Table Of ContentZOOSYSTEMATICA ROSSICA, 26(2): 307–314 25 DECEMBER 2017
New data on Lachnophorini and Odacanthini (Coleoptera:
Carabidae) of Vietnam, with description of two new species and
one new subspecies of Pseudophorticus
Новые данные о Lachnophorini и Odacanthini (Coleoptera:
Carabidae) Вьетнама с описанием двух новых видов и нового
подвида Pseudophorticus
D.N. FEDORENKO
Д.Н. ФЕДОРЕНКО
A.N. Severtsov Institute of ecology and evolution, Leninsky pr. 33, Moscow 119071, Russia.
E-mail: [email protected]
The Neotropical genus Pseudophorticus Erwin, 2004 is reported from the Eastern Hemis-
pere for the first time. Two new species, P. constricticollis sp. nov. and P. tetragonoderoides sp.
nov. are described from Vietnam; within the former species, two subspecies are recognized:
the nominotypical one (central Vietnam) and P. constricticollis tonkinensis ssp. nov. (northen
Vietnam). Additional records of Selina westermanni Motschulsky, 1858 in Vietnam are given.
Aeolodermus emarginatus (Chaudoir, 1872) is first reported from Vietnam.
Неотропический род Pseudophorticus Erwin, 2004 впервые обнаружен в Восточном
полушарии. Два новых вида – P. constricticollis sp. nov. с новым подвидом P. constricticol-
lis tonkinensis ssp. nov. и P. tetragonoderoides sp. nov. описаны из Вьетнама. Приведены
новые находки во Вьетнаме Selina westermanni Motschulsky, 1858. Aeolodermus emarginatus
(Chaudoir, 1872) впервые указан для Вьетнама.
Key words: ground beetles, taxonomy, Vietnam, Coleoptera, Carabidae, Lachnophorini, Oda-
canthini, Pseudophorticus, Selina, Aeolodermus, new species, new subspecies, new records
Ключевые слова: жужелицы, таксономия, Вьетнам, Coleoptera, Carabidae, Lachnophori-
ni, Odacanthini, Pseudophorticus, Selina, Aeolodermus, новый вид, новый подвид, новые
указания
INTRODUCTION in the subtribe Homethina Liebherr, 2016
of the tribe Odacanthini (Liebherr, 2016),
According to Erwin & Zamorano (2014), which makes the Lachnophorini restricted
the mostly Neotropical tribe Lachnophorini to the Neotropics, except for the monobasic
includes 19 genera with 159 described and Palaeotropical genus Selina Motschulsky,
many undescribed species. The genera Ae- 1858.
olodermus Andrewes, 1929 (monobasic Ori- During two expeditions sponsored by
ental), Homethes Newman, 1842 (nine spe- the Joint Russia-Vietnam Tropical Center
cies, southern Oriental Region to Australia), in 2015–2016, an unknown lachnopho-
and three Neotropical monobasic genera, rine was collected in central and northern
Diplacanthogaster Liebke, 1932, Quammenis Vietnam. Dr Terry L. Erwin (personal com-
Erwin, 2000, and Stenocheila Laporte de munication) has kindly identified a photo-
Castelnau, 1832, have recently been placed graphed specimen of that species as Pseu-
© 2017 Zoological Institute, Russian Academy of Scienсes
308 D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM
dophorticus sp. The genus Pseudophorticus TAXONOMY
Erwin, 2004 was erected for P. puncticollis
Order COLEOPTERA
Erwin, 2004 (type species) from Costa Rica
and three species earlier described in the Family CARABIDAE
genera Lachnophorus Dejean, 1831 or Eu-
Subfamily HARPALINAE Bonelli, 1810
phorticus Horn, 1881. In a later paper (Er-
win & Zamorano, 2014), the genus was said Tribe LACHNOPHORINI LeConte, 1853
to include one described species and many
Genus Pseudophorticus Erwin, 2004
undescribed ones.
