Adults of this subtribe can be recognized by combination of the following characters: mandibles moderately to strongly widened; apex of ligula with four or more setae; palpifers without seta; mentum with tooth simple or bifid; front angles of pronotum with some setae longer or shorter; pronotal base usually more or less lobed; elytral dorsal setigerous pores, if distinct, at least with one present on base of 5th interval; elytral apex truncate, outer angles completely rounded, not angulate, sutural angles not projected; apex of 7th and 8th intervals more or less tumid; penultimate pore of elytral umbilical series not displaced laterally or medially; males with terminal sternum more or less emarginate; 4th tarsomere strongly bilobed; claws pectinate; median lobe of aedeagus usually with apical orifice opened apically, dorsal surface with some setae subapically, such setae generally fine but sometimes very strong, present around apical orifice; internal sac generally with a long flagellum-like sclerite; right paramere trifurcate, apex usually widened; apical segment of ovipositor without spine, apex with extension usually membranous; spermatheca inserted on bursa copulatrix or joining of common oviduct and bursa copulatrix.

Based on widened mandibles, bifid 4th tarsomere, pectinate claws, and female ovipositor characters, Calleidina may be more closely allied with Physoderina than any other subtribe of Lebiini from the Oriental Region. Members of Calleidina can be readily distinguished from those of Physoderina by the absence of setae on the pronotum front angles, except Calleida sultana Bates. But Calleida sultana has a totally different shape of the aedeagus and no setigerous pore on 5th interval.

Reconstruction of phylogeny of Lebiini has been attempted by Ball et al. (1995) and Casale (1998), but their works did not focus on the systematic position of Physoderina. Their results support a close relationship between Physoderina and Metallicina + (Calleidina + Galerucidiina) (Ball et al. 1995) or Agra Fabricius (Casale 1998). Based on the study in the present work of all genera in Physoderina, we propose that Physoderina has more affinity with Calleidina than Metallicina. This is suggested by the following character states present in Physoderina and Calleidina but not in Metallicina: (1) apical segment of ovipositor without spine, usually spiculate, apex with extension usually membranous; (2) terminal labial palpomeres more or less widened in males; (3) mentum with tooth; (4) tempora ventrally without suborbital setigerous pore; and (5) males with terminal sternum emarginate.

So far, there is no rigorous phylogenetic analysis demonstrating that Physoderina is a monophyletic lineage in Lebiini. But, monophyly of this subtribe could be suggested by the following apomorphic character states: (1) setigerous pores present on 5th interval; (2) median lobe of aedeagus usually with apical orifice opened apically, internal sac usually with a long flagellum-like sclerite; (3) right paramere trifurcate; (4) spermatheca inserted on bursa copulatrix or joining of common oviduct and bursa copulatrix.

Chaudoir (1877) proposed the name Physodérides with insufficient definition, but ambiguously included Allocota Motschulsky, Cryptobatis Eschscholtz, Aspasiola Chaudoir and Lachnoderma Macleay. The original vernacular family-group name is available according to the Zoological Code of Nomenclature, and was first used in Latinized form by Bates (1883: 207) (Bouchard et al. 2011). Generally, Physoderina included seven genera (Jedlička 1963, Löbl and Smetana 2003, Lorenz 2005) before the present work.

In the present paper, we move three genera previously placed in Calleidina or Pericalina to Physoderina, propose two new generic synonyms, resurrect one generic name from synonymy, and describe one gen. n.. Hence, a total of ten genera is presently included in Physoderina: Paraphaea Bates, Anchista Nietner, Metallanchista gen. n., Physodera Eschscholtz, Diamella nom. n., Allocota Motschulsky, Lachnoderma Macleay, Dasiosoma Britton, Orionella Jedlička, and Endynomena Chaudoir.

The New World Cryptobatida group (sensu Erwin 2004) shares many characters with Physoderina from the Old World, such as: mandible widened, pronotum more or less angulate in middle, elytral disc depressed, and, above all, internal sac of male genitalia with flagellum-like sclerites. We propose that this group could be most closely related to Physoderina and perhaps should be included therein, similar to the concept including Cryptobatis in the original Physoderina (Chaudoir 1877). But, as the focus of the present work is the Old World fauna, we do not present a better subtribal arrangement for this New World group.

