ZOOSYSTEMATICA ROSSICA, 22(2): 271–284 25 DECEMBER 2013

New species of bombardier of the genera and (Coleoptera: Carabidae: ) from Vietnam Новые виды жуков-бомбардиров родов Brachinus и Pheropsophus (Coleoptera: Carabidae: Brachininae) из Вьетнама

D.N. FEDORENKO

Д.Н. ФЕДОРЕНКО

A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, 33 Leninsky Pr., Moscow 119071, Russia. E-mail: [email protected]

Eight new species of bombardier beetles are described from Vietnam: Brachinus dalatensis sp. nov., B. ovalipennis sp. nov., B. belyaevae sp. nov. (all in the melancholicus species-group), Pheropsophus (Stenaptinus) bidoupensis sp. nov., Ph. (S.) ngoclinhensis sp. nov., Ph. (S.) kalya- kini sp. nov., Ph. (S.) similis sp. nov. and Ph. (S.) obliteratus sp. nov. (all in the aptinoides species-group). A key to species of the Pheropsophus aptinoides group of the Vietnamese fauna is provided. Из Вьетнама описаны восемь новых видов жуков-бомбардиров: Brachinus dalatensis sp. nov., B. ovalipennis sp. nov., B. belyaevae sp. nov., Pheropsophus (Stenaptinus) bidoupensis sp. nov., Ph. (S.) ngoclinhensis sp. nov., Ph. (S.) kalyakini sp. nov., Ph. (S.) similis sp. nov. и Ph. (S.) obliteratus sp. nov. Описанные виды Brachinus входят в состав группы melancholicus, виды Pheropsophus – в состав группы aptinoides. Для определения вьетнамских представителей последней группы предложена таблица. Key words: bombardier , , distribution, Oriental Region, Vietnam, Coleop- tera, Carabidae, Brachininae, Brachinus, Pheropsophus, new species Ключевые слова: жуки-бомбардиры, таксономия, распространение, Ориентальная об- ласть, Вьетнам, Coleoptera, Carabidae, Brachininae, Brachinus, Pheropsophus, новые виды

INTRODUCTION Jedlička (1964) and Habu (1967) reviewed the truncatipenne carabids of Southeast The subfamily Brachininae Bonelli, Asia and Japan, respectively. Only a quar- 1810 is a speciose group of ground beetles distributed over warmer regions of the ter of about 120 Oriental species of Brachi- world. In the Oriental Region and adja- nus and Pheropsophus have been recorded cent lands, more than 150 brachinine spe- in Indochina so far. Numerous new species cies have hitherto been reported, all in described below suggest that Indochina, the tribe Brachinini, the genera Brachinus including Vietnam, does support a rich Weber, 1801, Pheropsophus Solier, 1833, bombardier fauna. Styphlomerus Chaudoir, 1875, and The purpose of this paper is to describe Fischer von Waldheim, 1828. Although eight new species from Vietnam: three ones many new species have been described of the genus Brachinus (the melancholicus from this region during the last two de- species-group) and six species of the ge- cades (Kirschenhofer, 1986, 2003, 2010, nus Pheropsophus (the aptinoides species- 2011; Tian & Deuve, 2007; Hrdlička, 2009 group). All of these look similar, sharing the a,b; Gao & Tian, 2010), its fauna remains combination of a large body, uniformly dark poorly known. No revision of the Orien- pronotum and elytra, and shoulders obliter- tal Brachinini have been undertaken since ated due to the apterous condition.

© 2013 Zoological Institute, Russian Academy of Scienсes 272 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM

MATERIAL AND METHODS si; antennomeres 5–11 of the same colour as tarsi, usually somewhat paler or darker than The material was collected over the the basal antennomeres. course of several expeditions sponsored by Head mostly short and wide, with large the Russo–Vietnamese Tropical Center. and convex eyes. Labrum strongly trans- The holotypes and some paratypes of the verse, four or more times as wide as long, described new species are deposited in the antennomere 3 longer than 1 and 2 com- Zoological Museum of the Moscow State bined. Mentum as usual in the genus, nei- University; the remaining paratypes, in the ther median tooth nor any pits. Last labial Author’s synoptic collection at the A.N. Se- palpomere subcylindrical, slightly dilated vertsov Institute of Ecology & Evolution, apicad, with apex obliquely truncate. Neck Moscow (SIEE). constriction usually weak laterally, deeper The parameters used were measured as in the middle. follows: total body length from mandible Pronotum rather wide and flat, pubes- tips to elytral apices; elytral length from cence varying from sparse to almost miss- base to apex along suture; widths of the ing, mostly scarce or very scarce; (antero) pronotum and elytra in their widest point; lateral seta tending to be reduced. Male maximum width of the head including com- protarsi not or hardly dilated, with tarso- pound eyes; and pronotal length along mid- meres 1–3 ventrally with two longitudinal line. Unless otherwise indicated, the num- stripes of adhesive vestiture. ber of specimens measured (n) is only given Macropterous to apterous. Elytra with for the first ratio in the description. shoulders distinct in winged species, re- The aedeagi were examined in glycerin, duced in wingless ones; apical membrane after being boiled for two minutes or put for absent or very narrow, much shorter than a day in a diluted KOH solution and then sutural flange at apex; apical fringe dense rinsed. To study inflated penial sacs, dry pe- and varying from short but distinct (B. chi- nes were first soften in water. Internal penile nensis) to hardly discernable. Elytral pubes- sacs were then everted, inflated to capac- cence moderately long and dense to (usual- ity using a syringe needle inserted into the ly) scarce and short, of two kinds, one dense basal orifice and afterwards dried in a hot- and decumbent, the other erect, very sparse air flow. All measurements were done under on disc, denser along sides and over an ex- the stereoscope using ocular-micrometer tensive posterolateral area. Elytral ridges within the accuracy of two decimal places. mostly weak to obsolescent; interspaces varying from very narrow to much wider TAXONOMY than ridges. Surface dull or feebly shining due to a rather sharp isodiametric micros- Order COLEOPTERA culpture, often combined with a more or Family CARABIDAE less dense punctation and dense strigae. Internal sac of penis with virgae. Subfamily BRACHININAE Bonelli, 1810 Remarks. This group first erected by Genus Brachinus Weber, 1801 Hrdlička (2009b) comprises about 30 de- scribed species widespread in the eastern The melancholicus species group parts of the Oriental Region, with general Diagnosis. Body large, 10.0–21.5 mm in range extending from Myanmar and south- length (may be a little smaller), uniformly ern China south to the Sunda Isles and New dark, head usually with two oblong pale Guinea. Several species have been described spots between eyes, or pale anteriorly; legs or reported from central Chinese provinces pale to dark, often bicolored, with pale fem- as well as from Japan (B. nigridorsis Nakane, ora and more or less infuscate tibiae and tar- 1962). Winged species, for example B. chi-

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM 273 nensis Chaudoir, 1850, are mostly wide- The diagnosis of this incertae sedis group spread. Wingless or brachypterous mem- seems to better fit the subgenus Neobrachi- bers of the group, including B. cychroides nus Erwin, 1970 than any other subgenus. Hrdlička, 2009, B. deuvei Gao & Tian, 2010, However, the apical elytral membrane is B. flores Kirschenhofer, 2011, B. hunanen- very narrow to non-existent in all members sis Kirschenhofer, 2011, B. jakli Hrdlička, of the melancholicus-group, which serves 2009, B. nigridorsis, and B. turnai Hrdlička, the feature to discriminate the African gen- 2009, as well as the species described below era Brachinulus Basilewsky, 1958 and Apti- seem to be local, primarily montane, deriva- noderus Hubenthal,1919 from Brachinus tives of various ancestral species. Weber, 1801 (Erwin, 1970). Clarification Hrdlička (2009b) defined the melan- of the status of the melancholicus-group lies cholicus-group without reference to any beyond the scope of the present paper. subgenus by merely listing its ten member species in addition to the five described Brachinus dalatensis sp. nov. in that paper. He, however, didn’t include (Figs 1, 8, 9) in the list the three species (B. votrubai Hrdlička, 2009, B. jakli Hrdlička, 2009 and Holotype. Male, “S[outh] Vietnam, Lam B. praestans Andrewes, 1931) described Dong Prov., Bi Doup – Nui Ba [Nature] Re- or re-described by him a few pages before serve, 12˚07´ N 107˚25´44´´E, Bi Doup Mt., (Hrdlička, 2009a), even though they clear- N[orthern]. slope, h = 1700–1900 m, 19–22. ly belong to the same group, with no expla- IV.2008, leg. D. Fedorenko”. nation. Furthermore, three species listed Paratypes. Male and 6 females taken together with the holotype, as well as on 10 and 12 Apr. by Hrdlička (2009b) as members of the 2008, and 3 May 2009; same locality, but envi- melancholicus group had been included by rons of Long Lanh, 12˚10´44´´N 108˚40´44´´E, Kirschenhofer (2003) in the orestes-group h = 1400–1600 m, 31 March, 3–6 and 7–8 Apr. of the subgenus Brachinidius Reitter, 1919, 2008, and 4 May 2009, 2 males, 2 females. sunk into the subgenus Brachinolomus Reit- Additional material. Vietnam: Dak Lak ter, 1919 decades before (Erwin, 1970). Prov., Chu Yang Sin National Park, 12˚22´40´´N

Figs 1–3. Brachinus, general appearance: 1, B. dalatensis sp. nov.; 2, B. ovalipennis sp. nov.; 3, B. belyaevae sp. nov.