A member of a rather large Neotropical Pseudophorticus constricticollis sp. nov.
genus found in Vietnam may suggest a re- (Figs 1, 2, 4–8, 12)
cent introduction; however, another Pseu-
Diagnosis. The species from Vietnam
dophorticus species was subsequently dis-
with distinctive colour pattern as described
covered in central Vietnam, which suggests
that both may be native to the area and ap- below. For additional differences from
parently still undescribed. the Neotropical species see ‘Remarks’ on
Descriptions of the two new species are P. tetragonoderoides sp. nov.
provided below, along with new records Description. Macropterous, body (Figs
and notes on habitats of Selina westermanni 1, 2). BL 4.7–5.9 mm. Moderately shiny
Motschulsky, 1858 and Aeolodermus emar- black, less shiny on pronotum and head
ginatus (Chaudoir, 1872). behind mid-eye level; elytra with nearly
indistinct blue violet tinge. Mouthparts,
including labrum, antennal scape and apex
MATERIAL AND METHODS
of pedicel reddish-yellow; mandibular api-
The acronyms used are as follows: ces slightly infuscated; legs flavous. Elytron
MPSU, Moscow Pedagogical State Uni- a third from apex with a yellow flexuose
versity; SIEE, the author’s reference collec- fascia spanning intervals 2–8 or 2–9; the
tion at A.N. Severtsov Institute of Ecology fascia narrow, generally consisting of three
& Evolution, Russian Academy of Sciences, subquadrate patches. Metaventrite red me-
Moscow; ZIN, Zoological Institute, Rus- dially, sides of metaventrite and often also
sian Academy of Sciences, St Petersburg; prosternum along middle reddish; abdomen
ZMMU, Zoological Museum of the Mos- medially reddish-black.
cow State University. Pubescence rather dense; eyes, center
The following parameters were ana- of head, clypeus, labrum and mandibles
lyzed: BL, maximum body length measured glabrous. Setae of dorsal pelage slightly
between apices of closed mandibles and ely- longer, directed slightly forward, slightly
tra; EL, maximum length of elytron; EW, longer on vertex and pronotal apex. Elytral
maximum width of elytra; HW, maximum pelage double, short and dense setae mixed
width of head across eyes; PL, length of with slightly longer and sparser ones. Ter-
pronotum along mid-line; PW, maximum minal maxillary and labial palpomeres very
width of pronotum; PWa, width of prono- sparsely setulose in basal half.
tum between apical angles; PWb, width of Microsculpture isodiametric, distinct on
pronotum between basal angles. frons before eyes, behind mid-eye level and
Measurements were taken using an over pronotal disc, with not well delimited
ocular micrometer within the accuracy of meshes here and there; almost obliterate on
two decimal places. The means are given in elytra and between eyes, obsolete on clyp-
round brackets for the ratios, with the num- eus, nearly so along pronotal apex and base.
ber of measured specimens (n) indicated for Head bisetose above each eye, posterior
the first ratio in the description. seta situated just in front of the level of eye
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM 309
Figs 1–3. Pseudophorticus, dorsal habitus. 1, P. constricticollis constricticollis ssp. nov., female para-
type; 2, P. c. tonkinensis ssp. nov., male paratype; 3, P. tetragonoderoides sp. nov., paratype.
posterior margin. Temples about a third as eral lobes and epilobes with pointed apices
long as moderately convex eyes, barely con- combined. Ligula short, apically trapezoidal
verging basad and extended into neck at and quadrisetose; inner setae being longer
very obtuse angle. Frons and vertex mod- and closer to outer ones than to each oth-
erately convex; neck constriction almost er. Terminal labial palpomere subfusiform;
imperceptible. Frons transversely, finely penultimate palpomere bisetose.
and densely, rugulose behind clypeus, with Pronotum cordate, convex, as wide as
a shallow subtransverse impression at the head, broadest about a fourth from apex.
level of eye anterior margin, often followed Sides rounded in apical half, often suban-
by small and shallow median pit or weak gulate at the widest point, converging ba-
V-shaped impression. Frontal foveae angu- sad, sinuate a fifth from base. Basal margin
late, deep and divergent before eyes; shal- evenly subconvex to nearly straight. Apex
lower and strongly diverging toward, and subtruncate medially to evenly sinuate
almost reaching, anterior supraocular setae. between acute and pointed apical angles.
