P. J. Darlington, Jr.
Australian Carabid Beetles VII. Trichosternus, Especially the Tropical Species.
Psyche 68:113-125, 1961.

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PSYCHE
Vol. 68 December, 1961 No. 4
AUSTRALIAN CARABID BEETLES VII.
TRICHOSTERNUS, ESPECIALLY THE
TROPICAL SPECIES
BY P. J. DARLINGTON, JR.
Museum of Comparative Zoology, Cambridge, Mass. This is one of a series of papers describing new Carabidae of zoo- geographic importance referred to in my account (1961b) of transi- tion of Australian wet forest carabid faunas. Some other papers of the series, including a list of my localities, are referred to below (p. 130). The present paper is concerned with Trichosternus, especially with the comparatively little known tropical species. Types of new species are placed, at least for the time being, in the Museum of Comparative Zoology; paratypes, with C. S. I. R. 0. at Canberra and usually in the Queensland Museum. Proportions given in the descriptions are calculated from actual measurements.
Items of geographic or evolutionary interest derived from Tricho- sternus are its total distribution; the occurrence of two very distinct (relict or primitive?) species, fax and mmtwurn, high on Mt. Bartle Frere (one reaching Mt. Bellenden Ker too) ; the distribution of T. frater and its apparent allies (subvirens and sirnplicipes of South Queensland and nudifies to fisheri in the tropics), which form a group of mostly allopatric forms in which the male front tarsi have apparent- ly been simplified in two different stocks and in which double invasion or hybridization may have occurred in a very limited area on part of the Atherton Tableland (pp. 122-125). Another case of hybridiza- tion (of eungella and mixtus) may have occurred on the Eungella Range (p. 127). See map for distribution of species on and north of the Atherton Tableland.
I cannot give an exclusive definition of Trichosternus. Characters used by Sloane (1894 etc.) and Tschitschirine (1902) fail among recently discovered species. Compared with Nurus, Trichosternus is usually smaller, more lightly built, with c? front tarsi usually dilated and with 3 segments squamulose below, while in Nurus the c? front tarsi are usually simple, rarely (e.g. in N. atlas Cast.) slightly dilated and with 2 segments squamulose. However, exceptional Trichosternus



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Voi.. 68, PI.A-I E 7




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19611 Darlington - Australian Carotid Beetles 11.5 resemble Nurus in having 8 tarsi simple (T. subvirens Chd., sim- plicipes Sl.,
nud'ipes Darl., soror Darl.) or with only 2 segments squamul~se (setosiceps Sl.) . Compared with Notononzus, Trichos- ternlus is usually larger, often with prosternal process setose (rarely in Notono7tzCus), often with alternate intervals of elytra raised, and often without distinct 10th intervals at outer edges of elytra, but all these characters fail in some species. A generic revision of the larger Australian pterostichines is much needed. The Australian genera should be compared with the New Zealand ones (Britton 1940) and phylogenies should be worked out if possible, and the old types, many of them in Europe, should be reexamined. However, I cannot under- take all this now.
The known range of T'richosternus is the eastern edge of Australia from the base of the Cape York peninsula (south of Cooktovvn) to central New South Wales (perhaps a little north of Sydney), with one species isolated in southwestern Australia ( Darlington I 953 ) and another on New Caledonia. All the tropical Australian species (ex- cept cordatus) live in rain forest, but some more-southern species occur in savannah woodland, and some enter or are confined to south tem- perate rain forest on the Dorrigo-Ebor and Mt. Royal plateaus. Before considering the tropical Trichosternus, I give the following tentative key to species south of the tropics, as a basis for comparison (cf. Sloane's key, 1899, pp. 567-569, and Tschitschirine's, 1902, pp. Known distribution of Truhosternus in tropical Queensland. The finely dotted line is the approximate eastern edge of high land (Atherton Table- land etc.). Arrows indicate occurrence southward, at increasing distances beyond the limits of the map, of the species indicated. The species are num- bered in the order in which they are treated in the text. No. 1, on Mt. B (artle) F(rere), is Trichosternus fax; 2, obscuripennis, which extends north beyond the limits of the map nearly to Cooktown; 3, montorum; 4, nudipes, on the Mt. Spec plateau; 5, soror; 6, frater, which occurs north to Mt. Lewis and south beyond the limits of the map on the Kirrama Range; 7, mutatus; 8, fisheri; 9, eungella, and 10, mixtus, both on the Eungella Range; 11, corda- tus, at the southern edge of the tropics; 12, spec, on the Mt. Spec plateau; 13, setosiceps, which is widely distributed on the south-central Atherton Table- land and occurs also south of the limits of the map on the Kirrama Range; and 14, kirrama, on the Kirrama Range. Nos. 1, 2, 3, 13, 14 are very distinct, phylogenetically isolated species. Nos. 4, 5, 6, 7, 8 are apparently interrelated, chiefly allopatric members of the frater group (see text). Nos. 9, 10, both on the Eungella Range, though structurally distinct, may be related to each other and may hybridize (see text). No. 11 represents a primarily south temperate rather than tropical stock. No. 12, on the Mt. Spec plateau north of Towns- ville, is apparently related to another south temperate (New South Wales) species.