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 274 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM

108˚21´11´´E, 1.5 km W of Chu Pan Phan Mt., cave; sides rather strongly rounded level h = 1650 m, 30 March – 14 Apr. 2012, leg. D. Fe- to broadest part, sinuate and divergent in dorenko, 1 male (SIEE); Khanh Hoa Prov., Hon front of acute hind angles; front angles rect- Ba Mt., ~1400 m, Apr. 2003, leg. A. Borisenko, 1 angular or slightly acute, with apices sharp male (SIEE). to rounded, slightly, if at all, prominent. Diagnosis. The combination of pale legs, Lateral bead very narrow. Both front and the absence of lateral seta on the pronotum, basal transverse impressions moderately the missing shoulders, the subglabrous ely- deep, the former mostly deeper, sometimes tral disc, and odd elytral ridges being a little obsolete medially; basal foveae wide, long higher than even ones easily differentiates and rather shallow, each with a fine line at this species from the other members of the bottom; these lines converging from base to melancholicus group. basal transverse impression, thence running Description. Body (Fig. 1) 10.0–12.1 mm forward parallel to side margins in basal in length, dark brown; elytra brown to black, 1/2–2/3 of the pronotum; lateral groove scutellum red or dark, appendages and two fairly wide in this basal area and behind oblong spots between eyes reddish-yellow, front angles, while narrow in the second sometimes knees slightly infuscate; often en- fourth. Median line moderately deep, short- tire forehead pale, without or with a darker ly impressed both at the preapical impres- spot at the frontoclypeal border. Microscu- sion and just in front of the basal one, often lpture reticulate, with meshes mostly con- widely sulcate at base. Disc rather smooth, spicuous and isodiametric, slightly trans- finely transversely rugulose and finely punc- verse on neck and sometimes also on each tate between forward extensions of basal side of pronotal median line; elytra with foveae, base and apex often longitudinally slightly longitudinal, squamose reticulation, strigose, coupled with a few rugosities; larg- with meshes up to twice as long as wide. er punctures scarce; pubescence very scarce Eyes convex but small, about as long as to absent at base, scarce to fairly dense in genae. Frons and vertex very convex, fron- anterior half, including lateral groove. Lat- tal sulci moderately deep and irregular, not eral setae reduced or indistinguishable from quite reaching front margin of eye; a shallow pubescence. Propleura slightly tumid, not to (mostly) deep, occasionally absent, V- or or hardly visible in dorsal view. U-shaped furrow just behind frontoclypeal Elytra ovate, convex, 1.27–1.42 (mean suture and sometimes also its mirror image 1.33) times as long as wide, 2.15–2.33 (mean on clypeus; neck constriction reduced to 2.27) times as wide as pronotum, with a a rounded median depression varying be- very narrow base and completely reduced tween small and shallow to fairly large and shoulders, broadest at the apical third, apex deep. Dorsal surface smooth and glabrous truncate or barely concave; apical hairy along the middle, rather densely punctate fringe barely visible. Ridges very weak, odd close to eyes and, especially, on neck (except ones slightly higher than even, some of even its middle); sometimes vertex punctate pos- ridges (mostly 2 and 6) often reduced. Pu- teriorly; neck transversely rugulose, usually bescence sparse and short, middle third of except in the middle. Clypeus and adjacent elytra between suture and ridge 5 usually U-shaped area on frons shining, with weak glabrous or subglabrous. Wings absent. microsculpture. Penis as in Figs 8 and 9, its apical lamella Pronotum cordate, 0.95–1.04 (mean bifid. 1.0, n = 7) times as wide as long, 0.88–1.00 Distribution. Dalat Plateau in southern (mean 0.93) times as wide as head, con- Vietnam (Dak Lak, Lam Dong and Khanh vex, broadest just behind front angles to a Hoa provinces). fourth from apex; base narrower than apex, Etymology. The species name refers to straight to convex, apex straight to con- the type locality, Dalat Plateau.

