ValimYeKavanaughDavid H.ShiHongliangLiangHongbinA key to species of subgenus Lithochlaenius (Coleoptera, Carabidae, Chlaeniini, Chlaenius), with descriptions of three new speciesZookeys9920112011128155210.3897/zookeys.128.1804 Lithochlaenius http://species-id.net/wiki/Lithochlaenius Kryzhanovskij, 1976Hemichlaenius Lutshnik, 1933:169 (nec Bates, 1892:307). Type species Chlaenius rambouseki Lutshnik, 1933; Kryzhanovskij, 1976:11Lithochlaenius Kryzhanovskij, 1976:9. Type species Chlaenius rambouseki Lutshnik, 1933; Morita, 1993:161Agilochlaenius Kirschenhofer, 1997:116. Type species Chlaenius latro LaFerté–Sénectère, 1851; Kirschenhofer, 2000:58Diagnosis.

Antennomere 3 distinctly longer than antennomeres 1 and 2 combined (Fig. 9); pronotum distinctly cordate (Fig. 1), disk glabrous or sparsely pubescent, each hind angle with single seta (Fig. 2); scutellar setiferous pore puncture present (Fig. 3); venter densely pubescent laterally, sparsely pubescent or glabrous medially (Fig. 4); prosternal intercoxal process punctate, bordered at apex; metepisterna long and narrow (Fig. 10); basal margination of elytra incomplete, absent from medial portion; anterior tarsomere 4 short and deeply emarginate apically (more distinctly so in male, Figs 5–8); aedeagus tubular, ejaculatory orifice long, extended to basal fifth of aedeagus, lamella short (Figs 11, 117–145); gonostylus of female ovipositor smooth, with one setiferous pore near apex and one slender spine at basal inner margin (Figs 12–15).

Description.

Length 12.0–18.0 mm, width 4.6–6.4 mm. Head and pronotum black, with green, blue or coppery metallic luster; elytra black, with blue or coppery luster in a few species; ventral surface black; legs black, brown or yellow; antennae yellow, brown, or dark brown.

Head with vertex nearly glabrous, or sparsely and coarsely punctate behind posterior level of eye; eyes moderately prominent; genae pubescent; antennae long, antennomere 1 (scape) coniform (Figs 52–53, 57–59), cylindrical (Figs 56, 60), or elongate-ovoid (Figs 54–55, 61–65); antennomere 3 sparsely setose, distinctly longer than antennomeres 1 and 2 combined (Fig. 9); mandibles triangular, hooked at apex; labrum with six setigerous punctures near apex; maxillary and labial palpi cylindrical, slightly compressed at apex; maxillary palpi glabrous; penultimate labial palpomere with a few setae, apical labial palpomere glabrous; glossal sclerite with two subapical setae; tooth of mentum bifid or emarginate at apex, with one pair of setae near the base; gula glabrous, slightly rugose.

Pronotum cordate (Fig. 1), widest at apical one-third, front angles obtuse, hind angles acute; lateral margins bordered; disk glabrous or sparsely pubescent, base longitudinally rugose, punctate; basal foveae small, deep, rugose, pubescent; hind angles each with one seta (Fig. 2).

Elytra moderately convex, oblong, with distinct isodiametric microsculpture at least laterally; scutellar striae long, with basal setiferous pore present (Fig. 3); at least outer intervals pubescent (Figs 80–93), pubescence on intervals 8 and 9 generally denser than on others; wings full-sized, functional.

Abdominal sterna densely pubescent laterally, sparsely pubescent or glabrous medially (Fig. 4); sterna IV to VI with single long seta at each side; sternum VII with one seta at each side in male, two in female; apex of sternum VII more rounded in male than in female.

Pro- and mesosterna and pro-, mes-, and metepisterna densely pubescent; metasternum pubescent laterally, nearly glabrous medially; prosternal intercoxal process punctate and bordered at apex; scutellum triangular, glabrous.