All of the ten genera have their center of distribution in the Oriental Region. One of them (Dasiosoma) has some African species; two genera (Physodera, Lachnoderma) have only a few Australian species; two species (Paraphaea binotata, Endynomena pradieri) have an Oriental origin, but are also widely distributed in the Pacific islands.

Paraphaea signifera Bates, 1873 [= Paraphaea binotata (Dejean)], by monotypy.

Paraphaea Bates can be readily distinguished from most genera of Physoderina except Anchista Nietner and Metallanchista gen. n. by the glabrous surface, moderately widened mandibles, and long setae around the aedeagal apical orifice.

Differences between Anchista and Paraphaea are: (1) pronotum lateral margins slightly angulate in middle in Anchista, but completely rounded in Paraphaea; (2) Anchista usually with distinct isodiametric microsculpture on the elytra, while Paraphaea has indistinct microsculpture; (3) in Paraphaea, males with adhesive hairs on the 1st metatarsomere, but such hairs absent in Anchista; (4) males with terminal sternum deeply emarginate in Paraphaea, but only shallowly emarginate in Anchista; (5) internal sac of aedeagus without main flagellum in Anchista, but main flagellum well developed and sinuous in Paraphaea; (6) in Paraphaea, the spermathecal gland inserted near the middle of the spermatheca, spermatheca more or less bent near the middle, but in Anchista the spermathecal gland inserted near the apex of spermatheca, spermatheca nearly straight.

Comparison between Paraphaea and Metallanchista gen. n. is presented in the diagnosis part under Metallanchista gen. n.

Dorsal side generally reddish-brown to dark brown, sometimes with faint metallic reflections; elytra unicolored or bicolored. Head glabrous; eyes hemispherical and strongly prominent; tempora shorter than half of eyes length, abruptly narrowed behind eyes; vertex flat. Antennae reaching elytral base; 1st antennomere slightly narrowed at base, 3rd slightly longer than 4th. Labrum smooth, without secondary setae; mandibles moderately widened, outer margin nearly straight (Fig. 149), glabrous on outer scrobe and dorsal ridge; terminal maxillary palpomeres fusiform in males and females; terminal labial palpomeres strongly securiform and truncate apically in the males, narrower in the females; ligula with apex slightly projected, with four long setae; paraglossae membranous, not longer than ligula, adnate; mentum tooth simple, with two setae near base; submentum with two long setae; genae glabrous beneath eyes. Pronotum slightly wider than head, disc glabrous or sparsely pubescent; mid-lateral setae present; front angles more or less setose, hind angles generally with a few additional short setae; pronotal base briefly but distinctly lobed; lateral margins completely rounded in middle (Fig. 155), more or less sinuate before hind angles; hind angles sharp, rectangular or subrectangular. Elytra wide, apex truncate, sutural angles not projected, outer angles completely rounded; sides slightly depressed in anterior third, disc with an indistinct depression near apical two-fifths; intervals glabrous or only odd intervals with a few additional setae; umbilical pores of 9th interval placed in one row (Fig. 147); basal margination nearly complete; basal pores well developed; 3rd interval with two to four setigerous pores, 5th interval with base slightly widened, with one setigerous pore; 7th and 8th intervals slightly tumid near apex. Ventral side nearly glabrous; males with terminal sternum deeply emarginate (Fig. 143), with one pair of setae; females with apex of terminal sternum straight or slightly emarginate, with two pairs of setae. Legs short; protibiae with cleaning spur well developed, distant from inner margin; tarsi widened, 4th tarsomere bifid, claws pectinate; males with adhesive hairs well developed (two whole rows) on 1st to 3rd pro-, 1st to 2nd meso- and 1st metatarsomeres, rudimentary (two rows but very weakly present near apex) on 3rd mesotarsomere. Male genitalia with median lobe of aedeagus twisted to left; apical orifice opened dorsally, strongly setose along basal margin; internal sac with main flagellum more or less sinuous, nearly reaching apical orifice, trumpet-form expansion small and strongly bent; apical bursa strongly sclerotized; three additional small sclerotized pieces placed near apical third; secondary flagellum short and indistinct (Fig. 67). Female genitalia. Spermatheca tubular, with distinct ring-sculpture, inserted on bursa copulatrix; spermathecal gland slender and long, inserted near middle of spermatheca; spermatheca more or less bent near middle. Apical segment of ovipositor scimitar-shaped, curved to outer side, inner margin slightly angulate near apex; with fine setae near apex; apex with elongate membranous extension.

