Lithochlaenius Kryzhanovskij, 1976

Valim, Ye, Kavanaugh, David H., Shi, Hongliang & Liang, Hongbin, 2011, A key to species of subgenus Lithochlaenius (Coleoptera, Carabidae, Chlaeniini, Chlaenius), with descriptions of three new species, ZooKeys 128, pp. 15-52 : 16-19

Subenus Lithochlaenius Kryzhanovskij, 1976

Hemichlaenius Lutshnik, 1933:169 (nec Bates, 1892:307). Type species Chlaenius rambouseki Lutshnik, 1933; Kryzhanovskij, 1976:11

Lithochlaenius Kryzhanovskij, 1976:9. Type species Chlaenius rambouseki Lutshnik, 1933; Morita, 1993:161

Agilochlaenius Kirschenhofer, 1997:116. Type species Chlaenius latro LaFerté-Sénectère, 1851; Kirschenhofer, 2000:58

Diagnosis.

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 ).

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. rambouseki is 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 Lithochlaenius and 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 solitarius groups 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 apica l lamella of the male aedeagus among New World species than we have seen among the Asian Lithochlaenius species. 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

1	All 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
2	Intervals 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
3	Antennomere 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
4	Intervals 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 stout	5
5	Tibiae 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
6	All 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
7	Antennomeres 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.
8	Intervals 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
9	Male 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

Figures

Figure 158. Map showing the known geographical distributions of Chlaenius (Lithochlaenius) spp.

Figures 1 - 15. Character states of Chlaenius (Lithochlaenius) spp. Figs 1 - 13 Chlaenius rambouseki Lutshnik 1 pronotum 2 site of hind angle seta 3 elytral base showing the site of basal pore 4 abdominal sterna showing the pubescence 5 male anterior tarsomere 4 in dorsal view 6 male anterior tarsomere 4 in ventral view 7 female anterior tarsomere 4 in dorsal view 8 female anterior tarsomere 4 in ventral view 9 antennomeres 1 - 5 showing antennomere 3 distinctly longer than 1 and 2 ones combined 10 metepisternum 11 aedeagus, showing the basic structure of the aedeagus in Lithochlaenius species 12 female gonostylus in ventral view 13 female gonostylus in lateral view 14 female gonostylus of Chlaenius propeagilis sp. n. in ventral view 15 female gonostylus of Chlaenius propeagilis sp. n. in lateral view.

Figures 117 - 130. Aedeagi of Chlaenius (Lithochlaenius) species 117 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 noguchii Bates (dorsal view) 130 Chlaenius noguchii Bates (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).

Figures 131 - 145. Aedeagi of Chlaenius (Lithochlaenius) species 131 Chlaenius wrasei Kirschenhofer (dorsal view) 132 Chlaenius wrasei Kirschenhofer (left lateral view) 133 Chlaenius agiloides Jedlicka (dorsal view) 134 Chlaenius agiloides Jedlicka, holotype (left lateral view) 135 Chlaenius agiloides Jedlicka (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 Jedlicka, paratype (dorsal view, black leg) 141 Chlaenius nuristanus Jedlicka, paratype (left lateral view) 142 Chlaenius nuristanus Jedlicka, paratype (dorsal view, yellow leg) 143 Chlaenius nuristanus Jedlicka, paratype (left lateral view) 144 Chlaenius rambouseki Lutshnik from Taiwan (dorsal view) 145 Chlaenius rambouseki Lutshnik 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).

Figures 52 - 65. Antennomere 1 of Chlaenius (Lithochlaenius) species 52 Chlaenius chuanqianensis sp. n., holotype 53 Chlaenius linwensini sp. n., holotype 54 Chlaenius propeagilis sp. n., holotype 55 Chlaenius agilis Chaudoir, holotype 56 Chlaenius formosensis Lorenz, CCCC 57 Chlaenius leishanensis Kirschenhofer, IZCAS 58 Chlaenius noguchii Bates, IZCAS 59 Chlaenius wrasei Kirschenhofer, IZCAS 60 Chlaenius agiloides Jedlicka, IZCAS 61 Chlaenius rambouseki Lutshnik from Ussuri region, IZCAS 62 Chlaenius anchomenoides Bates, paratype 63 Chlaenius nuristanus Jedlicka, paratype 64 Chlaenius nuristanus a. rubridipes Jedlicka, paratype 65 Chlaenius rambouseki Lutshnik from Taiwan, CCCC. Scale line = 1.0 mm.

Figures 80 - 93. Elytral intervals of Chlaenius (Lithochlaenius) spp. 80 Chlaenius chuanqianensis sp. n., holotype 81 Chlaenius linwensini sp. n., holotype 82 Chlaenius propeagilis sp. n., holotype 83 Chlaenius agilis Chaudoir, holotype 84 Chlaenius formosensis Lorenz, CCCC 85 Chlaenius leishanensis, IZCAS 86 Chlaenius noguchii Bates, IZCAS 87 Chlaenius wrasei Kirschenhofer, IZCAS 88 Chlaenius agiloides Jedlicka, IZCAS 89 Chlaenius rambouseki Lutshnik from Ussuri region, IZCAS 90 Chlaenius anchomenoides Bates, paratype 91 Chlaenius nuristanus Jedlicka, paratype 92 Chlaenius nuristanus Jedlicka, paratype 93 Chlaenius rambouseki Lutshnik from Taiwan, CCCC. Scale line = 1.0 mm.