Clypeus bisetose, convex, subsinuate api- Basal and apical beads traceable just lat-
cally. Antennae long, surpassing elytral base erally. Median line fine, moderately deep,
by apical 3–3.5 segments; antennomeres 3, deeper basally, reaching base, obliterate api-
4 and scape subequally long, slightly lon- cally. Basal transverse impression angulate
ger than antennomere 5, more than twice forward, deep medially, obliterate laterally;
as long as pedicel. Labrum truncate, sexse- apical transverse impression shallow. Basal
tose, with a distinct transverse impression. foveae linear, moderately deep, slightly
Mandibles imperceptibly concave laterally, converging apicad, almost reaching middle.
with widely explanate ventral scrobal ridge. Disc densely rugulose, more so basally. Lat-
Submentum bisetose; mentum bisetose at eral groove fine yet deep, with anterolateral
base of a large and acute median tooth; lat- seta at the widest point; lateral bead edged,
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
310 D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM
Figs 4–12. Pseudophorticus. 4–8, 12, P. constricticollis constricticollis ssp. nov.; 9–11, P. tetragonoderoi-
des sp. nov. 4–6, 9–11, median lobe of aedeagus, left (4, 9), ventral (5, 10) and dorsal (6, 11) aspects;
7, right paramere; 8, left paramere; 12, female reproductive tract; sab – spermathecal apical bulb,
sbb – spermathecal basal bulb, sg – spermathecal gland. Scale bar: 0.5 mm.
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM 311
reflexed, with much shorter posterolat- trapezoidal telescoped parts. Spermatheca
eral seta in or just before basal angle, 1–3 bipartite U-shaped, basal part being very
additional setae of similar length in apical slender, half as long as ovally elongated api-
angles, and a few shorter setae in basal half. cal part.
Propleura invisible in dorsal view. Etymology. Refers to the shape of the
Elytra square, convex, slightly flattened pronotum, from Latin constrictus (tied up)
on disc, EL/EW 1.41–1.51 (1.46, n=16), and collum (neck).
twice as wide as pronotum, nearly truncate Distribution. Central and northern Viet-
at bases, with square humeri, broadest a nam.
third from apex; outer angles widely round- Bionomics. A riparian psammophilous
ed; apical truncature oblique and almost species. All the adults were collected along
indistinctly sinuate; apices slightly pointed forest streams in bare fine gravel amid small
combined but blunt tips. Sides subparallel, stones, with negligible or no admixture of
diverging slightly apicad, usually gently mud.
sinuate a fourth from base. Basal ridge en- Remarks. The species description is
tire, straight, indistinctly angulate between based on two isolated populations treated
striae 4 and 5. Striae entire, indistinctly here as subspecies.
crenulate, deep, slightly deeper before apex.
Intervals nearly flat, convex in apical third, Pseudophorticus constricticollis
each with about three irregular rows of fine constricticollis ssp. nov.
setigerous punctures. Parascutellar striole (Figs 1, 4–8, 12)
long and deep; parascutellar setigerous pore
adjoining striae 1 and 2. Interval 3 with Holotype. Male, “Vietnam, Kon Tum
Prov[ince]., Chu Mom Ray Nat[io]n[al]. Park,
three discal setigerous pores (d1–d3); these
14°30´N 107°43´E, dak Car river bank, h = 700–
subequally spaced; interval 7 with two api-
850 m, 28.III–4.IV.2015, D. Fedorenko leg.”
cal setae; umbilical seta series continuous,
(ZMMU).
consisting of 15 setigerous pores adjoining Paratypes. 21 males and females taken to-
stria 8, except for pore 14 adjoining stria 9; gether with the holotype (SIEE, ZIN).
pores 1–3 and 10 adjoining both striae. Diagnosis. Body (Fig. 1): Labrum red-
Abdominal sternites IV–VI with one dish-yellow; antennomeres 7–11 flavous or
pair of ambulatory setae, sternite VII api- whitish; antennomeres 2–6 black to brown
cally bisetose in male, quadrisetose in fe- black, sometimes with antennomeres 5 and
male. 6 barely paler. Preapical elytral fascia span-
Legs. Metatarsomere 1 as long as those ning intervals 2–8 or 2–9. Some specimens
2 and 3 combined. Protarsomeres 1–3 not exhibiting slightly infuscated labrum or
dilated in male, each with adhesive ventral elytra with a vague posthumeral spot on in-
scales. Metafemur without fixed setae. tervals (6–)8.