(Maps 1, 2). This genus includes four species distributed in South and Southeast Asia. One of them (Paraphaea binotata) has a rather wide distribution, from Japan and India to the western Pacific islands (Map 1), but the other three species are more restricted.

Paraphaea is presumed to be the sister group of Anchista. The relationship is supported by these character states: (1) mandibles moderately widened; (2) terminal sternum with single seta on each side in males; (3) median lobe of aedeagus strongly setose around apical orifice; (4) apical segment of ovipositor with inner margin slightly angulate near apex.

Monophyly of Paraphaea is suggested by the following apomorphic character states: (1) males with adhesive hairs present on hind tarsi; (2) males with terminal sternum deeply emarginate; (3) median lobe of aedeagus twisted; (4) spermatheca more or less bent near middle.

Bates (1873) proposed the genus Paraphaea without comparing it with Anchista Nietner. Later, Chaudoir (1877) synonymized these two genera without detailed explanation. Based on external characters, it is difficult to find significant differences between these two genera, but the genital and male secondary sexual characters mentioned above provide clear differences and justify the generic separation of Paraphaea Bates.

Lectotype of Plochionus binotatus Dejean, designated herein (MNHN): male, body length = 7.8 mm, pin mounted, “TYPE” [red label]; “binotatus, mil. / in inf. Mariannes” [yellow label]; “Guerin” [yellow label]; “Plochionus”; “Paraphaea / signifera Bates”; “Ex Musaeo / Chaudoir” [red letters]; “Museum Paris / 1952 / Coll. R. Oberthür”; “LECTOTYPE ♂ / Plochionus binotatus / Dejean, 1825 / des. SHI H. L. 2011” [red label][Fig. 31]. Lectotype of Calleida discophora Chaudoir, designated herein (MNHN): male, body length = 8.0 mm; pin mounted; labia removed and separately pinned, “Calleida / discophora Ch.”; “Ex Musaeo / Chaudoir” [red letters]; “Museum Paris / 1952 / Coll. R. Oberthür”; “LECTOTYPE ♂ / Calleida discophora / Chaudoir, 1852 / des. SHI H. L. 2011” [red label][Fig. 32]. Lectotype of Paraphaea signifera Bates, designated herein (MNHN): male, body length = 7.9 mm; pin mounted, “TYPE” [red label]; “SATZUMA.”; “Ex. Musaeo / H. W. Bates, 1892”; “Museum Paris / 1952 / Coll. R. Oberthür”; “Paraphaea / signifera / Bates”; “LECTOTYPE ♂ / Paraphaea signifera / Bates, 1873 / des. SHI H. L. 2011” [red label][Fig. 33]. Paralectotype of Paraphaea signifera Bates (MNHN): a female, body length = 8.8 mm; pin mounted, labia removed and separately pinned, “SATZUMA.”; “PARATYPE” [red label]; “Ex. Musaeo /H. W. Bates / 1892”; “Museum Paris / 1952 / Coll. R. Oberthür”; “PARALECTOTYPE ♀ / Paraphaea signifera / Bates, 1873 / des. SHI H. L. 2011” [red label]. Holotype of Anchista eurydera Chaudoir, monotypy (MNHN): male, without head, pin mounted, “Ex Musaeo / Chaudoir” [red letter]; “TYPE” [red label]; “Museum Paris / 1952 / Coll. R. Oberthür”; “eurydera / Chaudoir / Indes Orient” [box label but pinned under specimen]; “HOLOTYPE ♂/ Anchista eurydera / Chaudoir 1877 / det. SHI H. L. 2011” [red label].

Plochionus binotatus Dejean: In the collection of MNHN, we found only one specimen (Fig. 31) from the Mariannas, apparently from Dejean’s collection. So this specimen could be the basis for the original description, although the original literature didn’t indicate or imply that the species was based on a single specimen. According to the Zoological Code of Nomenclature (4th Edition), Articles 61, 73 and 74, for taxonomic purpose of fixing the name to unique name-bearing type and preventing further uncertainty, we designate this specimen as the lectotype.