Anterior femora without tooth; tarsomeres short, sparsely setose dorsally; anterior tarsomere 4 short and triangular, deeply emarginate (more so in male), with two rows of long setae ventrally (Figs 5–8); basal three anterior tarsomeres dilated in male.

Male genitalia with aedeagus tubular, simple; ejaculatory orifice long (Fig. 11); apical lamella short, rounded or slightly truncated at apex. Gonostylus of female ovipositor smooth, with one setiferous pore near apex and a long slender spine at basal inner margin, outer margin without or with a very short spine (Figs 12–15).

Character states of Chlaenius (Lithochlaenius) spp. Figs 1–13 Chlaenius rambouseki Lutshnik1 pronotum2 site of hind angle seta3 elytral base showing the site of basal pore4 abdominal sterna showing the pubescence5 male anterior tarsomere 4 in dorsal view6 male anterior tarsomere 4 in ventral view7 female anterior tarsomere 4 in dorsal view8 female anterior tarsomere 4 in ventral view9 antennomeres 1–5 showing antennomere 3 distinctly longer than 1 and 2 ones combined10 metepisternum11 aedeagus, showing the basic structure of the aedeagus in Lithochlaenius species12 female gonostylus in ventral view13 female gonostylus in lateral view14 female gonostylus of Chlaenius propeagilis sp. n. in ventral view15 female gonostylus of Chlaenius propeagilis sp. n. in lateral view.

Habitus of Chlaenius (Lithochlaenius) spp16 Chlaenius chuanqianensis sp. n., holotype, male, dorsal view17 Chlaenius chuanqianensis sp. n., holotype, male, ventral view18 Chlaenius linwensini sp. n., holotype, male, dorsal view19 Chlaenius linwensini sp.n., holotype, male, ventral view20 Chlaenius propeagilis sp. n., holotype, male, dorsal view21 Chlaenius propeagilis sp. n., holotype, male, ventral view. Scale line = 5.0 mm.

Habitus ofChlaenius (Lithochlaenius) spp22 Chlaenius agilis Chaudoir, holotype, dorsal view23 Chlaenius agilis Chaudoir, holotype, ventral view24 Chlaenius formosensis Lorenz, male, in CCCC, dorsal view25 Chlaenius formosensis Lorenz, ventral view26 Chlaenius leishanensis Kirschenhofer, male, in IZCAS, dorsal view27 Chlaenius leishanensis Kirschenhofer, male, in IZCAS, ventral view. Scale line = 5.0 mm.

Chlaenius (Lithochlaenius) spp28 Chlaenius noguchii Bates, male, in IZCAS, dorsal view29 Chlaenius noguchii Bates, male, in IZCAS, ventral view30Chlaenius wrasei Kirschenhofer, male, in IZCAS, dorsal view31 Chlaenius wrasei Kirschenhofer, male, in IZCAS, ventral view32 Chlaenius agiloides Jedlička, male, in IZCAS, dorsal view33 Chlaenius agiloides Jedlička, male, in IZCAS, ventral view. Scale line = 5.0 mm.

Chlaenius (Lithochlaenius) spp34 Chlaenius rambouseki Lutshnik, male, collected from Ussuri region, in IZCAS, dorsal view35 Chlaenius rambouseki Lutshnik, male, collected from Ussuri region, in IZCAS, ventral view36 Chlaenius anchomenoides Bates, cotype, male, in BMNH, dorsal view37 Chlaenius anchomenoides Bates, cotype, male, in BMNH, ventral view38 Chlaenius nuristanus Jedlička, paratype, male, in MNHN, dorsal view39 Chlaenius nuristanus Jedlička, paratype, male, in MNHN, ventral view. Scale line = 5.0 mm.

Chlaenius (Lithochlaenius) spp40 Chlaenius nuristanus a. rubridipes Jedlička, paratype, male, in MNHN, dorsal view41 Chlaenius nuristanus a. rubridipes Jedlička, paratype, male, in MNHN, ventral view42 Chlaenius agiloides Jedlička, holotype, male, in NMPC, dorsal view43Chlaenius formosanus Jedlička, holotype, male, in NMPC, dorsal view44 Chlaenius rambouseki Lutshnik, from Taiwan, male, in IZCAS, dorsal view45 Chlaenius rambouseki Lutshnik, from Taiwan, male, in IZCAS, ventral view. Scale line = 5.0 mm.

