinean fauna is, of course, based on counts of species listed on my data sheets [16]. And that for the tropical Australian (North Queensland) fauna is based on a list extracted from my manuscript list of Australian Carabidae [4]. Many species described from "Queensland" are not known from more exact localities and may not be tropical, but on the other hand I have a number of tropical Queensland spe- ^ The carabid faunas of Celebes and the Moluc- cas are too little known to be included in this comparison. Celebes is about half again larger than Java, but only a]:)out one-third as many Carabidae (only about 150 species) have been recorded from it. (Professor E. O. Wilson calcu- lates, using data provided by me, that if Celebes were as well collected as Java and New Guinea, about 509 species of Carabidae should be known from the island. ) The Moluccas are much smaller, but tlie number of species that occur there is presumably increased by differentiation of species on different islands of the group. About 100 species of Carabidae have actually been found there ( including both those recorded in print and those found by myself on Morotai Island), but this is probably a minor fraction of the whole Moluccan carabid fauna. The following table shows the numbers of species in four principal tribes of Carabidae actually known from Celebes and the Moluccas. The figures suggest that Agonini and Lebiini are the dominant tribes on these islands as they are on New Guinea, but the figures should be considered preliminary indi- cations only. Celebes Moluccas Pterostichini 8 13 Agonini 23 18 Harpalini 20 8 Lebiini 36 35 208 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 < 2 o <0 CD .8 V/// yiHio iN,iNoan3Sd iNiNiHOvya INI,NVOVaO iNiia3"i INHVddVH INIIN3V1H0 ININOOtf iN.isoaaid iNiiaiaiN39 iNiiiyvos iNissnvd iN,woan3Sd iNiNiHovya INl.NVOVaO iNiiaan mnvdbivH lNIIN3VnH0 IN1N09V IN.lSOySld iNiiaia^3a iNiiiavos iNissnvd sq; < _J > < 1- < 2 UJ -^ o o c: c ./> ° .2 o that their distribution patterns really do reflect dispersals rather than local radi- ations, extinctions, and other complications. Carabidae are primarily winged insects vvhich do fly and can disperse rapidly, in ^pite of the fact that some have lost the power of flight. Their movements over the world and between adjacent continents have probably been very numerous and very complex. And, to judge from what has happened among other animals of which we have better fossil records (espe- cially the mammals), the multiple and successive dispersals of dominant carabid groups have probably been accompanied by frequent and widespread extinctions of ather groups. Therefore, I do not trust numbers clues" or "area clues" to show directions of movement of nondominant, discontinuously distributed groups, of vvhich the present distributions may be the result of withdrawals ( partial extinctions ) rather than of initial dispersals (cf. Lox- indrus, [82]). (Zoogeographic tracking is like tracking in snow in that the tracker 3an follow with the most confidence the clearest, most recent trails rather than )lder, partly obliterated ones.) With these criteria and their limitations n mind, I want now to survey the tribes )f Carabidae that are represented in New Guinea and to try to pick out the groups "hat clearlv show evidences of direction of dispersal. Evidence might come from the distributions of tribes as wholes, or of genera, or of species. In any single case, no matter how clear the evidence seems to be, the conclusions should be considered tentative. But if, of many separate cases, most seem to show dispersal in the same direction, the probability that dispersal has had a net direction will become strong. However, no matter how great the pre- ponderance of movements in one direction, some counter-movements are to be ex- pected too; this seems always to be the case in complex faunal movements. The groups of Carabidae that seem to show directions of dispersal are listed in Table 14. The table is derived mostly from the preceding survey of tribes (items [33- 57] ) . Most of the details that seem to me to indicate direction have been given in this survey (or under the groups con- cerned in Parts I-III and the present Taxonomic supplement) and will usually not be repeated here. Evidence of di- rection of movement may, of course, be derived not only from the distribution of a given genus or species but also from the occurrence of related fomis — whether they are Oriental or Australian. Tlae following abbreviations are used: Or, Oriental area; NG, New Guinea; Au, Australia beyond Cape York; CY Cape York; Mol, Moluccas. "Or to NG to Au" means that dispersal has apparently been from the Oriental area to New Guinea to AustraHa. In this table I have indicated movements from New Guinea to Cape York or from New- Guinea to the Moluccas only when they are con- tinuations of longer moxements from the Orient or Australia respectively. To in- clude other short-range movements would weight the results in favor of movements to\\'ard Australia, because the Carabidae of Cape York are better known than those of the Moluccas. [75] Summary of direetion to this point. The 128 groups (tribes, genera, and spe- cies) of Carabidae listed in Table 14, that seem to show relatively clear evidences of 222 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 Table 14. Groups of New Guinean Carabidae that seem to show directions of dispersal In tribe Ozaenini: Pseudozaena orientalis with siibsp. opaca. Or to NG. In tribe Paussini: Arthropteriis, An to NG. In tribe Scaritini: Geoscaptus cacus, An to NG; Syleter, Or to NG to CY; Clivina zebi. Or to NG to CY. ( Other Clivina in New Guinea are in part Austrahan- and in part Oriental-related, but I think they are not w ell enough known to justify deductions about their directions of dispersal. ) In tribe Bembidiini: Tachijs as whole, mostly Or to NG to Au; T. fasciattis. Or to NG to An; T. cetjlanicus. Or to NG; T. lilugi, Or to NG; T. convexus, Au to NG; T. fumicaius. Or to NG (closely related curticollis may not have moved Au to NG but may represent a first invasion from Or to NG to Au of an Or stock that later reinvaded NG as fiimicatus); T. umhrosus. Or to NG; T. coracinus. Or to NG; (some other Tachijs have distributions that suggest dispersal either from Or to NG or from Au to NG, but the evidence seems less clear than in the cases cited); Limnastus atricapillus. Or to NG. In tribe Trechini: Perileptus, Or to NG to Au. In tribe Panagaeini: Peronomerns xanthopus. Or to NG; Dischissus notulatiis. Or to NG; Craspedo- phorus gressittorum, Au to NG. In tribe Pterostichini: Morion of Or group, Or to NG to Au; (Mecyclothorax, see [80]); Brachi- dius crassicornis. Or to NG; Caelostomus (excluding picipes). Or to NG to GY; Ahacetus haplostermis. Or to NG to An; A. convexiusculiis. Or to NG; Lesticus, Or to NG to An; Prosopogmus, Au to NG to Mol; Platycoelus, Au to NG to Mol; Loxandrus, Au to NG to Or (Celebes) (see [82]). In tribe Agonini: tribe as a whole, mainly Or to NG to An; Euplenes, Or to NG; Dicranoncus ■ queenslandicus. Or to NG to Au; Lorostemma, Or to NG; Agomim (Sericoda) cetjlaniciim. Or to NG [ (see [80]); Notagonum dentellum, NG to An; N. suhmetaUiciim, Au to NG; Violagomim violaceum, NG to Au; Colpodes sapphyrinus with subsp. sloanei. Or to NG; C. habilis, NG to Au; (directions of. dispersal of some other Agonini are indicated, but the evidence seems less clear tlian in the cases cited ) . In tribe Perigonini: Perigona as whole. Or to NG to An; P. plagiata. Or to NG. In tribe Licinini: Bodister sundaiciis. Or to NG to Au; PhysolaestJins, Au to NG to Or (Java, Philip- pines); Dichrochile, Au to NG; (Microferonia, see [80]). In tribe Chlaeniini: Chlaenius ceylanicus. Or to NG to Au; C. tnacuUger, NG to Au; C. gtittula. Or to NG; C amplipennis. Or to NG; C. himacidatus group. Or to NG; (directions of dispersal of. some other Chlaenius are indicated, but the evidence is less clear than in the cases cited ) . j In the tribe Harpalini: Gnathaphaniis as whole, Au to NG to Or (SE Asia); G. licinoides, Au to; NG; G. upolensis, Au to NG to Or (to Malay Pen.); G. picipes, Au to NG; G. pulcher, Au to NG; Dia- phorurnerits, 2 stocks, An to NG to Mol, and Au to NG; Hypharpax, Au to NG to Or (Java, Sumatra); Lecanomerus, Au to NG; Chydaeus, Or to NG (see [80]); Platymetopus, Or to NG; Trichotichnus, 3 stocks. Or to NG; Coleolistis, Or to NG to CY; Egadroma, 3 stocks Or to NG to Au, and 1 stock Or to NG; Stenolophtis, 2 stocks Or to NG to An; Acupalpus, 3 stocks Or to NG to Au. In tribe Cyclosomini: Anaulacus siamensis. Or to NG; Caphora litimilis. Or to NG to CY; Saro-\ throcrepis, Au to NG to Or (Java, Philippines). In tribe Lebiini: Aristolebia, Or to NG to CY; Physodera, Or to NG; Lebia, Or to NG to Au; Holcoderus, Or to NG to Au; Minuthodes, NG to Au, NG to Or (Celebes); Catascopiis as whole. Or to NG to Au; C. facialis. Or to NG; C. elegans. Or to NG to Au; C. smaragdulus. Or to NG to CY; Pericalus, Or to NG; Coptodera as whole. Or to NG to Au; C. eluia. Or to NG; Agonochila, Au to NG; Mochtherus, Or to NG; Dolichoctis as whole. Or to NG to An; D. striata. Or to NG to An; D. aculcata, NG to Or (Celebes); Stricklandia, NG to An; PeUocypas, Or to NG; Syntorntis, Or to NG to An; Apris-i tus. Or to NG; (I have not counted Anchista binotata dnd Eiulynomena pradieri, hoihT^rohahXy curried by man from Or to NG); Demetrida, Au to NG to Mol (but see [80]); Phloeocarabus, Au to NG; Trigonothops, Au to NG; Nototarus, Au to NG. The carabid beetles of New Guinea • Darlington 223 Table 14. Continued In tribe Pentagonicini: Pentagonica as whole, Or to NG to Au; P. pallipes, Or to NG to CY; P. hlanda. Or to NG to Au; P. crichsoni, Or to NG to CY; P. rtificoUis, Or to NG to Au; (Scopodes see p. 228). In tribe Hexagoniini: Hexagonia, Or to NG to Au. In tribe Odacanthini: CoIIiuris, Or to NG to Au; C. fuscipennis. Or to NG; Clarencia, Au to NG; Dicraspeda, NG to Au, NG to Or ( Java, Philippines ) ; Lachnothorax, Or to NG. In tribe Dryptini: Dnjpta mastersi, Au to NG. In tribe Zuphiini: Planetes, Or to NG to Au. In tribe Helluodini: Pogonoglossits, NG to Au. In tribe Helluonini: Creagnis, Or to NG to Au; HeJhiosoma, Au to NG; Helliiodema, Au to NG; Gigadema, Au to NG. In tribe Brachinini: Pheropsophus, Or to NG to Au; Brachinus, Or to NG. In tribe Pseudomorphini: Adelotopus, 1 stock Au to NG to Or (Java) and 2 stocks Au to NG; Sphcdlomorpha, 3 stocks Au to NG; (Cryptocephalomorpha occurs from NG to the Malay Pen. and Thailand, but its direction of dispersal is not clear). directions of dispersal, are classified and totaled in Table 15. The grand totals are 89 groups that seem to have dispersed southeastward, including 44 groups that seem to have spread all the way from some part of the Oriental area to some part of Australia (including Cape York), and 39 groups that seem to have dispersed north- westward, including seven groups that seem to have spread all the way from Australia to some part of the Oriental area. It should be repeated and stressed that the groups considered to show clear evi- dences of directions of dispersal have, necessarily, been selected somewhat arbi- trarily. The totals would vary to some extent with judgments about which groups really show direction clearly. But on any reason- able basis of selection the tribes, genera, and species that seem to have dispersed from the Oriental area to New Guinea and Australia far outnumber those that seem to have dispersed from Australia to New Guinea and the Orient. This disparity is greatest among the groups that have moved the longest distances. Many pri- marily Asiatic or African-Asiatic genera include species that extend eastward across the Malay Archipelago to the mainland of Australia. But very few primarily Austra- lian groups include species that extend across the islands to the mainland of Asia; in fact Gnathaphanus, with one common Australian species reaching the Malay Peninsula and another reaching India etc., is the only carabid genus that seems to show this reverse patteni clearly. (Tlie Table 15. Summary of apparent directional DISPERSALS OF NeW GuINEAN CaRABIDAE Or to NG 37 Or to NG to CY 9 Or to NG to Au 35 NG to Au 8 SE ■ movements 89 Au to NG 25 Au to NG to Moluccas 4 Au to NG to Or 7 only to Celebes, 1 ( Loxandrus ) to Java, 1 (Adelofopus) to Java and Philippines, 2 (Physolaesthus, Sawthrocrepis) to Sumatra, 1 (Hypharpax) to Malay Pen., 1 (G. upolensis) to India, etc., 1 (Gnaihaphanus) NG to Or 3 only to Celebes, 2 {Minuthodes, Dolichoctis aculeata group) to Java, 1 ( Dicraspeda ) NW movements 39 224 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 mainly Australian tribe Pseudomorphini reaches the southeastern corner of Asia, but the only genus that does so (Crypto- cephalomorpha) is not Australian and its geographic history is doubtful. ) This whole situation suggests not only that move- ments from Asia toward Australia are much more numerous than the reverse, but also that the reverse movements are usually shorter. [76] Direction and doniinance. Thus far, I have been considering directions of dis- persal in the whole carabid fauna without distinguishing different fractions of it, ex- cept that I have compared certain relatively vagile fonns (especially very small ones) with the rest of the fauna. Now, I want to compare the apparent directions of dis- persal of different faunal fractions to see if the comparisons will give further in- formation about the origins and history of New Guinean Carabidae. Among the Carabidae of the Asiatic- Australian area, the genera Chlaenius (Part III, pp. 20ff) and Egadroma (Part III, pp. 69ff) are notably dominant. Each genus includes several species that are very widely distributed, ranging from Asia to Australia, and individuals of some of the species are numerous. The distribution patterns of these two genera have note- worthy characteristics in common. In