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM 275

Bionomics. The species inhabits semi- verse impressions mostly shallow, obsolete deciduous, broad-leaved montane tropical medially, the latter deepening laterad; basal forests at 1400–1900 m, sometimes co- foveae shallow to very shallow, with a line occurring with B. ovalipennis sp. nov. and at bottom usually running close to side mar- Pheropsophus bidoupensis sp. nov. gin, lateral groove thus being rather narrow in basal half, narrower than in the preceding Brachinus ovalipennis sp. nov. species. Propleura tumid, well visible from (Figs 2, 10, 11) above. Dorsal surface largely smooth and almost glabrous, pubescence reduced to just Holotype. Male, “S[outh] Vietnam, Lam a few setae. Dong Prov., Bi Doup – Nui Ba [Nature] Reserve, Elytra 1.36–1.47 (mean 1.42) times as env. Long Lanh, 12˚10´44´´N / 108˚40´44´´E, h = long as wide, 2.20–2.47 (mean 2.33) times 1400–1600 m, 1–2.IV.2008, leg. D. Fedorenko”. Paratypes.11 males and 8 females taken to- as wide as pronotum; all ridges subequally gether with holotype, as well as on various dates weak, sometimes almost obliterated, only between 31 March and 11 Apr. 2008, and be- suture elevated at base; glabrous discal tween 27 Apr. and 7 May 2009; same locality, but area tends to be more extensive; only ely- 12˚11´N 108˚42´E, ~4 km SSE of Mt Hon Giao, tral base, apical quarter, and sides laterad of h = 1500–1800 m, 2–3.IV.2008, 2 males. ridge 6 pubescent. Additional material. Vietnam: Khanh Hoa Apical lamella of penis simple (Figs 10, 11). Prov., Mt Hon Ba, ~1400 m, Apr. 2003, leg. Distribution. Dalat Plateau within Lam A. Borisenko, 3 males, 2 females (SIEE). Dong and Khanh Hoa provinces, southern Diagnosis. Very similar to the previous Vietnam. species, but with body, especially prono- Bionomics. Same as in B. dalatensis sp. tum, slightly slenderer; the elytral ridges of nov. equal height; body, and especially the head, Etymology. The species name refers to slightly paler; and the glabrous area on ely- the shape of elytra. tra more extensive. Penis in ventral view widely rounded apically. Brachinus belyaevae sp. nov. Description. Same as previous, except (Fig. 3) as follows: body (Fig. 2) 8.4–12.7 mm in length, slightly paler in general, forehead Holotype. Female, “South Vietnam, ~95 km predominantly pale, occasionally with a SEE of Ho Chi Minh, Xuyen Moc Nature Re- vague darker spot at the border between serve, 15.V.1991, leg. N. Belyaeva”. frons and vertex; sometimes head and pro- Diagnosis. The species distinctly differs notum entirely reddish-yellow except later- from the other species of the group in the ally. U-shaped furrow on frons moderately combination of the lacking shoulders, bicol- deep in general, a convex area between its ored legs and infuscate antennomeres 1–4. arms and clypeus glossy because of very fine No other member of the group has bicolored or obsolete microsculpture. legs combined with reduced shoulders. Pronotum long, 0.81–0.93 (mean 0.86, Description. Habitus as in Fig. 3. Body n = 7) times as wide as long, 0.82–0.93 length 11.8 mm, dorsum black, pronotal disc (mean 0.87) times as wide as head, apex as dark brown; two oblong spots between eyes, wide as or slightly wider than base, sides labrum, anterior 2/3 of clypeus, mouth- very slightly rounded; front angles mostly parts, basal half of scape, antennomeres 5 acute and sharp, occasionally subrectan- to 11, and tarsi reddish; four basal anten- gular or blunt; hind angles sharp, slightly nomeres slightly infuscate, femora yellow, acute to rectangular, base laterad barely their apices and tibiae rather dark brown. emarginate to slightly oblique forwards ac- Scutellum dark. Underside reddish-brown, cordingly. Both preapical and basal trans- paler medially. Dorsum dull because of the

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 276 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM heavy reticulation throughout (a little less Distribution. Known only from the type heavy within a small U-shaped postclypeal locality, Xuyen Moc Nature Reserve near area), with meshes isodiametric on head Binh Chau, southern Vietnam. and pronotum, slightly longitudinal, some- Etymology. Named after Dr. Natalia Be- what squamose, on elytra. lyaeva (Moscow), an Isoptera specialist, Eyes convex, longer than genae; frons and who collected the holotype. vertex very convex, smooth and glabrous, with setigerous punctures sparse and coarse Genus Pheropsophus Solier, 1833 along sides while dense behind of the level of posterior supraorbital setae; neck rugulose, Subgenus Stenaptinus Maindron, 1906 with a small, rounded, median depression. The aptinoides species-group Pronotum cordate, 1.05 times as wide as long, 0.96 times as wide as head, broad- The members of this group from Vietnam est about a fourth from apex; front margin share the following combination of charac- slightly concave, sides strongly rounded ters: Body uniformly dark, antennae and anteriorly and strongly sinuate basad; base legs yellow, femoral apices occasionally in- slightly narrower than apex, nearly straight; fuscate or with a darker spot on each side of front angles subacute, hind angles acute, all knee. Labrum anteriorly truncate to convex with blunted apices; disc convex; preapical (varying between individuals), with front impression shallow, deep only at middle; angles widely rounded. Elytra at base not or basal transverse impression obsolete; median just wider than base of pronotum, without line entire, moderately deep, deeper in basal shoulders, strongly widened backwards and third, wide in basal fourth; lateral bead very broadest just before apex, apical margins narrow, lateral groove narrow, wider at front recessed towards suture at a wide angle, su- angles, very wide and merging with basal tural angles rounded; disc largely glabrous, fovea behind middle; basal foveae deep and very sparsely and almost indistinctly ciliate, long, reaching mid-length, converging from with very short sparse erect setae that tend base to basal transverse impression, thence to be denser posterolaterally; interspace 1 running parallel to each other; each fovea with long dense pubescence throughout, in- anteriorly extended into a shallow line run- terspace 2 with four long setae, interspace ning parallel to lateral margin and ending 5 with one seta in basal 1/4 to 1/3; an ad- near preapical impression. Disc transversely ditional seta may be present in mid-length rugulose, base and apex densely longitudi- of the interval 4 and/or in interval 3 in front nally strigose; setigerous punctures rather of the apex. Elytral ridges, especially outer minute but dense almost throughout; lateral ones, more or less strongly carinate; ridges seta short, twice as long as pubescence. Pro- 2 to 5 flattened and broadened basally, with pleura invisible from above. ridge 4 not reaching base; ridge 8 often, Elytra ovate, 1.37 times as long as wide, and occasionally ridge 2, ending abruptly 2.26 times as wide as pronotum, broadest a and not quite reaching apex. Prosternum third from apex, shoulders reduced, apical rather densely pubescent. Abdominal ter- truncature transverse and straight. Ridges gite VII in female with 13–17 strong setae subequally well-developed, moderately along apical margin. Apical lamella of penis convex, barely separated from and about as large, mostly triangular and pointed (Figs wide as or slightly wider than interspaces, 16–18); interspecific differences in its shape subcostate in basal half, subcarinate in api- being mostly insufficient, if any. Internal sac cal half, ridge 7 a little narrower and higher of penis is rather simple in shape and often than others, ridge 2 almost reduced basally. (Ph. tamdaoensis Kirschenhofer, 2010; Ph. Pubescence short and sparse, scarce in the obliteratus sp. nov. and Ph. kalyakini sp. middle of disc. Wings absent. nov., as well as Ph. beckeri Jedlička, 1930