Calleida discophora Chaudoir: The original literature didn’t indicate or imply how many specimens were examined. It can be confirmed that the male (Fig. 32) from Chaudoir’s collection, and bearing Chaudoir’s hand-written label, belongs to the type series. The other three specimens in Chaudoir’s collection do not accord with the original literature, although they are mentioned later (Chaudoir 1877). We herein designate this first specimen as lectotype for taxonomic purpose of fixing the name to unique name-bearing type.

Paraphaea signifera Bates: This species was originally described from an unspecified number of specimens, but both sexes were mentioned, as well as type locality “Satzuma”. In the collection of MNHN, two specimens (one male and one female) from Bates’ collection are perfectly in accord with the original literature. We herein designate the male (Fig. 33) as lectotype for the purpose of providing the name with a unique name-bearing type. Another specimen from Bates’ collection with different locality was labeled as paratype by J. Mateu (not published), but it is neither a paratype nor a syntype and should not be part of the type series. We removed the paratype label.

Anchista eurydera Chaudoir: This species was originally described from a single specimen which head is missing. This male in MNHN (ex. collection of Chaudoir) is clearly the holotype.

(total 141 specimens). China: 1 female (IZAS), “Jiangxi, Xingguo, 1956.7.26, Chen Yong leg., early season rice”. 1 female (IZAS), “Jiangxi, Xingguo, 1956.7.2, Chen Yong leg., early season rice”. 1 female (IZAS), “Jiangxi, Yongxin, 1956.5.12, Long Changqi leg., early season rice”. 1 male, 2 females (IZAS), “Jiangxi, Ganzhou, 1956.8.1, Qiu Futao leg., barley”. 1 female (IZAS), “Guangdong, Zhongshan, Cuihengcun, 1957.8.4, grassland”. 1 female (IZAS), “Guangdong, Yingde, Chengguan Granary, 1957.8.22”. 2 males (IZAS), “Guangdong, Zhongshan, Baqu, 1957.7.29”. 1 male (IZAS), “Guangdong, Nanxiong, 1956.7.5, Wu Guangwen leg., rice land”. 5 males 6 females (IZAS), “Guangxi, Guilin, Yanshan Mt., 1953.4.24/ 1953.5.4/ 1953.7.17”[Figs 67, 95]. 1 female (HBUM), “Guangxi, Guilin, Yanshan Mt., 2003.V.17, Li Tianshan leg.”[Fig. 126]. 7 males 4 females (IZAS), “Guangxi, Guilin, Liangfeng, 1952.3.13/ 1952.5.17”.[Fig. 111] 2 males 3 females (IZAS), “Guangxi, Lingui, 1952.3.6/ 1952.3.10/ 1952.3.13/ 1958.4.22”. 2 males (IZAS), “Guangxi, Lingui, Dayu, 1958.4”. 1 female (IZAS), “Yaoshan, 1938.V.25”. 1 female (IZAS), “Yangshuo, 1938.VIII.1”. 1 male (IZAS), “Yunnan, Jinping, 1958.6.19, black fungi”. 1 female (IZAS), “Yunnan, Luxi, 1957.5.20, Zhao Wenguang leg., rice land”. 1 female (IZAS), “Yunnan, Cheli, 1958.VIII.29”. 1 female (IZAS), “Yunnan, Xishuangbanna, Yunjinghong, 900m, 1958.VI.26, Zhang Yiran leg.”[Fig. 1]. 1 female (IZAS), “Yunnan, Xishuangbanna, Damenglong, 650m, 1958.VIII.7, Zhang Yiran leg.”. 3 specimens (MNHN), “Chine. Kouangsi, P. Barriere, 1909”. 1 specimen (MNHN), “Chine, Foo Kien, M. dela Touche, 1899”. Japan: 1 specimen (MNHN), “Japan“; “Ex. Musaeo, H. W. Bates, 1892”. Vietnam:2 males 4 females (IZAS), “Tonkin, Hoa-Binh, 1939.VII/ 1940.VII/ 1940.VIII, leg. A. de Cooman”. 1 specimen (MNHN), “Honoï, Domonge, 1909”. 1 specimen (MNHN), “Cochinchine, aui nhon”. 2 specimens (MNHN), “cochinchine, Hasmand, 1872”. 1 specimen (MNHN), “Cochinchine, Env. De Saigon, Simard, 1902”. 