both genera, the relative numbers and diversity of species on different continents and the relative areas occupied seem clearly to indicate origin in the main part of the Old World and multiple dispersals toward and into AustraHa (Chlaenim, into the Americas too ) . Some species in each genus are now widely distributed in Australia as well as in the Orient. Considered singly, these species scarcely show the directions of their dispersals, although the geographic patterns of the genera as wholes do clearly indicate direction. These and most other really dominant Carabidae, for example Tachys fa.sciatus (Part I, p. 414), seem to have dispersed from Asia to Australia. This fact is significant because the most domi- nant members of a fauna are most likely to have dispersed recently and to be dis- persing now, and to show the main direction of dispersal of the whole fauna, if there is a main direction. Several of the most widely distributed and commonest species of these genera, e.g,., Chlaenius jlamguttatus and E<;adro7na smaraiidida, occur in a rather wide variety of wet places, although some other species of both genera have moved into drier haliitats. These are examples of what I think is a fact, that ground-living carabids associated with water ( hydrophilcs ) dis- perse and also cross climatic barriers more easilv than most other Carabidae (see [84] j. [77] Direction and size. Although small Carabidae disperse more easily than large ones, direction of dispersal is apparently not correlated with size of insects. The very small Carabidae that have reached New Guinea have probably dispersed more rapidly than most larger ones and may show stronger Oriental relationships for this reason [72], but the larger Carabidae too (including for example Lesticiis, Chlaenius, large Colpodes, and Cata- scopu.s) seem to be more Oriental than Australian in derivation. The two largest Carabidae in New Guinea are both Aus- tralian species, but one {Catadromus tene- hroides. Part I, p. 563) is probably man- carried and is probalily not established on the island, and the other ( Gigadenia tnaxil- lare. Part III, p. 234) is known from just the southern edge of New Guinea, and is a unique case and not an integral member of the New Guinean fauna. [78] Direction and it//i.i^.S'. Although winged, flying Carabidae disperse more easily than flightless ones, direction of dis- persal is not strongly correlated with presence or absence of wings in the present case. Almost all New Guinean Carabidae, whatever the present state of their wings, are or may be derived from ancestors that were winged when they reached the island [84, 88], regardless of the direction from The carabid beetles of New Guinea • Darlington 225 which they came. The only sure exception is the fhghtless tiger beetle Triconclyla, which came from the Orient, probably by rafting (cf. [21], footnote). The three other carabids most likely to have reached New Guinea without flying are all Aus- tralian, as would be expected from the fact that New Guinea was connected with Australia not long ago [17]. Tliey are species of Craspedophoriis ( Tax. Stippl. under Panagaeini), Coptocarpus (Tax. siippl., footnote under Oodini), and Noto- tarus (Part III, p. 185). These genera are all represented at low altitudes in tropical Australia (although the single individuals of Craspcdophonis and Coptocarpus thus far found in New Guinea were in the mountains) and all are now wholly flight- less in Australia as well as in New Guinea. They may have reached New Guinea with- out flying. However, their ancestors were prt)bably winged not long ago and may have dispersed partly by flight. [79] Direction and ecoJogij. Direction of dispersal is correlated with ecology to only a limited extent. The principal eco- logic groups of New Guinean Carabidae (mesophiles, hydrophiles, and arboreal forms) are all more Oriental than Austra- lian in relationships and in probable origins. However, the mesophiles divide into two subgroups: those (more numerous) that live on the floor of rain forest are mainly Oriental; those (fewer) that live in open country including open eucalyptus woodland are mainly Australian. The latter, the mesophiles that live in relatively open, relatively dry areas in New Guinea, are apparently the only ecologic group of New Guinean Carabidae in which Australian relationships and probable origins do pre- dominate. Because my ecologic division of the New Guinean fauna is rough at best, with many details in doubt, I see no point in attempting to find finer correla- tions between ecology and geographic origins. [SO] Direction and oltitude: mountain- hopping, across the Malay Archipelago. Direction of dispersal is not clearly cor- related with altitude. Most Garabidae on the high mountains of New Guinea seem to have been derived from lowland forms on the island and not to have had inde- pendent geographic origins [90]. Of the few endemic high-mountain genera that do seem to have independent geographic relationships, AnaJoma and Idiagomim probably have Oriental and Rhytiferonia Australian ties, but their origins are far from clear. Less differentiated mountain- hopping carabids that have reached New Guinea are considered in more detail be- low. Doubtful cases which should be dis- posed of first include Notagonum suh- metaUicum (see Tax. siippL), a common, winged, southern Australian carabid which has been found at moderate altitudes both in tropical Queensland and in New Guinea; it may have begun to mountain-hop north- ward and westward, but if so, it has not gone far. Craspedophoriis, Physolaesthus, and Coptocarpus, although thus far found only at middle altitudes in New Guinea, occur at low altitudes in tropical Australia and are probably not mountain-hoppers. The three genera MecyclotJiorax (Part I, pp. 498, 505; present part. Tax. suppl.), Mia'oferonia (Part III, p. 18), and Sco- podes (Part III, p. 197) have mutually similar distributions in the area mider discussion. All these genera are now chiefly Australian, but all have also en- demic species localized on mountains in New Guinea and in Java. (The distri- bution of MecycIotJwrax is complicated by occurrence also on the Hawaiian Islands, etc., and of Scopodes by occurrence on New Guinea not only of one Australian- related species but of a second endemic stock which has radiated on the island, but these are added complications which do not affect the Australia-New Guinea- Java pattern. ) The question is, have these genera mountain-hopped from Australia across New Guinea (and presumably Celebes ) to Java, or are the isolated species on New Guinea and Java relicts left by 226 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 withdrawals ( by partial extinctions ) of the Guinean species also well defined and genera into Australia? I cannot answer this flightless, but perhaps less closely related question, but at least in Mecijclothonix and to any particular Australian species (but Scopode.'i the New Guinean and Javan the species of this genus need further species seem fairly closely related to Aus- study), may be oldest of all. Tliis classi- tralian species, and I think westward fication by age may be wrong in detail, mountain-hopping is a possible explanation. However, it does seem probable that All the pertinent New Guinean and Javan Mecijclothorax, Scopodes, and Microfewnia species of these genera are now flightless, dispersed in the Malay Archipelago before but all three genera include winged species Sericoda and Clujdaeus did, and it is at in Australia, and all may have dispersed least possible that they represent stages by flight. in dispersal by a "sweepstakes" route A reverse pattern of distribution is across the mountains of the Archipelago shown by Sericoda (present part. Tax. from Asia to Australia. No single group suppl.) and Clujdaeus (Part III, p. 47), of mountain-hopping Carabidae occurs both of which are distributed as if they along this whole route now. However, have mountain-hopped from Asia across Sericoda and Clujdaeus have covered most the Malay Archipelago to New Guinea, of the route (and an Asiatic stock of In these cases direction of dispersal is Bemhidion, not related to the Australian clearer. An actual Oriental species of Bemhidion, has reached the mountains of Sericoda reaches New Guinea and is still Celebes — Darlington, 1959a), and Mecijclo- winged. And Clujdaeus includes slightly thorax, Microferonia, and Scopodes may differentiated species widely scattered on have follo\\'ed the whole route earlier, mountains on the Malay islands, some mountain-hopping from Asia to New flightless, but others still retaining wings Guinea and Australia, becoming extinct in (for example, C. hakeri Andrewes on Asia, and radiating in Australia especially mountains in Luzon is still dimoqohically south of the tropics. If so, the Javan and winged). New Guinean species of these genera may The five cases of probable or possible be geographic relicts, and the special New mountain-hopping summarized in the two Guinean group of Scopodes (Part III, p. preceding paragraphs can be arranged ac- 197, Notes under Scopodes) may represent' cording to apparent ages of dispersal, a separate radiation of the ancestral stock relative age being judged by state of wings which has paralleled the Australian radi- and by amount of differentiation of the ation rather than being derived from it. isolated species. Sericoda, with a winged This hypothetical history is diagrammed in species now extending from Asia to New Figure 14. Guinea, has presumably dispersed most I do not know how these five genera recently. Clujdaeus, with slightly differ- really have dispersed. But I think the entiated species scattered from the Hima- best way of attempting to decipher their layas to the Philippines and New Guinea, histories is to put the cases together, see and with wings still present in some insular whether they fit a common pattern ( as forms, is presumably a little older. Mecijclo- they seem to do), and see how that pattern thorax and Scopodes, each with well- compares with the distributions and ap- diffcrentiated, flightless species on Java parent histories of other Asiatic- Australian' and New Guinea, but with the Javan and Carabidae and of other animals and even New Guinean species apparently related of plants. to each other and related to specific Aus- Among New Guinean Carabidae, Deme- tralian species, are presumably still older, trida (Part III, pp. 140ff) may fit the pat- And Microferonia, with Javan and New tern suggested for Scopodes. The ancestor The carabid beetles of New Guinea • Darlington 227 6 < - 5 4 3 2 I -(N.AMER.)- ASIA JAVA N. TEMP- ERATE T ■> ? . ? ■ ■ ? - 7 CELEBES R 0 ^ ^ ->^ NEW GUINEA c AUSTRALIA I |S. TEMP- I ERATE Figure 14. (Hypothetical) diagram of dispersal of successive mountain-hopping Carabidae from Asia to Australia. Broken lines indicate presumed past occurrences; solid lines, present occurrences. Case 1 represents Psydrini (included in Ptero- stichini in text) other than Mecyc/ofhorax, now relict in North America and moderately diverse in Australia; 2, Mecyclotho- rax, now represented on mountains in Java and New Guinea and moderately diverse in south temperate Australia (and well represented in Hawaii, etc.); 3, Scopodes, now on mountains in Java and New Guinea with separate radiations in New Guinea and Australia; 4, Chydaeus, with slightly differentiated species on mountains from Asia to New Guinea but not Aus- tralia; 5, Agonum (Ser/codo) cey/onicum (Motschulsky); 6, 6emb/d/on of the bryanti-pendelburyi group (Darlington, 1959a), which has reached Java and Celebes but not New Guinea. See text for further details. of Deiiietiida must have reached the Aiis- trahan Region from some other part of the world, probably from Asia, since the genus does not have a subantarctic distribution. The New Guinean radiation of the genus (Part III, pp. 143-144), like that of Scopodes, may be a separate radiation of an original Asia-derived ancestor independent of the main Australian radiation. (However, I have scored Demctrida as probably halv- ing dispersed from Australia to New Guinea, see [74], Table 14.) A pattern of distribution that seems basically similar is shown by the native murid rodents of the Australian Region, among some of which radiation has pro- ceeded partly independently in New Guinea and in Australia. The subfamily Hydromyinae, for example, has diversified principally in New Guinea. In this case we know that the ancestral stock did come from Asia, and we can be reasonably sure that the ancestor of the New Guinean Hvdromvinae was Asia-derived and did not come from Australia even by counter- movement. Another basically similar pattern is shown by a geographically notable genus of plants: trees of the genus Nothofagus (southern beeches) are well represented on the mountains of New Guinea as well as in south-temperate Australia, New Zea- land, and South America. We know (from their pollen record) that the history of these trees in the southern hemisphere has been complex, but we can be reasonably sure that their ancestor came from the north. The numerous species on the mountains of New Guinea may therefore be products of radiation of an ancestor derived from Asia, not from Australia. (For further details and discussion see Darlington, 1965: 29-31 and other pages listed in index.) In summary of Carabidae that seem to have mountain-hopped for considerable distances in the Malay Archipelago, Seri- coda and Clnjdaeus (and Bemhidion, to 228 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 Celebes) have elearly dispersed southeast- ward, from Asia, relatively recently; and Mecijdothorax, Microferoniu, and Scopodes may have dispersed northwestward, from Australia, relatively long ago. However, all these genera may fit a common pattern of successive dispersals southeastward, from Asia toward Australia, with ancestral