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM 277 from China) supplied with a long process 8(5) Elytral ridges weak, ridge 7 either obliter- below the apex of left sclerotized lobe of ate or (rarely) subcostate at base. preputial field. 9(10) Pronotum subquadrate, sides faintly Secondary sexual differences are as in rounded in anterior half and slightly sinuate in front of base, both lateral bead and lateral other Stenaptinus: in males the profemora groove wider, the latter broadened behind are often incrassate, and the mesotibia with lateral seta; propleura very sparsely setulose; 24–34 spinules along the anterolateral mar- interspace 4 at mid-length with 10–13 ir- gin, arranged into two or more irregular regular rows of longitudinal carinulae. Penis rows, versus a single row of 14–18 spinules with apical lamella wide and rounded at tip in females. (Fig. 19) ...... Ph. kalyakini sp. nov. Distribution. This group is widespread 10(9) Pronotum cordate, its sides well rounded over the Oriental Region from Nepal and in anterior half and sinuate in front of base, China south to Celebes; however, the mem- lateral bead and lateral groove both very narrow all along; propleura sparsely but ber species tend to have vicariant ranges distinctly punctate-pubescent; interspace 4 confined to a particular montane region – at mid-length with less than nine irregular a pattern for which the apterous condition rows of longitudinal carinulae. Penis with seems to be largely responsible. apical lamella as in Fig. 22 ...... Remarks. This group fits well into Sten- ...... Ph. obliteratus sp. nov. in its original, strict, sense (Main- dron, 1906). The long lateral process on the Pheropsophus bidoupensis sp. nov. left side of the penile internal sac has beeen (Figs 4, 12, 16, 21) found to be characteristic of many consub- geners not only inside, but also outside the Holotype. Male, “S[outh] Vietnam, Lam Dong aptinoides group. These latter species are Prov., Bi Doup – Nui Ba [Nature] Reserve, env. Long Lanh, 12˚10´44´´N 108˚40´44´´E, h = 1400– as follows: Ph. javanus (Dejean, 1825), Ph. 1600 m, 20–21.IV.2008, leg. D. Fedorenko”. picicollis Chaudoir, 1876 and Ph. margini- Paratypes. 15 males and 13 females taken to- collis Motschulsky, 1854. gether with holotype, as well as on 31 March – Diagnoses of the species described be- 21 Apr. 2008 (pitfall traps), 5 May and 10 May low are provided in the key couplets. 2009; same locality, but 12˚11´N 108˚42´E, ~4 km SSE of Hon Giao Mt, h = 1500–1800 m, 7–8 Apr. 2008, 1 male. Key to Vietnam species of the Pheropsophus Additional material. Vietnam: Dak Lak aptinoides group Prov., Chu Yang Sin National Park, 12˚22´40´´N 108˚21´11´´E, 1.5 km W of Chu Pan Phan Mt., h 1(4) Neck almost or completely smooth. = 1650 m, 30 March – 11 Apr. 2012, leg. D. Fe- 2(3) Elytral ridges weak, subconvex (North dorenko, 7 males, 5 females (SIEE); same data, Vietnam) ...... Ph. tamdaoensis but leg. A. Abramov, 5 males, 4 females (Zoologi- 3(2) Elytral ridges prominent, outer ones sharp- cal Institute, Russian Academy of Sciences, St ly carinate; carinulate sculpture obsolete be- Petersburg) 5 km S of Dinh Kwo, 19–21 Apr. fore bases of ridges 3 to 5 (South Vietnam) . . 2010, leg. P. Udovichenko, 2 females; Khanh Hoa ...... Ph. bidoupensis sp. nov. Prov., Hon Ba Mt, ~1400 m, Apr. 2003, leg. A. 4(1) Neck rugose. Borisenko, 7 males, 5 females (SIEE). 5(8) Elytral ridges prominent, ridge 7 sharply Diagnosis. Similar to P. tamdaoensis sp. carinate and reaching base. nov. in appearance, but distinguishable in 6(7) Pronotum broadest at about a fourth from having much stronger elytral ridges and apex, nearly parallel-sided basally ...... Ph. ngoclinhensis sp. nov. acute hind pronotal angles. The two species 7(6) Pronotum broadest at about a third from are allopatric, with ranges widely separated. apex, sides more deeply sinuate in front of, Description. Body (Fig. 4) length 10.5– and divergent towards, hind angles ...... 14.5 mm. Head reddish-yellow, with a large, ...... Ph. similis sp. nov. subquadrate, sometimes vague, dark brown