Figures 146 - 151. Photographs of habitats for Chlaenius (Lithochlaenius) spp. 146 Chlaenius chuanqianensis sp. n., Dabaitang, Xishui county, North Guizhou, China 147 Chlaenius anchomenoides Bates, Goorais valley, Pakistan (Provided by Dr. Muhammad Abbas in Pakistan Museum of Natural History) 148 Chlaenius propeagilis sp. n., Gaoligongshan, Yunnan, China 149 Chlaenius leishanensis Kirschenhofer, Xiaodanjiang, Leigongshan, Leishan county, Southeast Guizhou, China 150 Chlaenius chuanqianensis sp. n. and Chlaenius agiloides Jedlicka, Jinshajiang, Sichuan 151 Chlaenius rambouseki Lutshnik, Ussri river, Heilongjiang, China.

Figures 152 - 157. Photographs of habitats for Chlaenius (Lithochlaenius) spp. 152 Chlaenius rambouseki Lutshnik, Baisha county, Hainan Island, China 153 Chlaenius rambouseki Lutshnik, Yuanbaoshan, Guangxi, China 154 Chlaenius rambouseki Lutshnik adult walking on concrete wharf of Ussri river at night 155 Chlaenius agiloides Jedlicka adults mating at night 156 Assorted molluscs, show one food of Chlaenius rambouseki Lutshnik from Hainan Island 157 Adults of Chlaenius agiloides Jedlicka preying on dragonfly nymph at night.

Figures 28 - 33. Chlaenius (Lithochlaenius) spp 28 Chlaenius noguchii Bates, male, in IZCAS, dorsal view 29 Chlaenius noguchii Bates, male, in IZCAS, ventral view 30 Chlaenius wrasei Kirschenhofer, male, in IZCAS, dorsal view 31 Chlaenius wrasei Kirschenhofer, male, in IZCAS, ventral view 32 Chlaenius agiloides Jedlicka, male, in IZCAS, dorsal view 33 Chlaenius agiloides Jedlicka, male, in IZCAS, ventral view. Scale line = 5.0 mm.

Figures 94 - 116. Abdominal sternum VII (ventral aspect) of Chlaenius (Lithochlaenius) species 94 Chlaenius chuanqianensis sp. n., male, holotype 95 Chlaenius chuanqianensis sp. n., female, paratype 96 Chlaenius linwensini sp. n., male, holotype 97 Chlaenius linwensini sp. n., female, paratype 98 Chlaenius propeagilis sp. n., male, holotype 99 Chlaenius propeagilis sp. n., female, paratype 100 Chlaenius agilis Chaudoir, male, holotype 101 Chlaenius formosensis Lorenz, male 102 Chlaenius formosensis Lorenz, female 103 Chlaenius leishanensis Kirschenhofer, male 104 Chlaenius leishanensis Kirschenhofer, female 105 Chlaenius noguchii Bates, male 106 Chlaenius noguchii Bates, female 107 Chlaenius wrasei Kirschenhofer, male 108 Chlaenius wrasei Kirschenhofer, female 109 Chlaenius agiloides Jedlicka, male 110 Chlaenius agiloides Jedlicka, female 111 Chlaenius rambouseki Lutshnik, from Ussuri region, male 112 Chlaenius rambouseki Lutshnik from Ussuri region, female 113 Chlaenius anchomenoides Bates, male, paratype 114 Chlaenius nuristanus Jedlicka, male, paratype 115 Chlaenius nuristanus Jedlicka, female, paratype 116 Chlaenius formosanus Jedlicka from Taiwan, male. Scale line = 1.0 mm.

Figures 22 - 27. Habitus of Chlaenius (Lithochlaenius) spp 22 Chlaenius agilis Chaudoir, holotype, dorsal view 23 Chlaenius agilis Chaudoir, holotype, ventral view 24 Chlaenius formosensis Lorenz, male, in CCCC, dorsal view 25 Chlaenius formosensis Lorenz, ventral view 26 Chlaenius leishanensis Kirschenhofer, male, in IZCAS, dorsal view 27 Chlaenius leishanensis Kirschenhofer, male, in IZCAS, ventral view. Scale line = 5.0 mm.

Figures 16 - 21. Habitus of Chlaenius (Lithochlaenius) spp 16 Chlaenius chuanqianensis sp. n., holotype, male, dorsal view 17 Chlaenius chuanqianensis sp. n., holotype, male, ventral view 18 Chlaenius linwensini sp. n., holotype, male, dorsal view 19 Chlaenius linwensini sp. n., holotype, male, ventral view 20 Chlaenius propeagilis sp. n., holotype, male, dorsal view 21 Chlaenius propeagilis sp. n., holotype, male, ventral view. Scale line = 5.0 mm.

Figures 40 - 45. Chlaenius (Lithochlaenius) spp 40 Chlaenius nuristanus a. rubridipes Jedlicka, paratype, male, in MNHN, dorsal view 41 Chlaenius nuristanus a. rubridipes Jedlicka, paratype, male, in MNHN, ventral view 42 Chlaenius agiloides Jedlicka, holotype, male, in NMPC, dorsal view 43 Chlaenius formosanus Jedlicka, holotype, male, in NMPC, dorsal view 44 Chlaenius rambouseki Lutshnik, from Taiwan, male, in IZCAS, dorsal view 45 Chlaenius rambouseki Lutshnik, from Taiwan, male, in IZCAS, ventral view. Scale line = 5.0 mm.