1 specimen (MNHN), “Cochinchine, Baudooin d’Aulne, 1897”. 2 specimens (MNHN), “Tonkin Sept., Ha-Lang, Lamey 151-97”. 37 specimens (MNHN), “Tonkin occ., Env. De Hoa-Binh, R. P. A. de Cooman, 1919.”. 9 specimens (MNHN), “Hoah binh, Tonkin”. Thailand: 2 specimens (MNHN), “Siam, Chantaboun, A. Battambang, A. Pavie, 1886”. Cambodia: 1 specimen (MNHN), “Cambodia“; “Ex. Musaeo, H. W. Bates, 1892”. 2 specimens (MNHN), “Cambodge, Rég de Chiehreng, G. Thomas 1912.”. 1 specimen (MNHN), “Kampong, Cambidge, Coll. J. Negre”. 1 specimen (MNHN), “Cambodge, 20/9/1912, R. Vitalis de Salvaza”. Myanmar:1 specimen (MNHN), “Bhamò, Birmania, Fea VII 1886”; “Ex. Musaeo, H. W. Bates, 1892”. 1 specimen (MNHN), “Minhia, Birmania, D. Comotto 1883”; “Ex. Musaeo, H. W. Bates, 1892”. 1 specimen (MNHN), “Birmanie, Theinzeik, P. Loizeau, 1914”. 1 male (MNPC), “Tharrawaddy, Burma, G. Q. Corbett”. Indian: 1 specimen (MNHN), “Ind. Or. Bor., Dr Bacon”; “Ex. Musaeo, Chaudoir”. 1 specimen (MNHN), “I. Andamman, H. Deyroll.”; “Ex. Musaeo, Chaudoir”. 1 specimen (MNHN), “Andaman”. 1 specimen (MNHN), “I. Andaman, Deyrolle 1877”. 1 specimen (MNHN), “Andaman, Coll. Borel”. 1 specimen (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, XI-XII 1896”. 2 specimens (MNHN), “Naga Hills.”. 2 specimens (MNHN), “N. Manipur”. 1 specimen (MNHN), “Sikkim, Guntok, Eté 1894, Chasseurs Bretaudeau”. The Philippines: 1 specimen (MNHN), “Philipines, Ch. semper”. Indonesia: 1 specimen (MNHN), “Java, Mt. Tengger. Mme. E. Walsh”. 1 specimen (MNHN), “Sumatra, Rég. De. Benkoelen, Tandjong Sakti, Mme. M. E. Walsh, 1935”. 2 specimens (MNHN), “Paggar Alam, Sumara, J. Bouchard”. 1 specimen (MNHN), “Boreno Occ., Pontianak, 1903”.

Pronotum glabrous, widest at apical third, lateral margins slightly sinuate before hind angles; elytra with bicolored pattern, dark brown background with a reddish yellow elongate spot on each side (Fig. 1); elytra with two setigerous pores on 3rd interval, and one setigerous pore on base of 5th interval; elytra without any secondary setigerous pores; median lobe of aedeagus with apical third strongly expanded, internal sac with screwed main flagellum (Fig. 67).

The unique pronotal shape, elytral pattern and number of setigerous pores easily distinguish this species from all other allied species.

Male genitalia. Median lobe of aedeagus with apical third strongly expanded; in dorsal view, left-lateral margin strongly sinuate medially, and then gradually narrowed to apex; apical lamella placed on left-ventral side, broadly triangular, apex slightly rounded, not distinctly extended apically; internal sac with main flagellum screwed; basal part of main flagellum strongly bent, so trumpet-form expansion reaching right margin, apical margin of trumpet-form expansion crenulate (Fig. 67). Female genitalia. Spermatheca very long and slender; spermathecal gland inserted near apical one-third of spermatheca, with a short branch near base; spermatheca slightly expanded, with ring-sculpture between the gland insertion and apex, basal part of spermatheca without sculpture; spermatheca strongly bent at apical third (Fig. 126). Apical segment of ovipositor scimitar-shaped, inner margin slightly angulate at apical third; length about four times basal width; inner margin setose in apical half; apex sharp, with membranous extension long and slender (Fig. 111).