Aedeagus (Figs 4–8) widely membra- Pronotum: PW/HW 0.96–1.01 (0.99,
nous dorsally, parameres subequally long. n=8), PW/PL 1.23–1.29 (1.26); sides
Internal sac behind middle with a patch of slightly diverging in basal fifth. Basal angles
several tiny spicules. projecting slightly laterad, right and sharp
Female genitalia (Fig. 12): basal gono- or with acute apices due to base gently in-
coxite (gonosubcoxite) with 10–11 apical dented just inside. Apex a fifth wider than
setae; apical gonocoxite triangular, with base, PWa/PWb 1.15–1.28 (1.21).
two outer (ventral) ensiform setae and a Elytra: EW/PW 1.96–2.10 (2.0); discal
double apical nematiform seta. Female re- setigerous pores slightly yet distinctly fove-
productive tract (Fig. 12) as described for ate, either adjoining stria 2 or d1 adjoining
the tribe (Liebherr, 1988; Liebherr & Will, stria 3.
1998); bursa copulatrix with four, inversely Distribution. Central Vietnam.
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
312 D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM
Pseudophorticus constricticollis h = 570 m, dak Lar Pa river bank, 8–20.V.2017,
tonkinensis ssp. nov. D. Fedorenko leg.” (SIEE).
(Fig. 2) Diagnosis. The species from Vietnam
with distinctive colour pattern as described
Holotype. Male, “N[orthern]-Vietnam, Phu below.
Tho Prov[ince]., ~ 90 km W of Hanoi, Xuan Description. Similar to P. constricticollis
Son N[ational]. P[ark]., h = 300 m, 21°07´58´´N
sp. nov., except as follows: Body (Fig. 3)
104°55´45´´E, in moist gravel, 6–15.VI.2014,
larger, BL 6.5–6.8 mm, yellow (including
D. Fedorenko leg.” (ZMMU).
gula, prosternum and mesosternum along
Paratypes. 34 males and females taken to-
middle), with whitish scape, maxillary
gether with the holotype (SIEE, ZIN).
palpomere 2 and femora. Head, pronotum,
Diagnosis. Distinctive from the nomino-
elytral base and epipleura, and metaster-
typical subspecies in the following points:
num laterally black. Elytra in middle third
Antennae more or less gradually paling
with a vaguely infuscated macula spanning
apicad, with segment 7 intermediate in
intervals 1–5 and laterally merging into a
colour between brown to dark brown seg-
darker subtriangular patch. Elytral dark
ments 3–6 and pale brown to brown seg-
parts, including central macula, with slight
ments 8–11; scape red, pedicel red to red-
(holotype) or no (paratype) metallic green
dish-brown; segments 7–11 slightly darker
tinge. Labrum dark brown, mouthparts
than legs (versus same coloured or slightly
largely yellow. Antennae rather pale brown,
paler than legs). Labrum mostly infuscated,
with reddish-yellow segments 2–4 and
indistinctly reddish. Elytral posthumeral
barely infuscated segment 5. Dorsum dull
spot absent, preapical fascia slightly nar-
due to coarse sculpture: head and pronotum
rower, spanning intervals 3–8, with inner
rugose and thence indistinctly microscu-
patch reduced to interval 3 and sometimes
lptured; frons rugulose, finely yet densely
separated. Pronotum on the average barely
punctate, except for a small smooth spot
more transverse, PW/PL 1.24–1.33 (1.28,
between eyes. Dorsal pubescence dense and
n = 8), and narrower basally, PWa/PWb
slightly longer than in P. constricticollis sp.