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116 Psyche [December
-3-DO). I have specimens of all the 16 full species named in this except cyaneotinctus.
Tentative key to Trichosternus of eastern Australia south of tropics
Elytral intervals 3, 5, 7 with seta-bearing punctures (Mt. Royal Range and highest part of Dorrigo-Ebor plateau ; 2 species?) australicus Sl.
.............................................................................. Only 3rd interval (if any) with seta-bearing punctures on elytral disc ................................................................................................ 2 Mesosternum setose anteriorly ................................................. 3 Mesosternum not setose anteriorly ........................................... 7 .............................. Male front tarsi not dilated, not squamulose 4
............
Male front tarsi dilated, 3 segments squamulose below 5
Alternate discal elytral intervals scarcely differentiated; greenish .... (extreme SE. Q. & adjacent N. S. W. border) s.iibvirens Chd.
Alternate discal intervals slightly elevated; black or nearly so, with cupreous or greenish margins (Bunya Mts., S. Q.) .............................................................................. simfdicipes Sl. Seventh elytral intervals sharply carinate at base; prothorax rela- tively broad at base; length c. 22 mm. (extreme NE. N. S. W. and adjacent Q. border) ...................................... angulosus Chd. Seventh intervals roundly costate at base .................................... 6 Very large, c. 33-40 mm. (SE. Q. to Blackall and Bunya Mts. etc.) ......................................................................... renardi Chd. Not so large, c. 25-32 mm. (N. S. W. S. to Dorrigo, Williams River, Booral, etc., and N. to Stanthorpe-Dalveen area, S.Q.) .................................................................................... vigors! Gory Three supra-ocular setae each side .............................................. 8 Two supra-ocular setae each side .............................................. 9 Purplish (savannah woodlands of N. N. S. W. chiefly west of .............................................. the rain forest zone) superbus Cast.
Greenish (unknown to me; may be another savannah woodland species) .......................... cyaneotinctus Boisd. incl. atroviridis Sl. Additional lateral prothoracic setae present (savannah woodland and drier rain forest of S.Q., N. into edge' of tropics; probably ............................................ includes hecate Tsch.) cordatus Chd.
................ Only usual 2 pairs lateral pi-othoi-acic setae present 10
Very large, 34 mm. or mose ...................................................... n ...... Smaller, 34 mm. (maximum size of cyaneus only) or less I 2
Prosternal process setose ; black ; length 37-42 mm. (extreme NE. ...........................
N. S. W. and adjacent Q. border)
perator Sl.