Detailed description of external characters has been provided by Habu (1967, 1982).

(Map 1). China [Jiangxi, Guangdong, Guangxi, Yunnan, Taiwan (only Orchid Island, by Kanô 1930a)]; Japan; Vietnam; Thailand; Cambodia; Myanmar; India; the Philippines; Indonesia; New Guinea (Darlington, 1968); Mariana.

Anchista eurydera Chaudoir was described from Indes Orientales which overlaps the range of Paraphaea binotata. The head of the male holotype is missing and its abdomen was badly destroyed by dermestid beetles, so it is impossible to examine its genitalia. Chaudoir (1877) noted that this species is extremely close to Paraphaea binotata, and wrote of some differences between these two species which are correct according to the holotype. But the differences in pronotal shape and elytral pattern should be regarded as mere individual variation, which is ubiquitous in this common and widely-distributed species. The paler color and more distinct yellow stripes on the pronotum are due to immaturity of the holotype. The transverse wrinkles on the pronotum are possibly an artifact caused by abnormal emergence. So we herein synonymize Anchista eurydera Chaudoir with Plochionus binotatus Dejean.

In Japan, this species is only recorded in South Kyushu and Satsunan Islands. Kanô’s (1930b) record of Paraphaea binotata from Tokyo is presumably a misidentification of Euplynes batesi Harold (Habu 1967). Kanô (1930a) recorded Paraphaea binotata from Taiwan (Orchid Island). It is probable that this record is also based on a misidentification. Moreover, we have studied a large series of carabid specimens from Taiwan, but didn’t find this species. So, it seems likely that this species is not distributed in Taiwan.

Holotype of Parena formosana Jedlička, by monotypy (NMPC): male, body length = 7.0 mm, board mounted, genitalia dissected and deposited in microvial pinned under specimen “Kuraru / Formosa. / 8-V.1935 / Col Y. Miwa”; “TYPUS” [red label with black frame]; “formosana / sp. n. / DET. ING. JEDLIČKA” [pink label][Fig. 34].

Type locality “Kuraru” is the old spelling of “Kueitzuchiao”, in the area of “Kenting National Park” at the southernmost part of Taiwan.

(Total 10 specimens). 1 male (CCCC), “Taiwan, Pingtung County, Lilong Mt., 2008.XI.5 N, Chou Wen-I leg.” [Fig. 2]. 1 male (CCCC), “Taiwan, Pingtung County, Fenggang, Longfengsi, 2010.VIII.8 N, Lin Wensin leg.”. 1 male (MTMB), “Taiwan: Pingtung County, Kenting National Park, Hengchun Research Center, 21°57'43.0"N, 120°48'50.0"E, Lanyu plant collection, sweeping, 2008.VIII.18., leg. Redei D & Tsai JF”. 1 male (CCCC), “Taiwan, Taitung County, Haiduan, 1996.V.18, Chou Wen-I leg.”[Fig. 68]. 1 male (CCCC), “Taiwan, Taoyuan County, Fusing, Shangbaling, 1200m, 1994.VII.30, Chen Changchin leg.”. 1 female (MTMB), “Taiwan, Taipei county, PiHu, at light, 3.IV.2002, leg. Gy. Fabian & O. Merkl”. 1 female (NHML), “yauo. F. 31. FORMOSA MC:AR1 Abatsusen?. XII. 07 y. yauo”; “H. E. Andrewes Coll. B. M. 1945-97”. 1 female (NHML), “Mokuriryo, Near Mt. Ari, Formosa 1-IV. 1938 Coll. Yoshio Yano”; “H. E. Andrewes Coll. B. M. 1945–97”. 1 male (NHML), “Mokuriryo, Near Mt. Ari, Formosa 1-IV. 1938 Coll. Yoshio Yano”; “H. E. Andrewes Coll. B. M. 1945–97”. 1 female (NHML), “Formosa KAGI MT. TAIKOU 1937 Coll. Y. Yano.”; “27-12-1937 Coll. Yoshio Tano.”; “No. 626 YOSHIO YANO COLLECTION”; “H. E. Andrewes Coll. B. M. 1945–97”.