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Figs 4–6. Pheropsophus, general appearance: 4, Ph. bidoupensis sp. nov.; 5, Ph. ngoclinhensis sp. nov.; 4, Ph. similis sp. nov. spot on vertex, fused to or barely separated with two to six (up to ten) shallow setiger- from the dark brown area behind eyes; the ous punctures along front margin and occa- spot is truncate or bifid anteriorly; prono- sionally 2–4 at base. Propleura impunctate. tum dark brown, often reddish at apex and Elytra 1.11–1.30 (mean 1.22) times as base; elytra brown to dark brown, densely long as wide, 1.91–2.06 (mean 1.96) times as speckled, with darker ridges. wide as pronotum, ridges rather strong, ridg- Head smooth, neck rugosities obliterate es 6 to 8 (especially 7 and 8) sharply carinate, or almost so, with several (2–7) setigerous much sharper towards base. Disc glabrous, punctures behind and sometimes also in carinulate sculpture fine, obsolete towards front of supraorbital seta; a small longitu- and around bases of ridges 3 to 5, carinulae dinal median depression at the border with being rather short and mostly forming 10–12 pronotum. Eyes flattened, but still rather (occasionally as few as seven) longitudinal convex. rows in interspace 4 at mid-length. Pronotum cordate, 0.89–1.02 (mean Penis (Figs 12, 16, 21). 0.96, n = 9) times as wide as long, 1.0–1.14 Variability. In individuals from northern (mean 1.05) times as wide as head, apex populations (Chu Yang Sin National Park), somewhat bisinuate, base slightly narrow- the pronotum is slightly broader, on average er than apex, tends to be slightly concave; 1.09 times as wide as long, with hind angles sides moderately rounded in front, sinu- somewhat lobed and protruding a little be- ate-divergent behind; both front and hind yond the straight base. angles subacute, blunted at apices. Lateral Distribution. Dalat plateau within Dak bead extremely fine, lateral groove narrow; Lak, Lam Dong and Khanh Hoa provinces, both preapical and basal transverse depres- Vietnam. sions indistinct, with a shallow median line Etymology. The species name refers to in between; surface smooth and glabrous, the Bi Doup location.

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Bionomics. The type series and the speci- Pheropsophus kalyakini sp. nov. mens collected by the author were taken by (Figs 14, 19) pitfall trapping or by hand in broad-leaved forests at 1400–1700 m, often near streams Holotype. Male, “Central Vietnam, Ha Tinh Prov., Vu Quang Nature reserve, h = 1200 m, in their upper flows. 5–26.VIII.1997, leg. M. Kalyakin”. Paratypes. 5 males and 5 females taken to- Pheropsophus ngoclinhensis sp. nov. gether with holotype, plus 1 female from the (Figs 5, 13, 17) same location but h = 400–700 m, 27 Aug. – 4 Oct. 1997. Holotype. Male, “Central Vietnam, Kon Tum Description. As compared to Ph. bi- Province, N[orthern] part., Ngoc Linh Nature doupensis sp. nov.: Body length 13.6–15.6 Reserve, Ngoc Linh env., pitfall traps, V.2005, mm. Dorsum dark, almost black, elytral leg. A. Anichkin”. Paratypes. 2 males and 4 females taken to- speckles very faint; pale color on head con- gether with the holotype. fined to median or anterior part of frons. Description. Same as above, except as Eyes flat to moderately convex; frontal fo- follows: body (Fig. 5) 12.3–14.4 mm in veae rather deep and deepening anterad; length, mostly brown, rather pale; vertex fine longitudinal striations between eye and with a dark, often vague, spot with trun- a smooth or finely rugulose frontal fovea; cate anterior margin; either apical mar- punctures behind supraorbital setae few; gins of elytra edged with pale or extreme neck rugose. apices of elytral ridges pale, yellow. Eyes Pronotum rectangular to subcordate, hardly more convex, neck rugose, without 0.82–0.88 (mean 0.84, n = 5) times as wide as depression, frontal foveae in posterior half long, 0.96–1.04 (mean 0.99) times as wide as densely and finely rugulose, punctures be- head, sides faintly rounded in anterior half, hind posterior supraorbital setae rather nu- only slightly sinuate in front of base; front merous. Pronotum barely shorter, cordate, angles blunt, hind angles acute to rectangu- 1.0–1.04 (mean 1.02, n = 5) times as wide as lar, with apices blunt. Lateral groove narrow long, 1.02–1.07 (mean 1.06) times as wide anteriorly and posteriorly, rather wide, but as head, base conspicuously narrower than shallow medially or behind lateral setiger- apex; front angles obtuse, hind angles very ous pore. Surface smooth and glabrous, base slightly acute, almost rectangular; sides and apex each with 4–14 small and shallow well rounded anteriorly, sinuate and paral- setigerous punctures. Propleura finely and lel behind, broadest at about a fourth from very sparsely punctate-pubescent. apex. Elytra 1.26–1.32 (mean 1.28) times as Elytra 1.32–1.49 (mean 1.41, n = 3) long as wide, 1.84–1.91 (mean 1.88) times times as long as wide, 1.92–2.40 (mean as wide as pronotum, elytral ridges as in the 2.12) times as wide as pronotum; ridges previous species, ridges 2 to 5 mostly much moderately convex, tend to fade basad, less strongly flattened basally, ridge 6 oc- ridges 4 and/or 6 flattened or obliterate at casionally widely interrupted just behind base, ridge 7 never sharply carinate basally base. Carinulate microsculpture fine, very and usually not quite reaching base; pubes- sparse at base; interspace 4 at mid-length cence sparse on disc, carinulae forming 10– with about ten rows of carinulae. 13 rows in interspace 4 at middle of elytron, Penis as in Figs 13, 17. sculpture extending to elytral base and Distribution. Known only from the type turning there from carinulate into aciculate. locality, Ngoc Linh Nature Reserve east of Apical lamella of penis large and round- Dak Glei, Kon Tum Province, central Viet- ed at tip (Figs 14, 19). nam. Distribution. Known only from the type Etymology. The species name refers to locality, Vu Quang Nature Reserve, Ha the type locality. Tinh Province, Vietnam.