stocks later becoming extinct in Asia, and with separate evolutionary radiations some- times (in Scopodes) occurring in New Guinea as well as in Australia. Some other Carabidae (notably Demetrida) may fit (a terminal stage of) this pattern. And some rodents (Hydromyinae) and some plants (notably Nothofagtis) may have followed the same pattern of southeastward dispersal, extinction in Asia, and radiation in separate centers in New Guinea and Australia. [81] Direction and age. Whether di- rection of dispersal is correlated with age (time of arrival) of different groups of Carabidae in New Guinea is an important question. It is conceivable that early arrivers might show different patterns of relationships and origins than later ar- rivers do, and the differences might indi- cate changes in the geographic or ecologic relations of New Guinea to the Orient and Australia. In the absence of a fossil record, age cannot be determined exactly, but it is at least a good working assumption that genera endemic to New Guinea (Table 13) are relatively old (but see [89]). Endemic genera of which the ancestors seem to have come from the Orient are Perileptodes, Analoma, Lyter, Odonto- masoreu.s\ Tarsagonum, Idiagonum, and additional chiefly mountain-living agonine genera, and those of which the ancestors seem to have come from Australia are Rhytiferonia, IleUuonidius plus Ilelluo- papua (one stock), and three endemic genera (perhaps only one stock) related to and perhaps derived from Loxandriis. (The relationships of Tiferonia and Minii- phloeus are doubtful.) Among these pre- sumably relatively older arrivers, therefore. Oriental stocks seem to be at least twice as numerous as Australian stocks, and the evidences of direction are clearer in the case of some of the Oriental stocks than of the Australian stocks, I think. The carabid stocks that have radiated on New Guinea may also be relatively old. The radiation of agonines derived from Notagonum- and Colpodes-\ike ancestors is unique on the island. Their ancestors were Oriental. The uni(|ueness of this case lies not only in the amount of differ- entiation of species and genera at all alti- tudes (Part II) but also in the ecologic radiation (of mesophiles, hydrophiles, and even a few arboreal forms) that has occur- red within the limits of New Guinea. A less striking radiation has occurred in the Pterostichini related to (derived from?) LoxandriLS; this group is Australian in pres- ent relationships and may have been derived from Australia (but see [82]). The radiation of this group has apparently produced two or three endemic genera in New Guinea {Haploferonia, Homalonesi- ota, and Nehrioferonia, if the latter is recognizable) but few species, and its ecologic radiation has been relatively slight: its members are confined to low and middle (not high) altitudes, and the habitats occupied are only those associated with standing water (most Loxandriis), the banks of running water (Nehrioferonia and probably Uomaloncsiota), and the rain-forest floor (probably Haploferonia and Loxandriis latus). Tlie most striking multiplication of spe- cies on New Guinea has occurred in the genus Demetrida, which may be Australian in origin. Most members of this genus in New Guinea apparently live in the foliage of rain forest. Other species-radiations of Carabidae in rain-forest foliage have oc- curred in Dolichoctis of the aculeata and' related groups, which may be derived from one Oriental ancestor, and in Dicraspeda, which is a mainly New Guinean genus ( six rather diverse species on the island, all in lower-story rain-forest foliage) of unde- The carabid beetles of New Guinea • Darlington 229 termined ancestry. Among the Carabidae differentiated species on Celebes. This that Hve on tree tiamks in rain forest, pattern suggests recent dispersal from moderate species-radiations have occurred Australia to New Guinea to Celebes. But in Catascopus of the icallocei and perhaps Loxandnis is well represented also in the other groups (Oriental in derivation) and warmer parts of North, Central, and South in Minuthodes, which now occurs chiefly America, and additional genera related to on New Guinea (nine species on the or derived from Loxandnis occur in the island) and of which the ancestor is vm- Americas as well as in Australia and New determined but was probably Oriental Guinea, but nowhere else in the world, as rather than Australian. Other carabid far as I know. The most likely explanation stocks in which moderate radiations of is that Loxandnis once occurred also in species have occurred on New Guinea and the Old- World tropics or at least in tropi- of which the ancestors probably came from cal Asia, that it reached Australia from the Orient include several subgroups of Asia long ago, that it later became extinct Tachijs (especially the serra group), some in Asia, and that still later it made minor Perigona, Trichotichnus, and Pogonoglossus, return movements from Australia to New all primarily ground-living mesophiles. And Guinea to Celebes. others of which the ancestors probably Most species of Loxandnis are hydro- came from Australia include some Clivina philes; some of them are among the most ( hydrophiles ) , Agonochila (arboreal), a aquatic of Carabidae, although some de- special group of Scopodes (mesophiles, or rived or related fonns are more terrestrial, on rotting logs), and Helhionidius (prob- Another subaquatic genus of Carabidae, ably mesophiles). On the whole, the groups the oodine Anatrichis (Part III, p. 31), has in which species-radiations have occurred a distribution that may correspond to that on New Guinea probably include more of an ancestral Loxandnis. Anatrichis oc- Oriental derivatives than Australian de- curs from southeastern Asia (including rivatives, and the Oriental origins are Japan, according to Ueno, personal com- clearer than the Australian ones, I think. munication) to Australia, and is widely Facts and probabilities considered in the distributed also in the warmer parts of the three preceding paragraphs suggest no Americas. Both Loxandnis and Anatrichis obvious correlation between direction of are primarily tropical. They extend into dispersal and age of Carabidae on New moderate temperate areas but do not have Guinea. Some Carabidae have probably the "subantarctic" distribution patterns of reached the island from Australia as well some other Carabidae that may have dis- as from the Orient in both older and more persed across Antarctica (for example, the recent times, but Oriental stocks have Migadopini, Darlington, 1965: 35-37). Tlie probabK' always been more numerous, close relationship of the American and This summary concerns only the direction Australian Loxandnis has been established of dispersal. Possible changes in rate of by modem methods of comparison (Moore, dispersal (numbers of stocks reaching New 1965), but Anatrichis needs study not only Guinea regardless of direction) from time of species now assigned to the genus but to time are considered in [85]. of some other Australian species that may [82] Australian-American discontinuities, prove to be related (Darlington, Part III, A special pattern of distribution is exempli- P- 31, paragraph 3). fied by Loxandnis (Part I, pp. 498, .549- Two other, nonaquatic genera of Carab- 557), which occurs in two widely separated idae should be mentioned in this con- regions. The genus includes numerous nection. The harpaline genus Ne7nagIos.sa species in Austraha, four Austrahan-related or Lecanomerus (Darlington, Part III, p. species on New Guinea, and one slightly 45) is supposed to occur in South America 230 Bulletin Mitseinn of Comparative Zoology, Vol. 142, No. 2 and Australia, but the relationships of the South American and Australian fomis need further study; whatever the earlier history of the group, the small species of Lecano- meriis in New Guinea have presumably been derived from an Australian stock rather recently. And Psetidaptimis {ThaJ- piiis) occurs in the warmer parts of the Americas (numerous species) and in Australia (one species); this genus too needs study. This pattern of Australian-American dis- continuity occurs also in leptodactylid and hylid frogs, chelyid turtles, and especially marsupials. These animals do not have "subantarctic" distributions. All of them, like the Carabidae just discussed (Loxan- clrus, etc.), inhabit principally tropical and warm-temperate areas, and none of them occurs on New Zealand (except as recently introduced by man). The pattern may have been fonned in somewhat dif- ferent ways in different cases, but it is probably usually a relict pattern, the result of widespread extinctions in the main part of the world. [83] Sirmmaiy of direction.'^ of dispersal. New Guinean Carabidae have been found to include 173 Oriental and 120 Australian "geographic units" [58]. This finding (to- gether with the fact that the carabid fauna of New Guinea seems to be mainly deriva- tive, and not a source-fauna from which many groups have radiated geographically [70] ) suggests that Carabidae have moved from the Orient to New Guinea more than from Australia to New Guinea. A com- parison of the distributions of rapidly dis- persing groups of Carabidae with those of more slowly dispersing groups [72] shows relatively strong Oriental relationships among the rapid dispersers, which again suggests movement mainly from the Orient toward Australia. Comparison of the dis- tributions of Carabidae with those of mammals [73] suggests that many cara- bids have moved from the Orient to New Guinea and Australia while the terrestrial mammals were isolated in the Australian Region; a number of carabid species seem to have dispersed from Asia to Australia so recently that populations are not or not much differentiated on the two contin- ents. Selected tribes, genera, and species of Carabidae that seem most clearly to show directions of movement [74, 75] include 89 groups that seem to have dis- persed southeastward and only 39 groups that seem to have dispersed northwest- ward, and the disparity is greatest over the longest distance: 44 carabid stocks seem to have dispersed all the way from the Orient to some part of Australia, while only seven stocks seem to have dispersed from Australia to the Oriental area, and only one primarily Australian genus seems to have reached the mainland of Asia. Before making a final summary, I want to re-emphasize the complexity of the situation, the difficulty of finding and assessing real evidence, and the tentative nature of the conclusions. Faunal move- ments are statistical, not co-ordinated one- way movements. They are the sums of very complex movements and countermove- ments of many families, tribes, genera, and species, which in turn are the sums of almost inconceivably complex movements of individuals. However, in spite of the difficulties and complexities, the evidences of net direction of movement of carabids in the Asiatic-Australian area are surpris- ingly good. The movements have been complex. Large numbers of Carabidae have apparently dispersed from Asia and the Oriental islands to New Guinea and Australia over a long period of time, while the numbers that have apparently dis- persed from Australia to New Guinea and Asia are significantly smaller. Conclusions about direction are more or less tentative in single cases, but when all the cases are put together they form a pattern which (I think) is as a whole overwhelmingly prob- able: dispersal has been predominantly toward the southeast, from Asia toward Australia. At any one point along the route the preponderance of southeastern The carabid beetles of New Guinea • Darlington 231 I ORIENTAL I AREA < I I MOLUCCAS |NEW GUINEA 'CAPE YORK I , I 37 T AUSTRALIA BELOW C. Y. 35 z. 25 V 4 4 7 I Figure 15. Diagram of apparent directional movements o'f Carabidae in the Oriental-Australian area. Lengths of arrows indicate distances moved; widths, numbers of stocks that seem to have made the movements; and numbers of stocks are given in figures on the arrows. See text for further details. against northwestern movements may not have been very great, but over the route as a whole movements from Asia to New Guinea and AustraHa seem to have been several times more numerous than move- ments from Australia and New Guinea toward Asia. The resulting, coniple.x but di- rectional faunal movement is diagrammed in an oversimplified way in Figure 15. I My conclusion is that there has in fact 'oeen direction in the sum of movements ,)f Carabidae in the Asia tic- Australian area: a continual flooding of Asiatic stocks into the Australian region, with much less movement of Australian stocks toward Asia, except for short distances. This process has probably been going on for a very long time, and is still going on. I think that it is only one feature — but a major one — of a complex pattern of directional movements which Carabidae are continu- ally making over the world as a whole, but which (in the absence of a fossil record) are very difficult to demonstrate. The 232 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 apparent world-wide pattern of movements has been briefly described in [67]. One other deduction: if, as I suppose and as apparently must be the case if Carabidae are not to multiply without limit everywhere, additions to carabid faunas tend to be balanced by extinctions, then the movements toward Australia of many new Asiatic stocks must have been accompanied by withdrawals of many other Carabidae, the withdrawals tending to begin in Asia and to progress toward New Guinea and Australia. Some sort of balance between spreading of new groups and withdrawal of old ones would explain several characteristics of the New Guinean carabid fauna. It would explain, for ex- ample, the various cases in which New Guinean Carabidae seem to find their closest relatives in the Philippines; I have not stressed these cases, because