Geographical Distribution.

China (Heilongjiang, Liaoning, Shaanxi, Gansu, Henan, Hubei, Hunan, Zhejiang, Fujian, Taiwan, Jiangxi, Guangdong, Hainan, Guangxi, Guizhou, Sichuan, Yunnan, Tibet), North Korea, Japan, Russia (Far East), Afghanistan, Pakistan, India. The known localities of Lithochlaenius species are shown in Fig. 158. Based on the work of Andrewes (1930) and Paik et al. (2006), members of this subgenus also occur in Indo-China (Laos, Cambodia, Vietnam), but we have not studied specimens from those countries.

Biology.

Members of this subgenus are typically collected on sandy beaches of rivers or streams (Figs 146–153). Adults of some species (e. g., Chlaenius agiloides, Chlaenius rambouseki) have been observed feeding on mollusks, worms, and dragonfly larvae at night (Figs 154–157). A few species have also been collected in light traps.

Remarks.

Based on the metallic body surface, single supraorbital seta, antennomere 3 longest and antennomeres 4–11 densely pubescent, Lithochlaenius species can be recognized as a member of the genus Chlaenius. Members of this subgenus are similar to those of subgenus Stenochlaenius in shape of the pronotum, but the latter are much smaller and have a glabrous body surface.

As presently conceived, the subgenus can be divided into two species groups: 1) the rambouseki group, members of which have all elytral intervals densely and more or less equally pubescent, and males have aedeagi slender in dorsal view; and 2) the agilis group, members of which have at least elytral intervals 1–5 glabrous medially with pubescence restricted to the strial depressions, intervals 6–9 densely pubescent, and males have aedeagi generally stouter in dorsal view. Most species of the subgenus have restricted geographical ranges, and only C. rambousekiis more broadly distributed.

In his treatment of the North American species of genus Chlaenius, Bell (1960) suggested that his solitarius species group, which included Chlaenius cordicollis Kirby, Chlaenius leucoscelis Chevrolat, Chlaenius prasinus Dejean, Chlaenius purpureus Chaudoir, and Chlaenius solitarius Say, represented a distinct group within his subgenus Chlaenius sensu stricto. He noted that the range of this group extends south into South America and that "Related forms occur in the Old World”. In the paper in which he proposed Lithochlaenius as a replacement name for Hemichlaenius Lutshnik, Kryzhanovskij (1976: 16) cited Bell's paper and suggested that Chlaenius solitarius might be a North American representative of that subgenus. Robert Davidson (personal communication) shares the view that species of Bell's solitarius group, and at least five additional species in Middle and South America, are likely related to some if not all Lithochlaenius species. All of these New World species share most of the diagnostic features of Lithochlaeniusand all of them have elytral pubescence as seen in members of the rambouseki group. However, members of all these New World species are distinguished in having the lateral and basal elytral margins smoothly continuous around the humeri without forming any trace of an angle and the elytral epipleura and basal regions smoothly continuous around the humeri, not separated by a carina of any kind. These were the main features that Bell used to distinguish members of his solitarius group. In contrast, members of all the Asian species of Lithochlaenius that we have studied have a distinct humeral angle formed at the junction of the lateral and basal elytral margins and a more or less distinctly carinate separation of the (lateral) epipleural from the basal (anterior vertical) elytral surfaces. Hence, Asian Lithochlaenius specimens could not be identified as members of the solitariusgroups using Bell's (1960) key. There is also greater variation in the development of the elytral basal margin (from complete to partially interrupted) and in the length and shape of the apical lamella of the male aedeagus among New World species than we have seen among the Asian Lithochlaeniusspecies. Consequently, we suggest that placement of any New World species in subgenus Lithochlaenius would be premature at this time and should await a more comprehensive, worldwide treatment of genus Chlaenius and analyses of phylogenetic relationships among the included species, species groups, and subgenera.