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19611 Darlington - Australian Carabid Beetles 117 Prosternal process not setose ; purplish ; length 34-39 mm. ................................
(Blackall Range, SE. Q.)
porphyriacus Sl.
....
Posterior-lateral prothoracic setae near base at least 3 species :
wilsoni Cast., nitidicollis Cast., and speciosus Sl. Posterior-lateral prothoracic setae about of prothoracic length
................................................................................ before base I 3
Smaller, length c. 21-27 mm. ; black sometimes slightly bluish or purplish (widely distributed in NE. N. S. W. and S. Q., from ............................
Dorrigo to Blackall Range
marginiferus Chd.
Larger, length c. 28-34 mm.; wholly purple or blue (Mt. Royal ................
Range, Dorrigo, etc. ; possibly 2 species) cyaneus Cast.
The following key to tropical Australian Trichosternus differs from the preceding key in form (ranges and authorities omitted) because the tropical species are discussed individually. All the species have rather long, strong, curved mandibles; rather short, irregular, sub- parallel frontal foveae; well developed, emarginate mentum. tooth; palpi not strikingly modified; elytra margined at base; 8th (submar- ginal) stria lined with very small ocellate punctures; lower surface im- punctate or nearly so except for usual "fixed" punctures; and (so far as my observations go) other usual characters of the genus, except as otherwise stated.
Key to Species of Trichosternus of
Tropical Australia
Mesosternum setose anteriorly; dorsal elytral intervals equal or nearly so (except in ~nontorum} ................. ... .............. 2 Mesosternum not setose anteriorly; alternate elyti-a1 intervals elevated (except scarcely so in kzrrama) .................................... 9
Seventh intervals of elytra not sharply carinate at base (at most somewhat swollen) ; elytra without dorsal punctures .............. 3 Seventh intervals sharply carinate at base; elytra with (small) dorsal punctures ......................................................................... 4
Subparallel, prothorax quadrate; humeri subdentate; color blue- purple
...................................................................................... fax
Elytra oval, prothorax subcordate; humeri not dentate; black, elytra dull ............................................................... obscuripennis Elytra with alternate intervals moderately raised; (form de- pressed ; color greenish or bluish) ................................ montorum Elytra with discal intervals equal or nearly so, except at extreme base (frater group) .................................................................. 5
Male front tarsi not dilated, without squamules ..................... 6 Male front tarsi more 01- less dilated, always with som'e squa-



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Psyche [December
mules below ......................................................................... 7 Broader ; greenish ........................................................ nudipes Narrower ; bluish or purplish ............................................... soror Greenish ..................................................................... frater Black or purplish .................................................................... 8 .............. Black or faintly purple ; smaller ( I 8-20 mm.) mutatus
............................................. Purple ; larger (23-25 mm.) fisheri
Only 2 pairs supra-ocular and 2 pairs lateral prothoracic setae (but extra apical ventral setae) present ........................ eungella Extra supra-ocular and/or lateral prothoracic setae present .... 10 Prosternal process setose ; if prosternal setae broken or not visible, note 3 supra-ocular and 4 or more median-lateral prothoracic ............... setae each side, and odd elytral intervals costate nzixtus
Prosternal process not setose .............................................. I I Only 2 pairs supra-ocular setae present, and odd elytral interva1.s raised ........................................................................................ 12 Extra supra-ocular setae present, or odd elytral intervals (except .................... 7th) scarcely raised and humeri scarcely dentate I 3
Black .............................................................................. cordatus Purple ..................................................................................... spec Only I pair median-lateral prothoracic setae present; odd elytral ......................................................... intervals costate setosiceps
Extra lateral prothoracic setae present; odd elytral intervals (ex- cept 7th) not costate ................................................. kirrama Trichosternus fax new species
Form as figured (Fig. I ) , subparallel, not strongly convex; black, head slightly metallic, pronotum with green 01- bluish marginal chan- nels and copper-purple on disc brightest in baso-lateral areas, elytra with marginal channels blue or greenish blue ; pronotum shining, head and elytra slightly duller. Head c. 3/4 or slightly more width pro- thorax; eyes moderate, genae c. wide as eyes, rounded and somewhat narrowed to neck; antennae rather short (in genus), passing base of prothorax by I or 2 segments; 2 supra-ocular setae each side. Pro- thorax subquadrate, c. 1/3 wider than long at middle, not much nar- rowed behind; base c. 1/10 or more wider than apex (variable) ; apex broadly emarginate but angles not otherwise advanced, not mar- gined; base subtruncate, slightly emarginate at middle, strongly mar- gined at sides; sides weakly arcuate for much of length, broadly, slightly sinuate before c. right, well defined posterior angles; lateral margins narrow, slightly wider posteriorly, each with a seta about 1/3 from apex and on margin at basal angle; disc rather flat, middle