most of the groups concerned are not well enough known taxonomically, but possible ex- amples will be found imder Tachijs exiil (Part I, Notes on p. 421), Tifcronia (Part I, Notes on p. 561), Peri<^ona erimae (Part III, Notes on p. 12), and Lehia popiiella ( Part III, Notes on p. 88 ) . Tlie explanation, of course, is or may be that these groups once occurred on the Greater vSunda Islands or even on the continent of Asia, but have become extinct there as competing groups have moved from Asia into the archipelago. Another, more important characteristic of the New Guinean carabid fauna is that it seems to include few or no phylogenetic rehcts [88, 89], and this may be not be- cause New Guinea is a young island but because there has been a relatively rapid overturn of the famia caused by the con- tinual, massive influx of new stocks, chiefly from Asia, with extinction of older stocks. And some kind of balance between arrixal of new stocks and progressive extinction of older ones may also explain the dis- tribution of the "mountain-hoppers" [80], of which new stocks seem to be coming from the Asiatic end of the archipelago while older stocks may be "retreating" to- ward Australia (Fig. 14). [84] Barriers; filtering, at Wallace's Line, Moluccas, and Cape York. During disper- sal across the Malay Archipelago, Carab- idae seem to have encountered principal barriers at Wallace's Line, in the Moluccas, and between New Guinea and Cape York. These have probably been main filter points in the "sweepstakes route" (Simpson, 1940) which many Carabidae have followed, in one direction or the other, between Asia and Australia. These three bankers are of different sorts, and their filtering effects have probably been somewhat different. At Wallace's Line, the barrier has been a gap of salt water — Makassar Strait — which has probably sometimes been narrower than now (perhaps only 25 miles wide in the Pleistocene) but which has separated Celebes from the continental shelf of Asia for a very long time. This is the first major barrier to eastward dispersal of Oriental stocks, and it is a relatively simple but effective barrier. Its effect on Carab- idae has probably been primarily to block more sedentary forms while more vagile forms, especially small, winged, active ones, have often crossed it. Among the Moluccas, too, water gaps have probably existed for a very long time. However, their filtering effect on dispers- ing Carabidae may have been less than the effect of Makassar Strait. Tlie most sedentary, least vagile elements of the carabid faunas of both the Orient and. Australia were probably filtered out by other barriers, so that the Carabidae that reached the Moluccas have been pre- selected for active dispersal. However, the Moluccas are relatively small islands. Their limitation of area has probably limited the numbers of species that could occur, and has probably given an extra advantage to dominant, successful stocks. The islands have therefore probably been a bottleneck through which only Carabidae with some degree of general dominance have been able to pass. (For further consideration of The carabid beetles of New Guinea • Darlington 233 the Caiabidac of the Moluccas see [62]. And for discussion of dominance among Carabidae see [68].) Finally, although New Guinea and Aus- tralia are separated by water now, they were broadly connected by land at times in the Pleistocene. Nevertheless only small fractions of the New Guinean and Austra- lian carabid faunas crossed the land con- nection. The New Guinean rain-forest carabids that reached Cape York or farther into tropical Queensland were not very numerous and some of them are patchily distributed now (see [63] and Table 9). And the Australian open-country carabids that reached even southern New Guinea were not very numerous and were ap- parently severely filtered, for very few of the many flightless carabid stocks of Aus- tralia reached New Guinea. Tlie barrier in this case seems to have been primarily ecologic, perhaps an alternation of areas of rain forest and open country like that which actually occurs on the Cape York Peninsula now, and which may have im- posed (different) barriers to both rain- forest and open-country stocks, regardless of the direction of their dispersal. Tliat the three principal barriers to dis- persal of Carabidae betAvcen Asia and Australia have probably acted in somewhat different ways is, I think, important. To- gether, however, they have had a common, net result: the accumulation on N^w Guinea of a carabid fauna composed almost entirely of small, winged, active carabids, many of them belonging to inherently successful, widely distributed, actively dis- persing groups. Among these actively dispersing groups, the hydrophiles are outstanding. Hydro- phile Carabidae, that live beside water, are usually winged and do fly relatively often either to escape rising water or to main- tain populations in shifting water-side habitats. Because they live in unstable, shifting habitats, they presumably have to maintain relatively large populations, and , this may be an additional advantage to them in dispersal. Water-side habitats are to some extent independent of forest cover, so that Carabidae associated with them can disperse relatively easily across areas like present Cape York, where the forest cover is discontinuous. And presence of water probably gives some protection against extremes of climate, so that hydro- philes can cross climatic barriers relatively easily too. Hydrophile Carabidae do in fact seem to be exceptionally good dis- persers. They have reached New Guinea in sufficient numbers not only to occupy their own habitats but also in several cases to invade the floor of rain forest, where they have apparently compensated for a deficiency of less vagile forms primarily adapted to the forest floor [97]. I have elsewhere (1959a; 1962a; 1965) suggested that not only the winged Australian Bembi- dion but also the now chiefly flightless and chiefly mesophile "Trechiis" of south- temperate Australia have been derived from winged ancestors which crossed the tropics from Asia to Australia at low alti- tudes in water-side habitats. [85] Amount of dispersal, noiv and in the past. Amount of dispersal — number of stocks dispersing between New Guinea and other areas regardless of direction — can be correlated with time, although the data are necessarily imprecise and the correlation is rather rough. Tlie method is to count and compare the numbers of stocks of New Guinean Carabidae at three taxonomic levels: at the level of non- endemic species, which are shared with the Oriental area and/or Australia, and each of which has had its own separate dispersal; at the level of endemic species or groups of species not in endemic genera, counting only species or species groups that seems to have separate relationships outside New Guinea; and at the level of endemic genera or groups of genera, count- ing only those with separate relationships outside New Guinea. A general correlation of taxonomic level with time is assumed. That is, it is assumed that, in spite of 234 Bulletin Museum of Comparative Zoology, Vol 142, No. 2 probable differences in rate of evolution of different stocks, nonendemic species have in general dispersed more recently than the ancestors of endemic species, which in turn have in general dispersed more recently than the ancestors of en- demic genera. The kind of unit to use for this puqoose must be considered carefully. To count only stocks that show direction of dispersal [74, 75] would weight the results in favor of the more recent stocks, because clues that show direction are likely to be clearest in the stocks that have dispersed most recently. And the stocks to be counted now are not the same as my "geographic units" [31, 58]. The latter were designed to show existing relationships, and some wide-ranging stocks were therefore counted twice: e. g., a New Guinean species that occurred also in both the Orient and Aus- tralia was counted as both one Or and one Au unit. Now, each stock will be counted only once. Also, in scoring "geo- graphic units," I have counted only stocks with discernible geographic relationships, while now I shall count all stocks that seem independent within the New Guin- ean fauna, whether or not their geographic relationships are discernible. The resulting units might be called amount-of-dispersal units. They are designed simply to show the numbers of separate stocks that make up the existing New Gmnean carabid fauna, correlated with taxonomic level and therefore presumably with age. Since the New Guinean carabid fauna is as a whole a relatively recent one [89], most amount- of-dispersal units probably represent move- ments into New Guinea, but this is not assumed in the calculation. Table 16 summarizes the amount-of-dis- persal units in the existing New Guinean carabid fauna. Before interpreting it, I should ask whether changes in my con- cepts of species and genera would sig- nificantly change the results — whether, if I "split" species and/or genera, the table would be significantly changed. I think Table 16. Summary of aaiount-of-dispersal units: NUMBERS OF SEPARATE STOCKS OF NeW Guinean Carabidae at three taxonomic levels Nonendemic species Endemic species and species groups in nonendemic genera Endemic genera and groups of genera Other than Agonini Agonini 128 9 ±129 ±9 11 ±5 the answer is that the counts would be changed, and that it might become neces- sary to tabulate units at additional taxo- nomic levels, including perhaps subgenera and natural groups of genera, but that if the classification were approximately phylogenetic, the table would still show many more dispersals at more recent than at the oldest level. Table 16 indicates that the existing New Guinean carabid fauna consists of many stocks that have not differentiated specifi- cally, many that have become distinguish- able species but not genera, but very few that have become distinct genera. This suggests that stocks that have dispersed more or less recently far outnumber those that dispersed longer ago. The difference in number of dispersals in proportion to time may have been greater than the figures show: the nonendemic species may have made their movements within a relatively short time; the endemic species may represent dispersals over a much longer time; and the endemic genera, a still longer one, so that the number of dispersals now represented in the New Guinean fauna by endemic genera may have been widely spaced over a very long period. This is diagrammed in Figure 16. For this and other reasons actual amounts of dispersal cannot safely be calculated from Table 16. Nevertheless, the table does strongly suggest that one of two things has happened. Either amount of dispersal — number of groups moving into or out of New Guinea — has increased The cababid beetles of New Guinea • Darlington 235 (0 o o a: UJ CD 2 iNON-E., ANCESTORS OF . |SPEC. , ENDEMIC SPEClESi IN NON-ENDEMIC ' GENERA ANCESTORS OF ENDEMIC GENERA I T HIS MORE RECENT SITUATION HYPOTHETICAL OLDER MORE RECENT OLDER i TIME Figure 16. Histogram of numbers of stocks in relation to time in tfie existing New Guinea carabid fauna. Division of the an- cestors of the endemic species and endemic genera into "more recent" and "older" categories is arbitrary. See text for further explanation. enormously among Carabidae in relatively recent times. Or faunal overturn [68] has eliminated a large proportion of older stocks as new ones have come in. I prefer the latter explanation. It is consistent with the relatively recent nature of the New Guinean carabid fauna as a whole [89], and it is consistent also with my general bypothesis of dispersals and replacements if successive carabid groups over the world [67]. My conclusion is that, although the ex- isting New Guinean carabid fauna is made up of many recently dispersed stocks and comparatively few old ones, this situation need not be the result of a recent increase in amount of dispersal. Carabidae may have been reaching New Guinea in num- bers for a very long time, and the present fewness of old stocks may be due to ex- tinction and replacement. I see no indi- cation that New Guinean Carabidae form 236 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 t\vo groups, one old and one relatively re- cent, separated by a time when few or none of the beetles reached the island. On the contrary, the concept of continual ar- rivals with continual overtinns seems to me to fit the observed situation. Tliere is therefore, as I shall say again [88], no initial starting point to be looked for in the accumulation of Carabidae on New Guinea. Of course this simple conclusion about the dispersal history of New Guinean Carabidae covers an immense ignorance of details and also ignores many complicat- ing factors and processes. For example, although I have noted that rates of evo- lution probably vary among different groups of Carabidae, I have not sufficiently emphasized how differences in rates of evolution may have affected some parts of the New Guinean carabid fauna. This is perhaps best shown by comparing the Agonini with the other carabids on the island. The Agonini are tabulated sepa- rately in Table 16. In general, the agonine figures confonn to the table as a whole in that number of dispersals seems to have been greater among more recent than among older stocks, but in the case of the Agonini the correlation has been blurred by radiations on New Guinea, the radi- ations having proceeded to the point where the number of initial ancestors can no longer be determined with any accuracy. But I think the Agonini do fit the main pattern reasonably well, and that so far as they differ, the difference is due not to their being older on New Guinea but to their having evolved there either more rapidly or more diversely. [86] Suuimanj of ii,eop,raphic