1.19–1.28 (1.24), due mainly to sides par-
nov., with pronotal lateral edge minutely
allel before, instead of diverging towards,
tuberculate in basal half; elytral intervals
base; base and apex less arcuate, often
each with 4–5 rows of setigerous punctures.
subtruncate medially; basal angles mostly
Pronotum: PW/PL 1.24–1.28, just as
not or barely projecting laterad. Elytral
wide as head, PWa/PWb 1.15–1.18; basal
discal setae not or barely foveate, situated
angles projecting slightly laterad, right and
mostly at middle of interval 3. PW/HW
sharp. Base finely beaded in lateral 1/4–1/5;
0.94–1.01 (0.97), EW/PW 1.94–2.11 (2.0).
apical bead fine yet entire, faint just medi-
Etymology. Refers to the provenance of
ally. Median line entire and deep, deeper ba-
the type series.
sally and apically. Apical transverse impres-
Distribution. Northern Vietnam.
sion fairly deep, basal transverse impression
wide and deep.
Pseudophorticus tetragonoderoides Elytra very slightly oblique at bases; hu-
sp. nov. meri slightly more widely rounded than in
(Figs 3, 9–11) P. constricticollis sp. nov.; apices narrowly
rounded; EL/EW 1.51–1.53, EW/PW
Holotype. Male, “Vietnam, Gia Lai Province,
1.96. Striae rather shallow; intervals flat,
~55 km ENE Pleiku, 14°17´45´´N/ 108°26´57´´E,
interval 3 along middle with three feebly
Kon Ka Kinh Nat[io]n[al]. Park, h = 600 m,
impressed discal setigerous pores; interval
8–20.V.2017, D. Fedorenko leg.” (ZMMU).
7 with two apical setae. Interval 9 with 17
Paratype. Female, “Vietnam, Gia Lai Pro-
vince, ~55 km ENE Pleiku, 14°17´24´´N/ umbilical setae, not constricted behind the
108°27´47´´E, Kon Ka Kinh Nat[io]n[al]. Park, middle (versus constricted at setigerous
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM 313
pore 10); pores of umbilical seta series ad- Genus Selina Motschulsky, 1858
joining stria 8, with pores 1–3 additionally
Selina westermanni Motschulsky, 1858
adjoining stria 9; pore 16 adjoining stria 9
only. Material. Vietnam. Dong Nai Prov.: 10 males
Legs. Protarsomeres 1–3 slightly dilated and females, Nam Cat Tien National Park,
and squamose ventrally in male. 11°25´18´´N 107°25´44´´E, Expedition of the
Aedeagus (Figs 9–11): apical lamella in Joint Russian-Vietnamese Tropical Center,
dorsal/ventral view triangular. 20.XI.2004 D. Fedorenko leg. (SIEE); 1 female,
same data, except for 21.X.2004, at light HQL
Remarks. The genus Pseudophorticus has
450 (SIEE). Song Be Prov.: 1 specimen, 60 km
not been described in detail; only a diagno-
N of Ho Chi Minh, Phu Gia Vill., 3–13.X.1994,
sis had been provided, a brief comparison
A. Napolov leg. (MPSU). Gia Lai Prov.: 2 speci-
with Euphorticus and Lachnophorus (Erwin,
mens, ~ 55 km ENE of Pleiku, Kon Ka Kinh
2004), and a key to genera of Lachnophorini National Park, 14°17´45´´N 108°26´57´´E, h =
(Erwin & Zamorano, 2014). As long as the 600 m, at light, 8–20.V.2017, D. Fedorenko leg.
genus is still formally monobasic (Erwin (SIEE).
& Zamorano, 2014), I can only compare Distribution. Africa, Madagascar, Sri
Pseudophorticus constricticollis sp. nov. and Lanka, India, Thailand, Cambodia, Viet-
P. tetragonoderoides sp. nov. with P. puncti- nam, southern China (Hong Kong, Macao),
collis and Pseudophorticus sp. from Peru (il- and Sumatra.
lustrated in Erwin & Zamorano, 2014, Fig. Bionomics. Bates (1886) noted that
19). In the latter two species, the prono- S. westermanni as the only member of “Sub-
tum is wider (versus as wide as, or slightly family Lachnophorinae” in Ceylon ran “in
narrower) than the head, and about three moist places in the half-dry river”. In the
quarters (versus half) as wide as the elytra. Cat Tien National Park, most of the adults
According to Erwin (2004), the two Neo- were taken at the extreme edge of a pool
tropical species, including P. puncticollis, where they dwelt in leaf litter on slimy mud.
are not periaquatic, have wings strongly re-
duced, and one species is wing-polymorphic Tribe ODACANTHINI Laporte, 1834
(Erwin, 2004; Erwin & Zamorano, 2014).