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1961 1
Dar/in.qton - '4 ustralian Carabid Beetles Fig. 1. Trichosternus fax n. sp. Fig. 2. T. montorum n. sp. Fig. 3. T. kirrama n. sp.
line deep, transverse impressions almost obsolete except for subbasal foveae each side midway between middle and sides, separated from margins by strong convexities; disc and foveae not punctate. Elytra c. 1/5 wider than prothorax, usually slightly narrowed anteriorly ; margin rectangular or almost acute at subdentate humeri; elytra with striae well impressed and punctate; intervals slightly convex but not costate, 3, 5, and 7 slightly wider than others but not more elevated, except 7th slightly elevated at base; narrow 10th interval indicated posteriorly; dorsal punctures lacking. Mesosternum anteriorly and prosternal process setose. Male front tarsi slightly dilated, 3 segments squamulose; d with I, 9 2 setae each side last ventral segment. Length 20-23; width 6.5-7.7 mm.
Holotype d (M. C. Z. Type No. 30,355) and 15 paratypes all from Mt. Bartle Frere, North Queensland, west slope 3000-5000 ft. (not found lower), Dec. 1957, collected by my wife, my son, and myself, in mountain rain forest.




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120 Psyche [December
This new species is sufficiently distinguished from others in the preceding key. It is not closely related to any other known species. It may prove to be generically distinct from Trichosternus, or it may be a relict of the ancestral stock of the genus. Trichosternus obscuripennis (Macl.)
Phis is the northernmost Trichosternus, and it descends to i-ela- tively low altitudes. Macleay (1887, p. 220) described it from the Mossman River. Sloane had it from Helenvale south of Cooktown. I have it from! Mt. Finnigan south of Cooktown, Thosnton Peak near Dainti-ee, Mt. Lewis near Mossman, and Black Mt. about 20 miles north of Kui-anda. Although most of my specimens were taken at 2000 or 3000 ft. altitude, several were found at or below 1000 ft. in strips of rain forest along the trail between Thornton Peak and the Dainti-ee River.
T. obscuripennis is a very distinct species but hardly a separate subgenus. Its lack of humeral teeth is approached in T. kirrama, and lack of dorsal elytral punctures is matched by fax, although neither of these species is close to obscurip'ennis in other ways. Trichosternus montorum new species
Form as figured (Fig. 2), rather broad, depressed; black with bluish, purplish, or sometimes greenish reflections, head and pi-onotal disc moderately shining, elytra a little duller. Plead about 4/5 width prothorax; eyes rather small, genae as wide or wider than eyes, isreg- ularly arcuate and somewhat narrowed to neck; antennae rather long (in genus) ; 2 supra-ocular setae each side. Prothorax cordate; 1/3 (å± wider than long at middle; base slightly (å 1/10) narrower than apex; sides rounded for about 3/4 of length, then rather strong- ly sinuate before c. right, scarcely blunted posterior angles; side mar- gins moderate, not much wider basally, each with a seta about 1/3 from apex and on margin at basal angle; apex not margined, broadly emasginate but angles not otherwise advanced ; base broadly emar- ginate at middle, slightly rounded toward sides, vaguely margined at sides but not at middle; disc depressed, middle line deep, transverse impressions weak ; baso-lateral foveae elongate but not strictly linear, slightly nearer margins than middle, not quite reaching middle of length, densely transversely rugulose, the rugulose areas meeting at middle posteriorly, with middle of base longitudinally rugulose ; disc otherwise impunctate. Elytra oval, usually slightly narrowed anteri- orly; margin somewhat obtusely angulate and finely toothed at hu- meri; elytra with finely punctate discal striae; intervals more or less convex, 3, 5, 7 increasingly elevated, 7th being sharply cai-inate for