origins of Netc Guinean Carabidae. The history of Carabidae in the Asiatic-Australian seg- ment of the world seems to have conformed in general (but of course not in detail) to the better documented histoiy of verte- brates. The largest favorable area (the Old-World tropics, including tropical Asia) has apparently been the principal center of evolution and dispersal of domi- nant Asiatic-Australian Carabidae. The smaller and less favorable area of Australiai has been a less important center. And the still smaller area of New Guinea has been least important in the evolution and dis- persal of dominant carabid stocks. Carabids have apparently been coming: into New Guinea continually during considerable period. There is no good evi- dence that arrivals were more numerous at some times than at others; arrivals have apparently been very numerous recently, but faunal overturns, with extinctions per- haps of many species, may have obscured the evidences of earlier arrival rates. Both Oriental and Australian stocks have prob- ably reached New Guinea at all times. The incoming Oriental have probably always exceeded the incoming Australian stocks in number and importance, except that among terrestrial mesophiles living in relatively dry, relatively open country Australian arrivals have been more nu- merous at least recently. Otherwise, incom- ing Oriental stocks have probably been more numerous than Australian ones re- gardless of size of insects, regardless of wing-state (almost all were winged), in all main habitats, and at all altitudes, as well as at all times. There must also, 1 think, have been continual extinctions correlated with the arrivals, and if the ar- rivals have been more from the Orient than from Australia, extinctions ("withdrawals") also have probably tended to begin at the Oriental end of the area and proceed toward Australia. The result has probably been a gradual shift of major distribution patterns from Asia toward Australia, caused by the procession of both dispersals and extinctions in this direction. But the details of this pattern have been excessively complex and have been further compli- cated and partly obscured by local evo- lutions and radiations of some groups within New Guinea and elsewhere. The broad movement of Carabidae from Asia toward Australia is, I think, part of The carabid beetles of New Guinea Darlington 237 the world-wide pattern of evolution of successive dominant groups in the great, climatically favorable area of the Old- World tropics (Africa and tropical Asia) and of dispersal into smaller and/or cli- matically less favorable areas, with re- placement of older by more recently dominant groups [67, 68]. And indications af extensive overturn in the New Guinean Fauna itself [85] and evidences (not given here) that Carabidae have dispersed from New Guinea eastward to smaller islands, 3n which numbers of species of Carabidae are at least roughly proportional to area \nd isolation, are consistent with Mac- Arthur and Wilson's (1967) theory of directional dispersal, faunal overturn, and "aunal equilibrium on small islands. (In ^act my diagram (Fig. 16) of the age itructure of the New Guinean carabid auna, although differently constructed, ?an be considered a model of faunal equi- librium comparable to the MacArthur- Wilson equilibrium model.) The New Guinean carabid fauna thus fits into and connects both the apparent world^^ide pattern and the local insular pattern of svolution, dispersal, and faunal balance. [ find this a very satisfying, unifying con- :ept. EVOLUTION [87] Evohitionanj perspective. For a rhorough discussion of animal species and ?volution, with the necessary background 3f modern biology and genetics, see Mayr, 1963. And for a careful statement of the ipparent role of evolution in detennining lattems of dispersal and resultant distri- jution patterns, see Darlington, 1965, Chap- ters 5 and 6. New Guinea can legitimately 36 treated as an evolutionary center of its u)wn, but it should be remembered that it ilso has its place in worldwide evolution ind dispersal patterns [67]. The patterns ire apparently determined by the relative ireas of different pieces of land, and by 'limate. The fact that New Guinea, al- hough a very large island, is still much smaller than Asia or Australia has presum- ably limited its importance as a center of origin and dispersal of major groups [70], but has probably not limited either rate or diversity of evolution of species upon the island. [88] Evolution of Neic Guinean carabid fauna as a whole. Geographic differenti- ation and overturn of Carabidae on New Guinea have been considered briefly in [85]. Now to be discussed in more detail are the evolution of the New Guinean cara- bid fauna as a whole, the differentiation and radiation of particular carabid stocks, and some general trends in the exolution of Carabidae on the island. The starting point for discussion of the evolution of the New Guinean carabid fauna is the assemblage of ancestral stocks from which the fauna has evolved. These stocks were evidently pre-adapted in several ways. They were pre-adapted for dispersal across barriers, the pre-adapta- tions including usually possession of wings (the whole New Guinean carabid fauna is or may be derived from ancestors that were winged when they reached the island), small (or at least not very large) size, probably other characteristics of structure and behavior including the habit of flying actively, and often adaptation to water-side habitats. Some, but not all, of the initial stocks (especially some Agonini) had also characteristics — general adapta- tions [68] — that made for general domi- nance, i.e., for success in a variety of situations. The pre-adaptations that favor dispersal and the adaptations that favor continued existence in island habitats are partly dif- ferent and opposed. For example, wings and flight pre-adapt a stock to dispersal, while existence in some habitats on some islands favors atrophy of wings. Or, for another example, small size favors disper- sal, but continued existence on some islands may in some cases ( perhaps on Madagascar [20] but apparently not on New Guinea) favor increase of size. There mav therefore 238 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 be a reversal of direction of selection and stocks in these habitats, the deficiency adaptive evolution after carabids reach an being compensated for in part by multi- island, and tliis in part explains the extraor- plication of the comparatively few rain- dinary distinctness of many insular cara- forest stocks that have reached the island bid faunas, including that of New Guinea and in part by ecologic shifts of hydro- especially at higher altitudes. To restate philcs onto the rain-forest floor. This is this important generalization in different consistent with the general rule stated by words: dispersal from a source fauna to Wilson (1961) for the ants of New Guinea an island selects as ancestors of the island and other islands: that dispersals tend to fauna fractions of the source fauna that occur in marginal habitats, and that after happen to be pre-adaptcd for dispersal; dispersal some stocks penetrate the rair but when these fractions evolve and radiate forest and evolve and diversify there, to form a whole fauna on the island, di- Water-side habitats apparently are mar- rection of selection is partly reversed, and ginal so far as the rain forest is concerned the new fauna that evolves is likely to be They are certainly less stable and also more very different from the source fauna in widely distributed than habitats on the gross taxonomic composition as well as in ground in rain forest. Carabidae that live many details. beside water do seem to disperse relativel) Most of the ancestral stocks from which easily and do apparently undergo e-cologic the New Guinean carabid fauna has shifts onto the rain-forest floor [84, 97]. evolved apparently reached the island at So, I take as the starting point of evo- low altitudes anel were adapted to lowland lution of the New Guinean carabid faunn tropical habitats. Carabidae in such habi- an initial assemblage of immigrants: small tats, including tropical rain forest, often do winged, dispersible forms concentrated al seem to be small and winged [21]. This low altitudes and adapted to existence ir may be partly because carabid populations the lowland tropics, but "disharmonic' in these places tend to have "patchy" distri- taxonomically, with a surplus of Agonini butions [22] maintained by continual redis- and disharmonic also ecologically, with a persals, so that the characteristics that surplus of hydrophiles but a moderate pre-adapt for dispersal to an island also shortage of stocks living in rain forest ane pre-adapt Carabidae to survive in lowland an extreme shortage of stocks adapted tc tropical habitats on the island. However, mountain habitats. I call this an initial dispersal across a tropical archipelago not assemblage, but I elo not think of it as ha\' only strongly selects small, winged, low- ing a single starting point. It was, rather land Carabidae but also probably has other a changing or evolving continuum, a sorl filtering effects. Carabidae tend to be of faunal germ plasm, of relatively un more active and more liable to dispersal in specialized forms. We do not know when some habitats than in others, so that some it began. We do know or at least suppose e^cologic groups disperse more readily anel that it has been continually changing b\ reach islands more often than the others "faunal overturn," by arrival of a succession do. Dispersal may be more rapid in rel- of ne^w stocks and extinction of old ones atively open country than in heavy forest, although some of the older stocks have And carabids that are associated with been evolving and radiating even while water apparently disperse more readily new ones have been continuing to arrive- than those that are not. The multiplication I think that overturn has probably beer of species in some groups (e. fi,., Deme- so extensive as to destroy most or all of the trida, see Part III, p. 140) in foliage and earlier stage\s of accumulation of Caraliieku on tree trunks in rain forest in New Guinea on New Guinea. There are two reasons foi suggests an initial deficiency of carabid thinking that overturn has been extensive i i The carabid beetles of New Guinea • Darlington 239 First, many species of Carabidae have From this changing continuum different reached New Guinea so recently that they carabid stocks on New Guinea have are not yet differentiated there, and if evolved to different extents and in differ- one accepts the idea of faunal balance as ent ways. The most striking processes have applying to all faunas everywhere ( except been multiplications of species and ecologic extremely young or very isolated ones), radiations. These processes are further New Guinea must have lost many older discussed below [91, 92]. As far as the stocks to compensate for the arrival of new fauna as a whole is concerned, the princi- ones. And second, the New Guinean cara- pal results of the multiplications and radi- bid fauna seems to possess no striking ations have been greatly to increase the evolutionary or geographic relicts, nothing number and diversity of both ground-living that seems very old, or taxonomically iso- and arboreal Caralyidae in rain-forest habi- lated, or geographically very distant from tats, and especially to fonn on the higher its relatives, as if all really old members mountains of New Guinea a complex alti- of the changing and evolving fauna have coline fauna which is ecologically like the been eliminated. New Guinea possesses carabid faunas of mountains elsewhere no endemic tribes; endemic genera are (Darlington, 1943) but which consists few; and all of them have or may have largely of genera and species which have relationships with other Carabidae still apparently evolved on and are confined to existing in the Oriental or Australian the island. Most of this alticoline fauna Regions. If such genera as Mecyclothorox seems to have been derived primarily from and Loxandras are geographic relicts, they the surrounding lowlands of New Guinea, have survived primarily in Australia rather by differentiations of alticoline stocks from than in New Guinea. Australia and New lowland ancestors and by ecologic radi- Zealand possess a number of geographi- ations especially of Agonini at high alti- cally isolated carabid stocks (e. g., the tudes. The "momitain hoppers" [80] that "carenums" [35] and various Pterostichini have reached New Guinea make up a com- [39] in Australia, and certain stocks with paratively small part of the mountain apparent northern relationships on New carabid fauna [90]. Zealand [89] ) which may have come long So, the New Guinean carabid fauna can ago via New Guinea but are not repre- be thought of as derived from a changing sented there now, and this is at least con- continuum of relatively unspecialized, sistent with disappearance of older stocks small, winged, lowland ancestors, including on New Guinea. I shall note again this many hydrophiles and some relatively un- point — the apparent absence of relicts specialized Agonini, which have been among New Guinean Carabidae — in con- coming in over a long period of time, and sidering the relative age of the fauna [89]. from which have evolved on New Guinea The actual rate of overturn (arrivals/ex- a great, partly endemic rain-forest fauna tinctions ) can perhaps be calculated and a diverse, unique mountain fauna. The eventually by formulae something like complexity of this process and of the whole those devised by Mac-Arthur and Wilson carabid fauna it has produced is, of course, ( 1967 ) . But neither existing data nor my far beyond what I can describe or even mathematics are adequate now. satisfactorily indicate. To summarize: the "initial assemblage" [89] Relative