Key to the species of subgenus Lithochlaenius
1All intervals punctate, pubescent, slightly convex (Figs 88–89, 93); aedeagus slender, basal portion rugose (rambouseki group)2
–At least basal half of intervals 1–5 smooth and glabrous medially, distinctively convex (Figs 80–87, 90–92); aedeagus usually stout, basal portion smooth (agilis group)3
2Intervals dull, with dense and regular punctures (Figs 89, 93); antennomere 1 elongate ovoid (Figs 61, 65); apical lamella of aedeagus thin (Figs 137, 145)Chlaenius rambouseki Lutshnik
–Intervals shining, with sparse and irregular punctures (large and small punctures mixed) (Fig. 88); antennomere 1 cylindrical (Fig. 60); apical lamella of aedeagus thick (Figs 134–135)Chlaenius agiloides Jedlička
3Antennomere 1 with apical end much thicker than basal end, coniform (Figs 52–53, 57–59)4
–Antennomere 1 with apical end as thick as basal end, elongate-ovoid (Figs 54, 55, 62–64) or cylindrical (Fig. 56)8
4Intervals 1–7 glabrous medially, with a row of pubescence laterally (near striae) (Fig. 86); legs dark brown or brown (Fig. 29); apex of sternum VII subtruncate (Figs 105, 106); aedeagus slender, depressed, apical lamella bent ventrally (Figs 129–130)Chlaenius noguchii Bates
–Intervals 6–7 pubescent, intervals 1–5 glabrous medially, with a row of pubescence laterally (near striae) (Figs 80–81, 85, 87); aedeagus stout5
5Tibiae and femora bi-colored, with tibiae black or dark brown (Figs 26–27), femora yellow or light brown; lamella of aedeagus rounded at apex (Fig. 127), thickened, and bent ventrally (Fig. 128)Chlaenius leishanensis Kirschenhofer
–Tibiae and femora concolorous, yellow or brown (Figs 16–19, 30–31)6
6All antennomeres brown or dark brown (Fig. 18); hind trochanters brown, nearly the same color as hind femora (Fig. 19); elytra with basal three-fourths of intervals 1–5 glabrous medially; lamella of aedeagus sub-truncate at apex (Fig. 119), thick (Fig. 120)Chlaenius linwensini Liu & Liang, sp. n.
–At least antennomere 1 yellow or brown, paler than the rest (Figs 16, 30); hind trochanters much darker than hind femora (Figs 17, 41); elytra with entire lengths of intervals 1–5 glabrous medially; lamella of aedeagus rounded at apex (Figs 117, 131), thin (Figs 118, 132)7
7Antennomeres 1–3 brown, color paler than antennomeres 4–11 (Fig. 30); apical lamella of aedeagus bent ventrally (Fig. 132)Chlaenius wrasei Kirschenhofer
–Only antennomere 1 yellow or brown, color paler than antennomeres 2–11 (Fig. 16); apical lamella of aedeagus straight (Fig. 118)Chlaenius chuanqianensis Liu & Liang, sp. n.
8Intervals 1–7 glabrous medially (Fig. 84); antennomere 1 cylindrical (Fig. 56); lamella of aedeagus subtruncate at apex (Fig. 125)Chlaenius formosensis Lorenz
–At most intervals 1–5 glabrous medially (Figs 82–83, 90–92); antennomere 1 elongate ovoid (Figs 54–55, 62–64); lamella of aedeagus round at apex (Figs 121, 123, 138, 140, 142)9
9Male with media lobe of aedeagus gradually bent near base (Fig. 122), lamella triangular at apex, left side of media lobe nearly straight in dorsal view (Fig. 121)Chlaenius propeagilis Liu & Kavanaugh, sp. n.
–Male with media lobe of aedeagus abruptly bent near the base (Figs 124, 139, 141, 143), lamella rounded at apex, left side of media lobe expand laterally in dorsal view (Figs 123, 138, 140, 142)Chlaenius agilis Chaudoir
Theagilisgroup