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19611 Darlington - Australian Carabid Beetles 121 most of length (most strongly so near base) ; 10th interval indicated but not well developed; each 3rd interval with 0, I, or 2 dorsal punc- tures behind middle (often asymmetrical in number and position). Mesosternum anteriorly and prosternal process setose. Male front tarsi slightly dilated, 3 segments squamulose below; with I, Q 2 setae each side last ventral segment. Length 23.5-27; width 7.9-9.1 mm.
Holotype (M. C. 2. Type No. 30,356) and 39 paratypes all from west slope Mt. Bartle Frere, North Queensland, 3000-5000 ft. (none lower), Dec. 1957, collected by the Darlingtons, in mountain rain forest. Also 6, not types, from east side Mt. Bellenden Ker, 3000-4500 ft., Dec. 1957, taken by myself, in mountain rain forest. This seems to be a typical, rather generalized (primitive?) Trichos- ternus, without known close relatives. It is of course distinguished from other species in the preceding key. Trichosternus nudipes new species
Form about average; greenish black, the green most distinct at sides and baso-lateral areas of pronotum and marginal channels of elytra; head and pronotum shining, elytra duller. Head c. 3/4 (by measure- ment) width prothorax; eyes small but abrupt, genae c. wide as eyes, convex, somewhat narrowed to neck; antennae of moderate length; 2 supra-ocular setae each side. Prothorax subcordate, moderately nar- rowed at base, c. 1/3 (A) wider than long at middle ; base slightly (less than 1/10) wider than apex; apex broadly emarginate with angles (lateral channels) slightly advanced, not margined; base broadly emarginate at middle, slightly rounded laterally, margined laterally; sides broadly rounded for much of length, moderately sinu- ate near c. right or slightly obtuse, scarcely blunted basal angles; later- al margins narrow, not wider basally, each with I or sometimes 2 setae each side 1/3 or less from apex and I seta on margin at basal angle; disc not very convex, middle line fine, transverse impressions. weak; baso-lateral impressions deep, c. 1/3 length prothorax, separated from lateral margins by convexities but extending inward somewhat vaguely nearly to middle; disc of pronotum impunctate except base irregularly rugulose. Elytra 1/3 or less wider than prothorax, sub- oval, slightly narrowed anteriorly ; margins forming c. right or slight- ly obtuse, finely raised-dentate angles at humeri; dorsal striae very fine, finely punctate; each elytron with 7 (including suture) rather weak but almost entire costae, 7th carinate basally; 10th interval in- dicated toward apex; 3rd interval usually 2-punctate, the punctures at or before apical 1/3 and near apex, but individual punctures some-