age of the Neic Guinean From which the New Guinean carabid fauna. If the New Guinean carabid fauna fauna has evolved should be thought of as has evolved not from an initial set of an- a changing continuum of which we cannot cestors beginning at one point of time but >ee the beginning but only a relatively from a constantly changing accumulation recent segment. of ancestors, including relatively general- 240 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 ized Agonini, to which additional incoming stocks have continually been added while other stocks have continually been elimi- nated, an exact determination of the age of the fauna may be impossible even in theory, and absence of a fossil record makes dating it in terms of geologic time impossible in practice in any case. We do not know the geologic age of New Guinea; we do know that some of the mountain ranges are geologically recent, but we do not know how long a significant piece of land has existed where New Guinea now is; we do not know when Carabidae were first able to reach and exist on proto-New Guinea, if there was one; and we do not know whether the ancestors of the existing fauna were the first carabids on the island or whether they were preceded by others that have disappeared during faunal over- turns. However, although we cannot de- termine absolute age, we can say some- thing about the relative age of the New Guinean carabid fauna, its age in relation to the faunas of other land areas. Both tropical Asia and Australia have carabid faunas that include noteworthy evolutionary and geographic relicts. Tropi- cal Asia, for example, has an endemic tribe (Idiomoqohini), as well as at least a few genera which are isolated taxonomically and/or widely separated geographically from their closest relatives (for example, Mouhotia, a genus of enormous, flightless scaritines confined to the Indo-Chinese Peninsula), and Australia has two endemic tribes (Agonicini in the southeast and Cuneipectini in the west), as well as Famhorus, a striking endemic assemblage of large scaritines [35], diverse and isolated pterostichines (and psydrincs) [39], and others. New Guinea has no comparable relict or isolated Carabidae. But perhaps no island, not even a large one, should be compared with continents. A more significant comparison can be made with the carabid faunas of New Caledonia and New Zealand. New Cale- donia has several genera of Carabidae so distinct that their relationships are doubt- ful, or so isolated geographically that their dispersal routes are lost. For example, the New Caledonian genus C\iphocoleus (eight species) is so distinct tliat it is not clear whether it belongs in the Agonini or in the Lebiini, and the New Caledonian scaritine genus Anomophaenus (eight species) is not related to anything now existing in Australia or New Guinea, its nearest relatives being (perhaps) in tropi- cal Asia. And New Zealand has an endemic tribe ( Zolini ) , an extraordinary relict genus {Maoripamhorus, related to the Australian Pamhorus), and endemic groups of Tre- chini, Agonini (Sphodrini), and perhaps Bemlndion of which the closest existing relatives seem to be in norf/j-temperate areas (for discussion of these cases see Darlington, 1965: 64). In contrast. New Guinea, although it has a much larger carabid fauna than either of the other islands, has no endemic tribe of Carabidae and relatively few (few in proportion to the size of the fauna) endemic genera, and all of the latter have or may have relatives in adjacent areas, in either the Oriental Region or Australia. I conclude that the carabid fauna of New Guinea is more recent in its origins than the faunas of tropical Asia or of Australia or of New Caledonia or New Zealand. It should be re-emphasized that the relative ages of the faunas do not neces- sarily indicate the relative ages of the islands. What the relatively recent age oJ the New Guinean carabid fauna probably does indicate, I think, is that, because oi the greater accessibility of the island. Carabidae have flooded into New Guinea in much greater numbers than into New Caledonia or New Zealand, and that fauna! overturnis have therefore been more rapic and more thorough in New Guinea. [90] Evolution of the mountain fauna The general characteristics of the moun- tain carabid fauna of New Guinea are the same as those of mountain faunas else- where (Darlington, 1943). The mountair The carabid beetles of New Guinea • Darlington 241 species are indicated on my data sheets [16]. Species are relatively few in any single mountain faunule, but geographic replacements are frequent, and the total number of species on all the mountains of New Guinea is very great [19]. The size range of the mountain species is within the range of the lowland species but with a single mode at 9-9.5 mm, which is larger than the larger of the two lowland modes; but this is apparently due not to increase af size of Carabidae on the mountains but to failure of many small forms to reach bigh altitudes [20]. Incidence of species ^vith atrophied wings increases with alti- ude, reaching about 95 per cent on the highest mountains; this is a result of itrophy of wings (and multiplication of >pecies following wing atrophy) in New Guinea, not of accumulation of wing- itrophied stocks from outside the island [21-23]. Ecologically, most of the moun- tain species are mesophiles, of which the majority live in wet montane forest, fewer in grassland above the forest zone; a few are hydrophiles living on the banks of nountain streams; and a few arc arboreal [24, 25]. The effects of altitude on Carab- idae [26] may be partly direct, but in- direct effects seem more important and probably include climatic control of vege- tation, limitation of areas, reduction of competition with ants, and presumably 3ther factors. The mountain carabid fauna of New i^uinea, of which some characteristics are reviewed above, is composed of very many separate stocks, derived mostly from the iurrounding lowlands, over a considerable period of time as indicated by their dif- erent degrees of differentiation. Tlie most ["ecent movements, probably still going on, nay be of lowland species into mid-altitude ?rass areas deforested by man [26]. Many )ther of the 161 species of Carabidae cnown to occur both lielow 500 and above 1000 m [19] may also have moved (spread) rom the lowlands to mid-altitudes re- t'litlv. In a few cases altitudinal differ- entiation has apparently just begun, for example, in Brachidius crassicornis (Part I, p. 508; Tax. siippL), in which mountain individuals are relatively large; in Proso- pogmiis garivagliae (Part I, p. 537), in which lowland and mountain individuals differ slightly in elytral striation and width of intervals; and in T richotichmis nigri- cans (Part III, p. 52), which is fully winged at low altitudes but dimorphically winged on the Bismarck Range. In many other cases differentiation has proceeded to the point where mountain-living (usu- ally mid-altitude) species now exist in genera which are well represented also in the lowland fauna. Such genera in which the mountain-living species are still winged include (this is far from a full list) Tachys, Notagonum (see also below), Colpodes, Iridagomim, T richotichmis, Horpcdoxenus, Hyphaereon, Catascoptis, Agonochila, and especially Demetrida. A few primarily winged lowland genera include mountain species in which the wings have atrophied; for examples see Clivino ] from base, with pos- terior angles sharply formed, c. right; disc convex, middle line distinct, anterior im- pression not sharply defined, posterior transverse impression deep, baso-lateral im- pressions small; entire base of pronotum punctate. Elytra: width elytra /prothorax 1.65; base margined, margin subangulate at. humeri; subapical sinuations obsolete; apices apparently broadly rounded ( slightly broken ) except angulate or vaguely sub- dentate at suture; striae impressed, not punctulate; intervals c. flat. Inner wings The cababid beetles of New Guinea • Darlington 279 fully developed. Lower surface: sides of sterna in part punctate; abdomen not pubes- cent. Legs: 4th hind-tarsal segment emarginate, scarcely lobed. Secondary sex- ual characters of i normal; 2 unknown. Measurements: length 9.7 mm; width 3.3 mm. Type. Holotype $ (Bishop Mus.) from Owen Stanley Rge., Papua, Goilala, Bome, 1950 m, Mar. 16-31, 1958 (W. W. Brandt); the type is unique. Notes. For comparisons, see preceding Key to Species of Reversum Group. Nofagonum ambulator n, sp. Description. With characters of genus and of reversum group, except wings atro- phied; form as in Figure 29, elytra more narrowed basally than usual; brown, head and pronotal disc darker, appendages yel- low; moderately shining, reticulate micro- sculpture faint on head, very light and transverse on pronotum; irregularly trans- verse on elytra. Head 0.87 and 0.84 width prothorax; eyes small but more abruptly prominent than usual, with posterior supra- ocular setae behind posterior eye level. Prothorax: width/length 1.15 and 1.15; base/apex 1.16 and 1.23; base and apex margined; sides broadly sinuate well before base, with basal angles sharply formed, c. right or slightly acute; lateral margins very narrow; disc convex, with middle line and anterior transverse impression nonnal, pos- terior transverse impression deeper (in the type), and baso-lateral impressions small but rather deep; whole basal area strongly punctate. Elytra: width elytra prothorax 1.60 and 1.55; base margined, margin faintly subangulate at humeri; subapical sinuations slight or obsolete; apices independently rounded then sinuate to denticulate sutural angles; striae deep, not punctulate; intervals convex. Inner wings atrophied, reduced to vestiges c. Mi as long as elytra. Lower stir- face: sides of pro- and mesosterna punc- tate; abdomen not pubescent. Legs: 4th hind-tarsal segment emarginate, with very short lobe on outer side. Secondary sexual characters of i normal; ? unknown. Mea- surements: length c. 7.8; width 2.7-2.8 mm. Types. Holotype $ (CSIRO) from Mur- mur Pass (W of Mt. Hagen), N-E. N. G., 8600 ft. (c. 2620 m), Nov. 1961 (W. W. Brandt); and 1 ? paratype (Bishop Mus.), 32 km E Wapenamanda (Western High- lands), N-E. N. G., 2500-2700 m, June 9, 1963 (M. Sedlacek). Notes. Although evidently related to N. reuersior (Darlington 1952, Part H, p. 137), this species seems distinct by prothorax much less sinuate with the sinuations nearer base, as well as by atrophy of wings. These differences may conceivably all be due to a single mutation, but I cannot assume this without further evidence, and the occur- rence of a short winged species of Notago- num is worth recording now. Nofagonum externum Darlington Darlington 1952, Part II, pp. 131, 138. Additional material. Papua: 2, Popon- detta, 25 m, June 1966 ( Shanahan-Lippert, Bishop Mus.), hght trap; 1, Mt. Lamington, 1300-^1500 ft. (c. 400-450 m) (C. T. Mc- Namara, South Australian Mus.). Notagonum sigi Darlington Darlington 1952, Part II, pp. 131, 143. Additional material. N-E. N. G. : 3, Wau, 1100 m, Sept. 9, 1961 (Sedlacek). Notes. N. sigi is superficially very much like vile but lacks the sparse but distinct ventral pubescence of vile and has a slightly narrower prothoracic base, although the proportions of both species vary and may overlap. The fact that the three \A^au in- dividuals were all taken at one time and place suggests that they came from one particular habitat, while vile is apparently much more widely distributed altitudinally and perhaps ecologically. Notagonum sinuum Darlington Darlington 1952, Part II, pp. 131, 139. Additional material. Papua: 1, Biniguni, 280 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 Gwariu R., 150 m, "No. 3," July 27-Aug. 14, 1953 (Geoffrey M. Tate, AMNH). N-E. N. G.: 1, Adelbert Mts., Wanuma, 800-1000 m, Oct. 24, 1958 (Gressitt); 1, Finisterre Rge., Budemu, Stn. No. 51, c. 4000 ft. (c. 1220m), Oct. 15-24, 1964 (Bacchus, British Mus.); 2, Herzog Rge., Vagau, Stn. No. 137, 147A, c. 4000 ft. (c. 1220 m), Jan. 4-17, 1965 ( Bacchus, British Mus. ) . Notes. This species, now known from all three political divisions of New Guinea, occurs chiefly at moderate altitudes in the mountains, but evidently descends to or nearly to sea level. Notagonum altum Darlington Darlington 1952, Part II, pp. 131, 144. Additional material N-E. N. G.: 4, Mt. W'ilhelm, 2800-2900 m, July 6, 1963 (Sed- lacck); 2, Sarua Kup, Kubor Rge., Oct. 31, 1965 (Dept. Agr. Port Moresby); 6, Finis- terre Rge., S side Mt. Abilala, Stn. No. 100, 8000 ft. (c. 2440 m), Nov. 17, 1964 ( Bacchus, British Mus. ). Nofagonum margaritum Darlington Darlington 1952, Part II, pp. 131, 145. Additional material. One hundred one, from all three political di\'isions of New Guinea, including die following. N-E. N. G. : 16, Wau & vie. (incl. Mt. Missim, Mt. Kaindi), 1100, 1150, 1200, 12.50, 1600-1650, 1800, 2.300 m, dates in Jan., Feb., May, June, Sept., Oct., Dec, 1961-1966 (Sedlaceks and others). WestN. G.: 60, Star Rge., various localities, 1260, 1300, 1500 m, dates in May, June, July, Aug., 1959 ( Leiden Mus., Neth. N. G. Exp.), many at light including ultra- violet light; 1, Waigeu Is., Camp Nok, 2500 ft. (c. 760 m), Apr. 1938 (Cheesman). Notes. This species is apparently com- mon over an exceptionally wide altitudinal range, from near sea level in some cases (see localities given in 1952) to middle altitudes in the mountains. Notagonum subpuncfum Darlington Darlington 1952, Part II, pp. 131, 146. Additional material. Papua: 10, Kiunga, Fly R., dates in July, Aug., Sept. 1957 (W. W. Brandt, Bishop' Mus.'); 1, Kokoda, 360 m, Mar. 20, 1956 (Gressitt), light trap; 1, Mamoo Plantation, Northern Dist., Mar. 22, 1956 (Dept. Agr. Port Moresby), at Hght. N-E. N. G.: 2, Minj, Western Highlands, 5200 ft. (c. 1600 m) May 20, 1960 (J. H. Barrett, Dept. Agr. Port Moresby), at m. v. lamp. West N. G.: 1, Star Rge., Sibil, 1260 m, June 16, 1959 (Leiden Mus., Neth. N. G. Exp.), at light. Notes. This species, including subspecies capitis Darlington (1952: 147), is now known from nearly the whole length of New Guinea, from sea level to moderate altitudes in the mountains. However, it has not yet been found in the Morobe area. The specimens from the Fly River repre- sent a population which, in size and elytral microsculptiue, is more like subspecies capi- tis from the Vogelkop than like typical suhpunctum from Dobodura (northern Papua) etc., but which differs slightly from capitis in form of elytral apices, which are usually a little more distinctly angulate than in capitis. However, extremes of the two forms are hardly distinguishable. I do not wish to describe a Fly River form now, but mention it as an example of geographic variation at low altitudes within New Guinea. Tliis kind of local geographic varia- tion will probably be found to occur ini many other lowland carabids when ade- quate series from different parts of the island are available. Notagonum denfellum Darlington Darlington 1952, Part II, pp. 132, 147. 