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 280 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM

Etymology. Named after Dr Mikhail Ka- 1.4 times as long as wide, 1.92 times as wide lyakin (Moscow), a reputed ornithologist, as pronotum, ridges entire, carinate, sharply my friend and companion during several carinate laterally, ridges 6 and 7 reach ely- expeditions to Vietnam. tral base and sharply carinate there. Cari- Remarks. Pinned after a long preserva- nulate sculpture reaches elytral base. Pe- tion in alcohol, many specimens of the type nile lamella narrow, with apex pointed, and series show integument darkened to black smaller (Figs 15, 18). and elytra deformed/twisted, which may Distribution. Same as in the previous have affected the accuracy of measure- species. ments. Several specimens, mostly females, Remarks. In the holotype, both lateral show rather robust heads. pronotal setae are abraded, which suggests wear that might have also caused the lost of Pheropsophus similis sp. nov. vestiture on the disc, thus making the latter (Figs 6, 15, 18) look glabrous. Etymology. The species name reflects Holotype. Male, “Central Vietnam, Ha Tinh great similarity in appearance between this Prov., Vu Quang Nature Reserve, h = 400– species and the sympatric/syntopic Ph. 700 m, 27.VIII–4.X.1997, leg. M. Kalyakin”. kalyakini sp. nov. Description. Hardly separable from the previous species based on external charac- Pheropsophus obliteratus sp. nov. ters. Body (Fig. 6) 14.2 mm in length. Head (Figs 7, 20, 22) pale before eyes, with pale area reaching mid-eye level medially. Eyes moderately Holotype. Male, “S[outh] Vietnam, Dak Lak convex. Pronotum cordate and shorter, 0.93 province, Chu Yang Sin Natn. Park, 12˚23´48´´N times as wide as long, 1.09 times as wide as 108˚20´59´´E, upper flow of Krong Kmar river, head; disc and propleura glabrous. Elytra h = 1000 m, 30.III–14.IV.2012, leg. D. Fedo- renko”. Description. Body (Fig. 7) 12.6 mm in length, dark brown, almost black, ap- pendages, labrum, anterior half of clypeus, supra-antennal plates and a fairly small, transverse spot between anterior halves of eyes reddish-yellow; frons, base of clypeus, and mandibular apices infuscate to brown, rather dark, elytra dark brown, with a hint of darker speckles. Eyes convex. Neck widely rugose, frons and vertex widely smooth along middle, with rather numerous but fine setigerous punctures and very fine longitudinal stria- tions laterally; frontal foveae moderately deep, deeper at clypeus, with shallow U- shaped depression around pale spot on frons. Pronotum cordate, 0.98 times as wide as long, 1.06 times as wide as head, broadest at the apical third, base about as wide as apex, much narrower than maximum width, base Fig. 7. Pheropsophus obliteratus sp. nov., general slightly concave, apex subtruncate, sides appearance. rather strongly rounded anteriorly, sinuate

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Figs 8–11. Brachinus, penis: 8, 9, B. dalatensis sp. nov.; 10, 11, B. ovalipennis sp. nov. Left lateral (8, 10) and posteroventral (9, 11) aspects. Scale bar 1 mm. and barely divergent in front of base, front interspaces between them; ridges 7 and 8 angles subrectangular, hind angles sub- carinate in apical two thirds, not reach- acute, all prominent. Preapical and basal ing base and somewhat bent in basal third, transverse depressions shallow; pronotum where ridges 4 and 6 are fading or interrupt- smooth, with several small setigerous punc- ed. Longitudinal carinulae in interspace tures on each side of base as well as along 4 forming less than nine rows, carinulate the apical and side margins in anterior half. sculpture turning into aciculate one at the Propleura sparsely punctate-pubescent. scutellar level. Elytra 1.26 times as wide as long, 1.85 Penis (Figs 20, 22) with apical lamella times as wide as pronotum. Elytral ridges large and widely emarginate on right side, slightly convex and rather wide, ridges 3 its tip in dorsal view narrow, parallel-sided to 5 at mid-length about twice as wide as and apically rounded. Inflated endophallus

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 282 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM

Figs 12–15. Pheropsophus, penis with right paramere, right lateral aspect: 12, Ph. bidoupensis sp. nov.; 13, Ph. ngoclinhensis sp. nov.; 14, Ph. kalyakini sp. nov.; 15, Ph. similis sp. nov. Scale bar 1 mm.