Genus Aeolodermus Andrewes, 1929
On the other hand, both species from Viet-
nam are macropterous and riparian, like Aeolodermus emarginatus (Chaudoir,
many Anchonoderus. This, combined with 1872)
the disjunct range, suggests that the Ori-
Material. Vietnam. Dong Nai Prov.: 1 male,
ental Pseudophorticus may deserve at least
Nam Cat Tien National Park, 11°25´18´´N
subgeneric status. The relationships within
107°25´44´´E, Expedition of the Joint Russian-
Pseudop horticus need further study.
Vietnamese Tropical Center, at light, 10.VI.2005,
Etymology. Reflects similarity between
D. Fedorenko leg. (SIEE). Lam Dong Prov.: 2
the new species and some psammophilous
males, 25 km NNW of Bao Loc, Loc Bao env.,
species of the carabid genus Tetragonoderus 11°44´18´´N 107°42´08´´E, h = 800 m, 5–20.
Dejean, 1829 in elytral pattern. IV.2013, D. Fedorenko leg. (SIEE).
Distribution. Known from the type local- Distribution. Malay Peninsula, Borneo,
ity only. Sumatra, Celebes, Philippines. The first re-
Bionomics. The holotype specimen was cord for Vietnam.
collected in bare fine sand along a forest Andrewes (1929, 1933) reported
stream, and the paratype in plant debris on Ae. emarginatus from Celebes, Borneo, Su-
sandy riverbank with a thick facial layer of matra (Medan), and the Malay Peninsula,
mud. This means riparian and, given the and supposed that Melbourne, Australia,
distinctive body pattern, apparently psam- was wrongly indicated as the type local-
mophilous way of life. ity. Darlington (1956) also noted that the
© 2017 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 26(2): 307–314
314 D.N. FEDORENKO. NEW DATA ON LACHNOPHORINI AND ODACANTHINI OF VIETNAM
species was not probably Australian, while Impériale des naturalistes de Moscou, 45(2):
occurring from the Malay Peninsula to the 382–420.
Philippines. Darlington P.J. 1956. Australian carabid beetles
III. Notes on the Agonini. Psyche, 63(1):
Bionomics. The listed specimens from
1–10.
Loc Bao were taken on moist ground with
Erwin T.L. 2004. The beetle family of Costa Rica
dense vegetation at the edge of a swamp.
and Panama: Descriptions of four new genera
Remarks. This species was illustrated
and six new species with notes of their way
by Louwerens (1952) for comparison with of life (Insecta: Coleoptera). Zootaxa, 537:
Homethes microguttatus (Louwerens, 1952), 1–18.
with reference to neither examined material Erwin T.L. & Zamorano L.S. 2014. A synopsis
nor locality. of the tribe Lachnophorini, with a new genus
of Neotropical distribution and a revision of
the Neotropical genus Asklepia Liebke, 1938
AKNOWLEDGEMENTS
(Insecta, Coleoptera, Carabidae). Zookeys,
430: 1–108.
I am indebted to Dr T.L. Erwin (Smithso-
Liebherr J.K. 1988. Redefinition of the super-
nian Institution, Washington, D.C., U.S.A.)
tribe Odacanthitae, and revision of the West
and Dr Laura S. Zamorano (Universidad de
Indian Lachnophorini (Coleoptera: Carabi-
los Andes, Colombia) for discussion of the new
dae). Quaestiones Entomologicae, 24: 1–42.
species. I thank Vassili Belov (College Station,
Liebherr J.K. 2016. Cyphocoleus Chaudoir (Co-
TX, USA) for suggestions that have improved
leoptera, Carabidae, Odacanthini): descrip-
the manuscript. The study was supported by
tive taxonomy, phylogenetic relationships,
the Russian Academy of Sciences Biodiversity
and the Cenozoic history of New Caledonia.
Program.
Deutsche Entomologische Zeitschrift, 63(2):
211–270.
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