1963, Breviora ( Museum of Comparative Zoolog>0, No. 183: 3,5. Additional material. Twelve, from all three political divisions of New Guinea and' Goodenough Island, including the following localities. N-E. N. G.: 1, Bulolo (Morobe area), 730 m, Aug. 31, 1956 (E. J. Ford, Jr., Bishop Mus.); 1, Western Highlands: Baiyer, 1150 m, Oct. 19, 1958 (Gressitt), light trap; 1, Torricelli Mts., Mobitei, 750 m, The carabid beetles of New Guinea • Darlington 281 Apr. 1-15, 1959 (W. W. Brandt, Bishop Mus. ) . Notes. This variable species is widely distributed in New Guinea at low and mod- erate altitudes (subspecies chimhu Darling- ton (1952: 149) is on the Bismarck Range at at least 5000 ft. ( c. 1525 m ) ) and occurs also in tropical northern Australia. Nofagonum subimpressum Darlington Darlington 1952, Part II, pp. 132, 149. Additional material. Eleven, from all three political divisions of New Guinea and Fergusson Is.; none above 500 m (pre- vious highest record, 800 m, in Snow Mts. ); 1, Sangeman Village, nr. Busu R., NE of Lae, 25 m, Aug. 30, 1957 ( D. Elmo Hardy, Bishop Mus. ), at light; not found near Wau. Nofagonum paludum Darlington Darlington 1952, Part II, pp. 132, 150. Additional material. Papua: 1, Popon- detta, 25 m. May 1966 ( Shanahan-Lippert, Bishop Mus.); 1, Normanby Is., W'akaiuna, Sewa Bay, Nov. 11-20, 1956 {W. W. Brandt, Bishop Mus. ) . Nofagonum addendum Darlington Darlington 1952, Part II, pp. 132, 153. Additional material. West N. G.: 5, Waigeo Is., Camp Nok, 2500 ft. (c. 760 m), April 1938 (Cheesman). These specimens are in addition to 2 paratypes from the same locality recorded in 1952. Notes. For comments on the relation of this species to Altagonum vallicola, see the latter, below. Nofagonum angulum Darlington Darlington 1952, Part II, pp. 132, 154. Additional material. West N. G. : 5, Star Rge., various localities, 1260, 1300, 1500, 1800 m, dates in May, June, July 1959 (Leiden Mus., Neth. N. G. Exp.); 1, Wissel Lakes, Arabu Camp, 1800 m, 1939 (H. Boschma, Leiden Mus.); 1, Bokondini, 40 km N of Baliem Valley, c. 1300 m, Nov. 16-23, 1961 (no collector given), Malaise trap. N-E. N. G.: 20, Wau and vie. (Mt. Missim, Mt. Kaindi, Nami Ck., Edie Ck., Bulldog Rd.), 1100, 1200, 1250, 1700, 2000, 2200, 2300, 2350 m, Jan., Feb., Mav, June, Aug., Sept., Oct., Nov., Dec. 1961-1965 (Sedlaceks), some at light including m. v. light, also in Malaise trap; 8, same area (Edie Ck. and Mt. Kaindi), Stn. No. 10, 20, 7000, 8000 ft. (c. 2135, 2440 m), Sept. 17, 22, 1964 (Bacchus, British Mus.); 2, Fera- min, 1200-1500 m. May 11-22, 2.3-31, 1959 ( W. W. Brandt, Bishop Mus.); 1, Swart Vy., Karubaka, 1450 m, Nov. 12, 1958 (Gressitt), light trap; 3, Okapa, May, June, 1965 ( Hornabrook ) ; 2, same localitv, Stn. No. 170, c. 5000 ft. (c. 1525 m),>eb. 4-15, 1965 ( Bacchus, British Mus. ) ; 1, Wonenara, nr. Kratke Mts., 1450 m, June 14, 1966 (Gressitt), hght trap; 1, 11 km S of Mt. Hagen (town), 2000-2300 m. May 20, 1963 (Sedlacek); 1, Sarua Kup, Kubor Rge., Oct. 31, 1965 (Dept. Agr. Port Moresby); 8, Finisterre Rge., Budemu and Moro, Stn. No. 51, 78, 4000, 5500 ft. (c. 1220, 1675 m), Oct. 15-24, Oct. 30-Nov. 15. 1964 (Bacchus, British Mus.). Papua: 1, Owen Stanley Rge., Goilala: Tapini, 975 m, Nov. 16-25, 1957 (W. W. Brandt, Bishop Mus.). Notes. The occurrence of this ver\' dis- tinct species seems worth gixing in full, as an example of the distribution of a widely spread but strictly mountain living New Guinean carabid. Nofagonum subrufum Darlington Darlington 1952, Part II, pp. 133, 156. Additional material. N-E. N. G.: 1, Finisterre Rge., Moro, c. 5550 ft. (c. 1690 m), Oct. 30^Nov. 15, 1964 (Bacchus, British Mus.). Notes. This very distinct species has been previously known only from two specimens from Rattan Camp, Snow Mts., West N. G., at 1200 m. Nofagonum subspinulum Darlington Darlington 1952, Part II, pp. 133, 158. 282 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 Additional material. Papua: 1, Good- enough Is., "E. Slope No. 10," 900 m, Oct. 24-30, 1953 (Geoffrey M. Tate, AMNH); 1, Mt. Rill, Sudest Is., "No. 10," 250-350 m, Aug. 22, 1956 (L. J. Brass, AMNH). N-E. N. G.: 1, "Krisa, N. New Guinea, Vanimo," Apr. 1939 (Cheesman, South Austrahan Mus.). Notagonum astrum n. sp. Description. With characters of genus; form as in Figure 30; black, legs bicolored (femora dark at base pale at apex, tibiae dark, tarsi pale with small black spots at articulations), antennae dark with basal segment pale; moderately shining, reticulate microsculpture light, isodiametric on front, transverse on pronotum and elytra. Head 0.88 width prothorax; eyes large, normal. Prothorax: width/length 1.39; base/apex 1.18; sides rounded almost to base, then slightly sinuate before very obtuse, narrowly rounded posterior angles; lateral margins narrow; baso-lateral impressions small, vaguely punctate; base indistinctly, apex not margined at middle; disc convex, with distinct middle line, indistinct or irregular transverse impressions. Elytra: width elytra/pro thorax 1.72; margins broadly rounded at humeri; subapical sinuations weak; apices narrowly independently rounded; striae impressed, outer (not inner) ones slightly punctulate; intervals slightly convex. Lower siu-face not punctulate; ab- domen not pubescent. Legs: 4th liind- tarsal segment strongly lobed, outer lobe longer than inner. Secondary sexual char- acters of $ normal; ? unknown. Mea.sure- ments: length 8.0 mm; width 3.1 mm. Type. Holotype S (Leiden Mus.) from Star Rge., West N. G., 1300 m, "Bivak 39," June 28, 1959 (Neth. N. G. Exp.); the type is unique. Notes. In my key to species of Notago- num (Part II, pp. 130ff) this new species runs to couplet 2 but fits neither part of the couplet, the sides of the prothorax being sinuate relatively near the base but the abdomen not being pubescent. The new species may actually be most closely related to N. gibbum ( couplet 12 ) but has a wider head and bicolored legs, the latter distin- guishing the present species from any other Notagonum known to me Nofagonum exactum n. sp. Description. With characters of genus; form as in Figure 31, with elytra more oval than usual (but inner wings long and folded ) ; brownish black, lateral margins of elytra and (less distinctly) of prothorax narrowly testaceous, appendages testaceous except antennae browner from 4th segments; shining, elytra subiridcscent, reticulate mi- crosculpture light and so strongly transverse on elytra as to be scarcely distinguishable. Head 0.82 width prothorax; eyes normal. Prothorax cordate; width/length 1.39; base/ apex 1.07; lateral margins moderate; poste- rior angles right and exactly defined; base margined, apex not margined at middle; disc normal, baso-lateral impressions mod- erately deep, vaguely subpunctate, and base slightly longitudinally wrinkled at middle. Elytra quadrate-suboval, each slightly (in- dependently) impressed before middle; width elytra /prothorax 1.64; subapical sinu- ations moderate; apices subtruncate with sutural angles very briefly dehiscent and weakly subdentate; striae impressed, slightly irregular but not distinctly punctulate; in- tervals slightly convex. Lower surface c. im- punctate; abdomen not pubescent. Legs: 4th hind-tarsal segments rather strongly lobed, outer lobe longer than inner. Secondary sexual characters of c^ normal; 9 unknown. Meastirements: length 6.7 mm; width 2.6 mm. Type. Holotype 6 (Bishop Mus.) from Wau, Morobe Dist., N-E. N. G., 1200 m, July 16-22, 1962 (M. Sedlacek); the type is unique. Notes. In my key to the species of Nota- gonum (Part II, pp. 130ff) this runs to dentcllum, but the present species has the posterior angles of prothorax much better defined and the elytra more oval. The dif- The CARABiD BEETLES OF New Guinea • DarUngtotx 283 ference in shape of prothorax is striking on comparison of specimens. Nofagonum quadruum n. sp. Description. \\\\\\ characters of genus; form as in Figure 32, rather depressed, elytra impressed near or slightly before middle; black, lateral margins of prothorax and elytra narrowly testaceous-translucent, appendages testaceous except antennae slightly darker except at base; shining, retic- ulate microsculpture absent or faint on front and disc of pronotum, light but distinct and strongly transverse on elytra. Head 0.76 and 0.75 width prothorax; eyes normal. Fro- thorox: width/length 1.45 and 1.49; base/ apex 1.11 and 1.09; lateral margins moderate and moderately reflexed; apex finely mar- gined, base not or indistinctly so; disc with usual impressions, impunctate except vaguely subpunctate in baso-lateral impres- sions. Elytra: width elytra/prothorax 1.51 and 1.53; subapical sinuations strong; apices acutely angulate then emarginate to acutely denticulate (almost spined) sutural angles; striae impressed, not distinctly punctulate; intervals slightly convex. Loxcer swiace not or not much punctate; abdomen not pubes- cent, he OS: 4th hind-tarsal segment rather strongly lobed, outer lobe longer than inner. Measurements: length 8.2-9.8 mm; width 3.2-3.7 mm. Types. Holotype <^ (Bishop Mus.) and 9 paratypes (some in MCZ, Type No. 31822) from Wau, Morobe Dist., N-E. N. G., 1200 (1 para type 1200-1300) m, dates in Mar., Apr., Oct., 1961-1964 (holotype, Oct. 11, 1962) (Sedlaceks); and 1 additional para- type from Wau, 3400 ft. ( slightly over 1000 m), "3.8.62" (J. J. H. Szent-Ivany, Dept. Agr. Port Moresby ) . Additional material. Papua: 1, Tapini, (Owen Stanley Rge., c. 1200 m). May 17-19, 1961 (Gressitt). N-E. N. G.: 1, Torricelli Mts., Mobitei, 750 m, Mar. 16-31, 1959 ( W. W. Brandt, Bishop Mus. ) . West N. G. : 1, Star Rge., Sibil, 1260 m, June 16, 1959 (Neth. N. G. Exp., Leiden Mus. ). Measured specimens. The i holotype and 1 2 paratype from Wau, 1200 m. Notes. In my key to species of Notago- num (Part II, pp. 130ff), the present new species runs to margaritum Darlington (couplet 20) but has elytral denticles much more prominent than margaritum, elytral striae not punctulate, and differs in other ways. The individual from the Torricelli Mts. has dark rather than pale legs and may represent an independent population. Some of the specimens recorded above were taken in light traps including mercury vapor light traps. Nofagonum sectum n. sp. Description. With characters of genus; fomi as in Figure 33, with elytra indepen- dently impressed c. % from apex; black, lateral margins prothorax and elytra nar- rowly slightly translucent, appendages dark; moderately shining, reticulate micro- sculpture in part light or indistinct on head and pronotum, more distinct and transverse on elytra. Head 0.72 width prothorax; eyes large, normal. Prothorax wide-subcordate; width length 1.45; base/apex 1.24; lateral margins rather wide; apex margined, base faintly margined; disc weakly convex, baso- lateral impressions moderate, poorly de- fined, irregular but scarcely punctate. Elytra long-subquadrate, width elytra/ prothorax 1.46; apices strikingly modified, with outer-apical angles c. right and sharply defined, each apex then strongly emarginate to second angulation c. opposite end 3rd stria, then again emarginate to moderate spine at sutural angle; striae moderately impressed, sHghtly irregular but not punctu- late; intervals flat or slightly convex. Lower surface virtually impunctate; abdomen not pubescent. Legs: 4th hind-tarsal segments moderately lobed, outer lobe longer than inner. Measurements: length 9.8 mm (in- cluding spines); width 3.6 mm. Type. Holotype ? (Bishop Mus.) from Mokai Village, Torricelli Mts., N-E. N. G., 750 m, Dea 8-15, 1958 (W. W. Brandt); the type is unique. Notes. Although based on a single fe- 284 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 male, this species is so strikingly character- ized by form of elytral apices that it is worth describing. In my key to the species of Notagonum (Part II, pp. 130ff ) it runs to N. externum (couplet 7) because the outer- apical elytral angles are sharply defined, but the elytral apices are otherwise much more modified than in externum. These two species are probably not directly related. Genus VlOLAGONUhA Darlington Darlington 1956, Psyche 63, p. 8. Diagnosis and description. See reference given. Type species. Colpodes violaceus Chaud- oir. Generic distribution. Sec following Notes. Notes. This genus of medium-sized Agonini is distinguished from Colpodes by shorter head, and from Notagonum by posi- tion of elytral spines. The principal species of the genus is V. violaceum (Chaudoir), which occurs in New Guinea, New Britain, the Solomons, and NE Australia, with sub- species goa Louwerens (1956, Treubia, 23: 221, 231 ) in the Moluccas. V. CColpodes") piceiis ( Andrewes ) represents the group on Samoa. And an undescribed species occurs on the Palau Islands (Darlington, 1970: 23). Violagonum violaceum (Chaudoir) Colpodes