Aedeagi of Chlaenius (Lithochlaenius) species117 Chlaenius chuanqianensis sp. n., holotype (dorsal view)118 Chlaenius chuanqianensis sp. n., holotype (left lateral view)119 Chlaenius linwensini sp. n., holotype (dorsal view)120 Chlaenius linwensini sp. n., holotype (left lateral view)121 Chlaenius propeagilis sp. n., holotype (dorsal view)122 Chlaenius propeagilis sp. n., holotype (left lateral view)123 Chlaenius agilis Chaudoir, holotype (dorsal view)124 Chlaenius agilis Chaudoir, holotype (left lateral view)125 Chlaenius formosensis Lorenz, holotype (dorsal view)126 Chlaenius formosensis Lorenz, holotype (left lateral view)127 Chlaenius leishanensis Kirschenhofer (dorsal view)128 Chlaenius leishanensis Kirschenhofer (left lateral view)129 Chlaenius noguchiiBates (dorsal view)130 Chlaenius noguchiiBates (left lateral view). Scale lines: A = 1.0 mm (Figs 117, 119, 121, 123, 125, 127, 129); B = 0.5 mm (Figs 118, 120, 122, 124, 126, 128, 130).

Aedeagi of Chlaenius (Lithochlaenius) species 131 Chlaenius wrasei Kirschenhofer (dorsal view) 132 Chlaenius wrasei Kirschenhofer (left lateral view)133 Chlaenius agiloides Jedlička (dorsal view) 134 Chlaenius agiloides Jedlička, holotype (left lateral view) 135 Chlaenius agiloides Jedlička (left lateral view) 136 Chlaenius rambouseki Lutshnik from Ussuri region (dorsal view)137 Chlaenius rambouseki Lutshnik from Ussuri region (left lateral view) 138 Chlaenius anchomenoides Bates, paratype (dorsal view) 139 Chlaenius anchomenoides Bates, paratype (left lateral view)140 Chlaenius nuristanus Jedlička, paratype (dorsal view, black leg) 141 Chlaenius nuristanus Jedlička, paratype (left lateral view) 142 Chlaenius nuristanus Jedlička, paratype (dorsal view, yellow leg)143 Chlaenius nuristanus Jedlička, paratype (left lateral view)144 Chlaenius rambousekiLutshnik from Taiwan (dorsal view)145 Chlaenius rambousekiLutshnik from Taiwan (left lateral view). Scale lines: A = 1.0 mm (Figs 131, 133, 136, 138, 140, 142, 144); B = 0.5 mm (Figs 132, 134, 135, 137, 139, 141, 143, 145).

Antennomere 1 of Chlaenius (Lithochlaenius) species52 Chlaenius chuanqianensis sp. n., holotype53 Chlaenius linwensini sp. n., holotype54 Chlaenius propeagilis sp. n., holotype55 Chlaenius agilis Chaudoir, holotype56 Chlaenius formosensis Lorenz, CCCC57 Chlaenius leishanensis Kirschenhofer, IZCAS58 Chlaenius noguchii Bates, IZCAS59 Chlaenius wrasei Kirschenhofer, IZCAS60 Chlaenius agiloides Jedlička, IZCAS61 Chlaenius rambouseki Lutshnik from Ussuri region, IZCAS62 Chlaenius anchomenoides Bates, paratype63 Chlaenius nuristanus Jedlička, paratype64 Chlaenius nuristanus a. rubridipes Jedlička, paratype65 Chlaenius rambouseki Lutshnik from Taiwan,CCCC. Scale line = 1.0 mm.

Elytral intervals of Chlaenius (Lithochlaenius) spp. 80 Chlaenius chuanqianensis sp. n., holotype81 Chlaenius linwensini sp. n., holotype82 Chlaenius propeagilis sp. n., holotype83 Chlaenius agilis Chaudoir, holotype84 Chlaenius formosensis Lorenz, CCCC85 Chlaenius leishanensis, IZCAS86 Chlaenius noguchii Bates, IZCAS87 Chlaenius wrasei Kirschenhofer, IZCAS88 Chlaenius agiloides Jedlička, IZCAS89 Chlaenius rambouseki Lutshnik from Ussuri region, IZCAS90 Chlaenius anchomenoides Bates, paratype91 Chlaenius nuristanusJedlička, paratype92 Chlaenius nuristanus Jedlička, paratype93 Chlaenius rambouseki Lutshnik from Taiwan,CCCC. Scale line = 1.0 mm.