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Figs 16–19. Pheropsophus, apex of penis, ventral aspect: 16, Ph. bidoupensis sp. nov.; 17, Ph. ngoclin- hensis sp. nov.; 18, Ph. similis sp. nov.; 19, Ph. kalyakini sp. nov. Scale bar 1 mm.

Figs 20–22. Pheropso- phus, inflated internal sac of penis: 20, 22, Ph. obliteratus sp. nov.; 21, Ph. bidoupensis sp. nov. Left lateral (20, 21) and posteroventral (22) aspects. Scale bar: 1 mm.

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(2): 271–284 284 D.N. FEDORENKO. NEW SPECIES OF BOMBARDIER BEETLES FROM VIETNAM with a long lateral process below the apex of Six new species and one redescription of ge- left sclerotized lobe of preputial field. nus Brachinus from S.E. Palaearctic and Ori- Distribution. Known from the type local- ental region. Studies and reports of District ity only. Museum Prague-East, Taxonomical Series, 5(1–2): 103–114. Etymology. The species name refers to Jedlička A. 1964. Monographie der Truncati- elytral striae weakening or obliterated to- pennen aus Ostasien. Lebiinae – Odacanthi- wards base. nae – Brachyninae (Coleoptera, Carabidae). Bionomics. The holotype was taken Entomologisсhe Abhandlungen und Berichte by hand near a tiny stream flowing into a aus dem Staatliches Museum für Tierkunde in tributary of Krong Kmar River in a tropical Dresden, 28(7) [1962–1964]: 269–579. monsoon broadleaf forest. Kirschenhofer E. 1986. Neue Arten truncati- penner Carabidae der palaearktischen und orientalischen Region unter besonderer Be- AKNOWLEDGEMENTS rücksichtigung der Aufsammlungen Eigin Suenson in Ostasien (Coleoptera Carabi- I am greatly obliged to Vasily V. Belov (Ma- dae). Entomofauna, 7(23): 317–346. plewood, USA) for reviewing the manuscript Kirschenhofer E. 2003. Neue und wenig be- and a linguistic revision of the text. kannte truncatipenne Carabidae aus Süd- und Südost Asien aus dem Ungarischen REFERENCES Naturwissenschaftlichen Museum Budapest (Coleoptera: Carabidae, Brachinini, Lebi- Erwin T.L. 1970. A reclassification of Bombar- ini). Annales Historico-Naturales Musei Na- dier beetles and a taxonomic revision of the tionalis Hungarici, 95: 5–18. North and Middle American species (Cara- Kirschenhofer E. 2010. New and little-known bidae: Brachinida). Quaestiones Entomologi- species of Carabidae from the Middle East cae, 6: 4–215. and Southeast Asia (Coleoptera, Carabi- Gao Q. & Tian M. 2010. Two new species of the dae: Lebiini, Brachinini). Annales Historico- genus Brachinus Weber, 1801, from China Naturales Musei Nationalis Hungarici, 102: [Coleoptera, Carabidae, Brachinini]. Revue 1–40. française d’Entomologie (N.S.), 32(1–2): Kirschenhofer E. 2011. Neue Arten der Gat- 97–100. tungen Craspedophorus Hope, 1838, Brachi- Habu A. 1967. Fauna Japonica. Carabidae, nus Weber, 1801, Trigonotoma Dejean, 1828, Truncatipennes group (Insecta: Coleoptera). faunistische Meldungen der gattung Chla- Tokyo: Biogeographical Society of Japan, enius Bonelli, 1810 aus der Orientalis. Acta Tokyo Electrical Engineering College Press. Coleopterologica, 27(1): 38–60. xiv+338 p.+xxv pl. Maindron M. 1906. Description d’une nouvelle Hrdlička J. 2009a. Contribution to the tribe espece de Pheropsophus et remarques sur ce Brachinini (Coleoptera: Carabidae) – II. genre. Bulletin de la Société Entomologique de Two new species and one redescription of France: 15–16. genus Brachinus from Indonesia. Studies and Tian M. & Deuve T. 2007. Review of four spe- reports of District Museum Prague-East, Tax- cies of the genus Brachinus Weber (Cole- onomical Series, 5(1–2): 95–102. optera: Caraboidea: Brachinidae) from the Hrdlička J. 2009b. Contribution to the tribe Philippines. Zootaxa, 1546: 15–22. Brachinini (Coleoptera: Carabidae) – III. Received May 29, 2013 / Accepted November 27, 2013

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