violaceus auct. including Darlington 1952, Part II, pp. 159, 160. Additional material. Two hundred sixty- eight, from all three political divisions of New Guinea and Goodenough, Normanby, Woodlark, Rossel, Fergusson, Sudest, and Biak Is., and including the following. N-E. N. G.: 126, Wau and vie, altitudes from 1050 to 1700-1800 m (most from 1050 to 1300 m), dates in every month, 1961-1963 (Sedlaceks), some at light, some in Malaise trap. Papua: 2, Mt. Giluwe, 2500, 2,550 m. May 1, May 27-June 6, 1963 (Sedlacek). West N. G.: 4, Star Rge., various localities, 1220, 1260, 1500 m, dates in May, July, Aug. 1959 (Leiden Mus., Neth. N. G. Exp.). Notes. This is one of the commonest Carabidae in New Guinea. It evidently occurs throughout the island from sea level to moderate altitudes in the mountains, and rarely at higher altitudes. It is easily recog- nized by its rather broad form, size ( length c. 10 mm), purple or blue color, and elytra each with a single, moderate apical spine c. opposite the end of the second interval. It is partly arboreal, occurring in vegetation including clumps of leaves on low branches in rain forest and also in piles of dead leaves on the ground in forest. It often flies to light. Genus COLPODES Macleay Darlington 1952, Part II, pp. 115, 158. Notes. As I (and many other authors) have used it, this is a "genus of convenience" to which can be referred relatively large, usually tropical Agonini not now referable to more exactly defined generic groups. The single new species described below is probably not directly related to any previ- ously known New Guinean ^'Colpodes" but may be independently derived from Nota- iionum. Colpodes guega n. sp. Description. With characters of genus as restricted (Part II, pp. 158-159); form as in Figure 34; dark brown with lateral mar- gins of prothorax and (less distinctly) of elytra translucent, appendages paler; mod- erately shining, reticulate microsculpture faint and c. isodiametric on front, light and strongly transverse on pronotum, still more strongly transverse on (faintly iridescent) elytra. Head 0.74 width prothorax; eyes normal; front not wrinkled. Prothorax: width length 1.39; base/apex 1.41; lateral margins moderate; apex strongly margined, base very narrowly indistinctly so; disc con- vex, middle line distinct, transverse impres- ' sions slight; baso-lateral impressions poorly defined, not distinctly punctate. Elytra: width elytra prothorax 1.47; base margined, margin rounded-obtuse at humeri; apices \ The carabid beetles of New Guinea • Darlington 285 with outer angles not defined, subapical sin- uations broad, actual apices acutely angulate c. opposite ends 3rd intervals and obtusely angulate (almost minutely denticulate) at suture; striae moderately impressed, not distinctly punctulate; intervals slightly con- vex, 3rd 3-punctate as usual, Sth and 9th not specially modified at apex. Lower surface c. impunctate, without special pubescence. Legs: 4th hind-tarsal segments lobed, outer lobe longer than inner. Secondary sexual characters: ? with only 1 seta-bearing puncture each side apex last ventral seg- ment; i unknown. Measurements: length 11.5; width 4.3 mm. Type. Holotype 9 (sex detennined by dissection) (Bishop Mus. ) from Guega, W 3f Swart Valley, West N. G., 1200 m, Nov. 14, 1958 (Gressitt); the type is unique. Notes. In my key to Colpocles of New Guinea (Part II, pp. 159-160), this runs to couplet 9 but fits neither half of the couplet, differing from acuticauda in having elytra with entire basal margins and elytral 3rd intervals 3-punctate, and differing from sinuicauda and simplicicauda in having ely- tral apices conspicuously angulate. See also Notes under genus, above. Co/podes saphyrinus sloanei Maindron Dadington 1952, Part II, pp. 160, 161. Additional material West N. G.: 1, Star Rge., Bivak 39A, 1500 m, July 3, 1959, and 1, Star Rge., Sibil, 1260 m, June 21, 1959 (both Netherlands-New Guinea Exp., Leiden Mus.); 1, Waigeu Is., Camp Nok, 2500 ft. (660 m), Apr. 1938 (Cheesman). N-E. N. G.: 52, Wau, 980-1100, 1200, 1250, 1300, 1200-1500 m, Jan. (most), Feb., Mar., May, Aug., Sept., Oct., Dec, 1961-1961 (Sed- laceks), some in light traps; 1, Mt. Kaindi, 1000 m, July 9, 1963 (Sedlaceks); 1, Karimui, 1080 m, July 11-12, 1963 (Sedlacek); 5, Eliptamin Vy., 1200-1350, 1350-1665, 1665- 2530 m, June, Aug., Sept., 1959 (W. W. Brandt, Bishop Mus.). Papua: 1, Mt. Lamington, 1300-1500 ft. (c. 400-450 m) (C. T. McNamara, South Australian Mus.). Notes. This species is evidently charac- teristic of the lower mountain slopes in New Guinea. It rarely occurs near sea level (3 at Dobodura, previously recorded) and has rarely been found above 1500 m. Co/podes helluo Darlington Darlington 1952, Part II, pp. 160, 162. Additiomd material. West N. G.: 1, Star Rge., Tenma Sigin, 1800 m. May 20, 1959 (Leiden Mus., Neth. N. G. Exp.), at light (the types came from Rattan Camp in the Snow Mts. at 1150 m). N-E. N. G.: 6, Wau, altitudes from 1200-1400-1500 m, Mar., Apr., June, Sept., Dec, 1961-1964 (Sed- lacek, Gressitt), 1 taken at light; 9, Elipta- min Vy., 1200-1350 m, dates in June, July, Aug., Sept., 1959 (W. W. Brandt, Bishop Mus. ) ; 5, Okapa & vie, some at 1800-1900 m, Aug. 27, 1964 (Hornabrook and Sedla- ceks), some under stones; 3, Wanatabe Vy., nr. Okapa, Stn. No. 174, c. 5000 ft. (1525 m), Feb. 5, 1965 (Bacchus, British Mus); 1, Koibuga, E Highlands, 1500 m, July 5, 1963 (Sedlaceks); 1, Finisterre Rge., Budemu, Stn. No. 51, c. 4000 ft. ( 1220 m), Oct. 15-24, 1964 (Bacchus, British Mus.). Papua: 2, Owen Stanley Rge., Goilala: Tororo, 1560 m, Feb. 15-20, and Rome, 1950 m. Mar. 8-15, 1958 (W. W. Brandt, Bishop Mus.). Notes. This very distinct species is easily recognized by form ( Fig. 35 ) and form of elytral apices. It is evidently widely distrib- uted at moderate altitudes in New Guinea. Co/podes laefus (Erichson) Darlington 1952, Part II, pp. 160, 163. Additional material. Seventeen, from all three political divisions of New Guinea; most at low altitudes (usuallv near sea level) but 1, Wau, 1200 m, Jan. 29, 1963 (Sedlaceks), light trap, and 3, Eliptamin Vy., 1350-1665 and 1665-2530 m, dates in June 1959 (W. W. Brandt, Bishop Mus.). Notes. Colpodes laetus laetus is now known on the Bonin Is. as well as the Philippines, Celebes, New Guinea (not Australia), Solomons, and New Hebrides, and C. I. pacificus Andrewes is on Samoa 286 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 and the E Caroline Is. ( Kusaie ) ( Darling- ton, 1970: 24). Co/podes habilis Sloane Darlinj^non 1952, Part II, pp. 160, 164. 1963, Breviora (Museum of Comparative Zoology), No. 183: 5. Additional material. One hundred thirty- four, from all three politieal divisions of New Guinea, few near sea level, most be- tween 1000 and 2000 m, none specifically higher. Included in this total are 52 from Wau, 1050 to 2000 m, dates in every month, many specimens at light. Note.s. Occurs also on Biiru, New Brit- ain, Solomons, and Santa Cruz Is., and in tropical northern Australia. Colpodes bennigseni Sloane Darlington 1952, Part II, pp. 160, 165. Additional material. N-E. N. G.: 34, Wau & vie, 1200 m (most so labeled, but some without altitude), Feb., Mar., Apr., May, June, July, Aug., Nov., Dec, 1961-1966 (Sedlaceks and others), some in light traps; 1, Okapa, Mar. 20, 1964 ( Hornabrook ) ; 4, Eliptamin Vy., 1200-1350, 1665-2530 m, June, July, Aug., 1959 (W. W. Brandt, Bishop Mus. ); 1, TorricelH Mts., Mokai Village, 750 m, Dec. 16-31, 1958 (W. W. Brandt, Bishop Mus.). West N. G.: 4, Star Rge., Sibil, 1260 m, dates in Apr., May, June, 1959 (Leiden Mus., Neth. N. G. Exp.), some at light. Notes. This species, like habilis (above), occasionally occurs near sea level ( see local- ities cited in 1952) but is mainly character- istic of middle altitudes in the mountains. Colpodes rex Darlington Darlington 1952, Part II, pp. 160, 167. Additional material. West N. G.: 11, Star Rge., Bivak 34A at 850 m, Bivak 39A at 1500 m (most), Sibil at 1260 m, dates in May, June, July, Aug., 1959 (Leiden Mus., Neth. N. G. Exp.), at least 1 at light; 6, Japen Is., Camp 2, Mt. Eiori, 2000 ft. (610 m), Sept. 1938 (Cheesman); 7, Waigeu Is., Camp Nok, 2500 ft. (660 m), Apr. 1938 (Chees- man). N-E. N. G.: 6, Eliptamin Vy., 1200- 1350, 1350-1665, 1665-2530 m, dates in June, Julv, 1959 (W. W. Brandt, Bishop Mus.); 2,'Feramin, 1200-1500 m, June 7-14, 15-18, 1959 (W. W. Brandt, Bishop Mus.); 4, TorricelH Mts., Mobitei & Mokai Village, 750 m, dates in Dec. 1958, Feb., Mar., 1959 (\V. W. Brandt, Bishop Mus.); 1, Finisterre Rge., Saidor: Matoko, Aug. 28-Sept. 5, 1958 (W. W. Brandt, Bishop Mus.). Notes. It is surprising that this striking endemic species, which has been found in all three political divisions of the island ( Papuan examples are recorded in 1952 ) , has not been found at Wau or anywhere in the Morobe area. Otherwise the species seems to be widely distributed in New Guinea at moderate altitudes, rarely below 1000 m. Colpodes acuticauda Darlington Darlington 1952, Part II, pp. 160, 170. Additional material. N-E. N. G.: 1, Wau, 2500 m, Dec. 28, 1961 (Sedlacek); 1, Finisterre Rge., Saidor, Matoko Village, (c. 1500 m), Sept. 6-24, 1958 (W. W. Brandt, Bishop Mus.). West N. G.: 1, Star Rge., Bivak 39, June 28, 1959 ( Leiden Mus., Neth. N. G. Exp.). ,y Colpodes sinuicauda Darlington Darlington 1952, Part II, pp. 160, 171. Additional material. N-E. N. G.: 1, Wau,, 1700-1800 m, Oct. 7, 1962 (Sedlaceks); 1, Caves near Telefomin, Aug. 1964 ( B. Craig, South Australian Museum). Papua: 1, Mt. Dayman, Maneau Rge., 1550 m, N Slope No. 5, June 30-July 13, 1953 ( Geoffrey M. Tate,AMNH). Notes. The 2 types were from Sigi Camp, Snow Mts., West N. G., at 1500 m. This distinct species is therefore now known; from all three political divisions of New Guinea, but from a total of only 5 specimens. The carabid beetles of New Guinea • Darlington 287 Colpodes simplicicauda Darlington Darlington 1952, Part II, pp. 160, 172. Additional material. West N. G.: 2, Wissel Lakes, Enarotadi, 1550 m, July 25- 29, 1962 (Sedlacek) and Arabii Camp, 1800 m, 1939 (H. Boschma, Leiden Mus. ). N-E. N. G.: 11, Wau & vie. (incl. Edie Ck. and Mt. Kaindi), altitudes 1200 to 2400 (most over 2000) m, Jan., May, June, Oet., 1961- 1963 (Sedlaceks), some in light traps; 1, 16 km NW of Banz, 1700-2100 m, June 28-29, 1963 (Sedlacek); 1, 32 km E of Wapenamanda, 2500-2700 m, June 9, 1963 (Sedlacek); 1, Finisterre Rge., Saidor, Matoko Village, Sept. 6-24, 1958 (W. W. Brandt, Bishop Mus.). Papua: 2, Owen Stanley Rge., Goilala: Borne, 1950 m, Mar. 8-15, 1958 (W. W. Brandt, Bishop Mus.). Notes. This species too, like several of the preceding ones, is widely distributed at middle altitudes in the mountains of New Guinea, but it ranges somewhat higher than most of the others. Genus PUCAGONUM Darlington Plicagonum rugifrons Darlington Darlington 1952, Part II, p. 174. Additional material. West N. G.: 3, Star Rge., Bivak 36, 1220 m, July 29, 1959, and Bivak 39A, 1500 m, July 1, 3, 10, 1959 (Leiden Mus., Neth. N. G. Exp.). N-E. N. G.: 2, Edie Ck. (nr. Wau), 2000, 2100 m, Oct. 4-10, 1961, May 31, 1962 (Sedlaceks). Papua: 1, Popondetta, 60 m, Sept. 3-4, 1963 (Sedlacek); 1, Mt. Dayman, Maneau Rge., 1550 m, N Slope No. 5, June 30-July 13, 1953 (Geoffrey M. Tate, AMNH). Plicagonum fulvum Darlington Darlington 1952, Part II, pp. 174, 175. Additional material. West N. G. : 4, Star Rge., Bivak 40, 2330, 2360 m, July 19, 22, 29, 1959 (Leiden Mus., Neth. N. G. Exp.). Notes. This species is, so far as known, confined to West N. G.; the types were from the Snow Mts. However, the following spe- cies from the Morobe area, N-E. N. G., is apparently a geographic representative. Plicagonum kaindi n. sp. Description. With characters of genus; form c. as in P. fulvum Darlington ( Part II, Fig. 3); dark brown, head almost black, margins of prothorax and appendages more rufous; reticulate microsculpture absent on front, slightly transverse on pronotum, c. isodiametric ( at most slightly transverse ) on elytra. Head 0.78 and 0.80 width prothorax; front longitudinally wrinkled at sides, scarcely so at middle, irregularly slightly impressed, sparsely punctulate. Prothorax c. as in fulvum; width length 1.33 and 1.31; base apex 1.25 and 1.19; lateral margins wide, moderately reflexed, each with usual 2 setae; disc with usual impressions; base and apex strongly margined; baso-lateral impressions formed mainly by angles be- tween base and lateral margins, not dis- tinctly punctate. Elytra: width elytra/ prothorax 1.60 and 1.73; base margined, margin rounded at humeri; subapical sinua- tions moderate, apices more or less (obtusely but usually distinctly) angulate opposite ends 3rd intervals; striae moderately im- pressed, not distinctly punctulate, intervals flat or slightly convex, 3rd with only 1 ( the posterior) dorsal puncture; outer intervals not specially modified apically; no 10th intervals. Lower sutiace virtually impunc- tate; abdomen not pubescent. Legs: 4th hind-tarsal segment emarginate but not lobed; 5th segments without obvious acces- sory setae. Secondary sexual characters normal. Measurements: length 15.5-17.5 mm; width 5.6-6.3 mm. Types. Holotvpe 6 (Bishop Mus.) from Mt. Kaindi, 16 km SW of Wau, N-E. N. G., 2300 m, Oct. 5-7, 1962 (Sedlacek), m. v. light trap; 15 paratvpes (some in MCZ, Type No. 31825), same locality, 2200, 2300, 2400 m, dates in Jan., June, Oct., 1962-1963 ( Sedlacek ) , most in m. v. light trap; 1 para- type (AMNH), same localit\-, 2050 m. May 25, 1959 (L. J. Brass); and 2 paratypes labeled W'au (but perhaps actually from 288 Bulletin Museum of Comparative Zoology, Vol. 142, No. 2 Mt. Kaindi), 2400 m, Jan. 9-12, 1962 (Sed- laceks and others ) . Additional material N-E. N. G.: 1, Bull- dog Rd., 19-29 km S of ^^'an, 2200-2500 m, May 28, 1962 (Sedlacek). Measured specimens. The S holotype and 1 9 paratype from Mt. Kaindi. Notes. This may be a geographic form of Plica