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122 Psyche [December
times absent or obscured. Mesosternum anteriorly and prosternal process (sometimes slightly) setose. Male front tarsi narrow, without squamules; c? with I, $2 2 setae each side last ventral segment. Length 22.^-26; width 7.9-9.5 mm.
Holoty~e
(M. C. 2. Type No. 30,357) and 87 paratypes all from Mt. Spec plateau (Paluma Range), c. 40 miles north of Towns- ville, North Queensland, 2000-3000 ft., Dec. 1957 and Feb. 1958, taken by the Darlingtons, in rain forest; and I, same locality, June 7- 10, 1955, E. 0. Wilson.
This is similar to and perhaps related to T. subvirens Chd. of South Queensland, but subvirens has an entire mentum tooth (emarginate in nudipes), posterior lateral pronotal setae just inside (not on) the margin, and lacks or nearly lacks rugulose sculpture on the base of the pi-onotum.
Trichosternus soror Darl.
I described this species (1953) from 3 specimens from Millaa Mil- laa, on the Atherton Tableland. I did not find it at this locality dur- ing my last trip, but found it on both the lower and upper western slopes of Mt. Bartle Frere, at 2000-3500 and 3000-5000 ft., and on the east side of Mt. Bellenden Ker, at 3000-4500 ft. Although this small, bluish species has simplified male tarsi, like the preceding species, it may not be directly related to it but may be independently derived from a common ancestor perhaps frater ( below) .
Trichosternus frater new species
Slightly more slender than average of genus, rather convex; green- ish 01- bluish black, head and pronotuin moderately shining, elytra somewhat duller. Head 4/5 or more width prothorax; eyes small, genae about wide as eyes, rounded and somewhat narrowed to neck; 2 supra-ocular setae each side; antennae of moderate length. Pro- thorax subcordate, c. I/& wider than long at middle; base slightly narrower than apex; apex broadly e~niai-ginate but angles otherwise scarcely advanced, apex not margined; base emarginate at middle, rounded toward sides, margined at sides ; sides broadly arcuate through much of length, moderately sinuate toward base; basal angles c. right or slightly obtuse, scarcely blunted ; side margins rather narrow; not much wider posteriorly, each with I seta near or before anterior 1/3 and I on margin at basal angle; disc with distinct middle line, less distinct transverse impressions, and moderate baso-lateral foveae which reach posterior margin but are separated from lateral margins by con- vexities; surface of disc nearly smooth except with some (variable)



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19611 Darlington - Australian Carabid Beetles 123 transverse strigulation in baso-lateral impressions. Ely tra slightly more than 1/4 wider than prothorax, subparallel, slightly narrowed anteriorly; margin forming obtuse (nearly right), slightly dentate angles at humeri; dorsal striae formed by fine vaguely punctulate im- pressed lines in low spaces between intervals; latter subequal on disc, slightly raised but not costate, except 7th strongly raised and costate basally, 10th interval indicated posteriorly; each 3rd interval I- or 2-punctate posteriorly, position of punctures variable. Mesosternum anteriorly and prosternal process (sometimes inconspicuously) setose. Male front tarsi moderately dilated, 3 segments squamulose below; d" with I, Q 2 setae each side apex last ventral segment. Length 20-23 ; width 6.5-8.0 mm.
Holotype ~3
(M. C. 2. Type No. 30,358) and 73 paratypes all from Kirrama Range, inland fromi Cardwell, North Queensland, 2000-3000 ft. altitude, Dec. 1957 and Feb. 1958, collected by the Darlingtons, in rain forest. Also the following specimens, not types: I 2, mountains above ( SW of) Atherton, Atherton Tableland, 3000- 4000 ft., Dec. 1957 and Feb. 1958; I, near Black Mt., c. 20 miles north of Kuranda, probably near 2000 ft. altitude, May 1958; and 3, Mt. Lewis, near Mossman, probably near 3000 ft., Dec. 1957 ; all collected by the Darlingtons, in rain forest. These localities are all on the Dividing Range system, which has probably been the principal path of north-south dispersal of wet forest Carabidae in North Queensland.
T. frater may represent the ancestral stock, with dilated, squamu- lose male tarsi, from which nudipcs of the Mt. Spec plateau, subvirens and s'lmplicipes of South Queensland, and also (independently) soror of the Atherton Tableland etc. have been derived. These 5 species (including frater itself) are all allopatric and are generally similar in form and technical characters except for the differences in the tarsi. The two following new species may be local derivatives of the same stock with d tarsi still dilated and squamulose. See map for distribution of tropical species of the frater group. Trichosternus mutatus new species
Small; form about average for genus but rather convex; black, sometimes with faint purplish reflections; moderately shining, elytra slightly duller. Head 3/4 to 4/5 width prothorax; antennae rather short, scarcely extending beyond base of prothorax; eyes moderate, genae c. wide as eyes, rounded and narrowed to neck; 2 supra-ocular setae each side. Prothorax subcordate, 1/4 (2) wider than long at middle; base equal to or slightly narrower than apex; sides broadly