Map showing the known geographical distributions of Chlaenius (Lithochlaenius)spp.

Photographs of habitats for Chlaenius (Lithochlaenius) spp. 146 Chlaenius chuanqianensis sp. n., Dabaitang, Xishui county, North Guizhou, China147 Chlaenius anchomenoides Bates, Goorais valley, Pakistan (Provided by Dr. Muhammad Abbas in Pakistan Museum of Natural History)148 Chlaenius propeagilis sp. n., Gaoligongshan, Yunnan, China149 Chlaenius leishanensis Kirschenhofer,Xiaodanjiang, Leigongshan, Leishan county, Southeast Guizhou, China150Chlaenius chuanqianensis sp. n. and Chlaenius agiloides Jedlička, Jinshajiang, Sichuan151 Chlaenius rambouseki Lutshnik, Ussri river, Heilongjiang, China.

Photographs of habitats for Chlaenius (Lithochlaenius) spp. 152 Chlaenius rambouseki Lutshnik, Baisha county, Hainan Island, China153 Chlaenius rambouseki Lutshnik, Yuanbaoshan, Guangxi, China154 Chlaenius rambouseki Lutshnik adult walking on concrete wharf of Ussri river at night155 Chlaenius agiloides Jedlička adults mating at night156 Assorted molluscs, show one food of Chlaenius rambouseki Lutshnik from Hainan Island157 Adults of Chlaenius agiloides Jedlička preying on dragonfly nymph at night.

Abdominal sternum VII (ventral aspect) of Chlaenius (Lithochlaenius) species94 Chlaenius chuanqianensis sp. n., male, holotype95 Chlaenius chuanqianensis sp. n., female, paratype96 Chlaenius linwensini sp. n., male, holotype97 Chlaenius linwensini sp. n., female, paratype98 Chlaenius propeagilis sp. n., male, holotype99 Chlaenius propeagilis sp. n., female, paratype100 Chlaenius agilis Chaudoir, male, holotype101 Chlaenius formosensis Lorenz, male102 Chlaenius formosensis Lorenz, female103 Chlaenius leishanensis Kirschenhofer, male104 Chlaenius leishanensis Kirschenhofer, female105 Chlaenius noguchii Bates, male106 Chlaenius noguchii Bates, female107 Chlaenius wrasei Kirschenhofer, male108 Chlaenius wrasei Kirschenhofer, female109 Chlaenius agiloides Jedlička, male110 Chlaenius agiloides Jedlička, female111 Chlaenius rambouseki Lutshnik, from Ussuri region, male112 Chlaenius rambouseki Lutshnik from Ussuri region, female113 Chlaenius anchomenoides Bates, male, paratype114 Chlaenius nuristanus Jedlička, male, paratype115 Chlaenius nuristanus Jedlička, female, paratype116 Chlaenius formosanus Jedlička from Taiwan, male. Scale line = 1.0 mm.

AndrewesHE (1930) Catalogue of Indian Insects. Part 18–Carabidae. Government of India Central Publication Branch, Calcutta, xxii + 389pp.PaikJKTracDHWillK (2006) Carabidae from Vietnam (Coleoptera).Journal of Asia-Pacific Entomology 9: 85-105.BellRT (1960) A revision of the genus Chlaenius Bonelli (Coleoptera, Carabidae) in North America.Miscellaneous Publications of the Entomological Society of America 1: 97-166.KryzhanovskijOL (1976) A survey of the Callistini tribe (Coleoptera, Carabidae) in the Far East. Proceedings of the Institute of Soil Biology (N.S.) 43 (146): 8-17.