Pronotum glabrous, widest at middle, lateral margins sinuate before hind angles; elytra dark brown without pattern, or disc with a faint reddish patch; elytra with three or four setigerous pores on 3rd interval and one setigerous pore on base of 5th interval, in some specimens odd intervals with a few secondary setae; median lobe of aedeagus with apical third slightly expanded, internal sac with sinuous main flagellum (Fig. 68).

This species is closest to Paraphaea minor. Distinguishing characters are provided in the diagnosis of Paraphaea minor.

Body length 7.0–7.8 mm; head and pronotum reddish brown to dark brown, lateral explanate areas of pronotum paler; mouthparts, antennae and legs yellowish brown or reddish brown; elytra with luster, uniformly dark brown or disc somewhat reddish, in some specimens with a faint reddish patch on central area, the patch at most reaching 6th interval; ventral side yellowish brown to dark brown. Head glabrous, without punctures or microsculpture. Pronotum cordiform, widest near middle; ratio PW/PL 1.31 to 1.40; pronotal base briefly but distinctly lobed; disc glabrous, microsculpture indistinct, without punctures; front angles with a few setae, sometimes abundant and long (in specimens with additional interval setae), or sparse and short, but always distinctly longer than in Paraphaea binotata; lateral margins completely rounded in middle, sinuate before hind angles; front angles wide and rounded, not projected; hind angles rectangular or nearly so, slightly pointed; disc moderately convex; basal foveae wide and even, with a few punctures. Elytra with striae distinct, finely punctate along striae; intervals slightly convex, without punctures or microsculpture; 3rd interval with three or four setigerous pores, the apical one placed about apical one-eighth, adjacent to 2nd stria, position of the other two or three pores variable, pore number and position variable even in same specimen; 5th interval with one setigerous pore near base, adjacent to 5th stria; in some specimens, the odd intervals with some secondary setae; 17-19 umbilical pores on 9th interval. Male genitalia. Median lobe of aedeagus with apical third slightly expanded; in dorsal view, left-lateral margin moderately sinuate medially, and then gradually narrowed to apex; apical lamella placed in left-ventral side, narrowly triangular, apex rounded, slightly prolonged; internal sac with main flagellum sinuous; basal part of main flagellum moderately bent, trumpet-form expansion adjacent to left margin, apical margin of trumpet-form expansion not crenulate (Fig. 68). Female genitalia not studied.

(Map 2). This species is endemic in Taiwan.

This is a highly variable species, although not widely distributed. According to the eleven specimens examined, three forms exist on Taiwan island: (1) The typical form is found in the southernmost part of Taiwan (four specimens examined), with elytra uniformly dark brown or disc slightly reddish and intervals without secondary setae; (2) specimens from the Alishan Mountains (four specimens examined), with a faint reddish patch in the middle of the elytra; (3) specimens from the lowlands of Taiwan (three specimens examined from Taitung, Taoyuan and Taipei) with secondary setae on odd intervals, and setae on front angles of pronotum distinctly more abundant and longer than the other two forms. These three forms are different only in color and secondary setae. We dissected the holotype, two typical form from Pingtung and a lowland form from Taitung, and found no significant differences between their male genitalia. We didn’t study male genitalia of the Alishan form.

Holotype (MNHN): male, body length = 5.8 mm, board mounted, genitalia dissected and deposited in microvial pinned under specimen, “Hoa-Binh / Tonkin / A. de Cooman”; “MUSEUM PARIS / Coll. Ch. ALLUAUD”; “HOLOTYPE ♂/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label][Figs 69, 96, 35]. Paratypes (Total 1 male, 6 females): Vietnam: 1 female (MNHN), “Tonkin / Dap-Can”; “MUSEUM PARIS / Ex. Coll. M. MAINDRON / Coll. G. BABAULT 1930”; “Anchista /circumdata / sp. n.”; “PARATYPE ♀/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label]. Thailand: 1 female (MNHN), “MUSEUM PARIS / BANGKOK / LARNAUDIE 349-64”; a round yellow label probably meaning Oriental with backside written “3kg / Gk”; “PARATYPE ♀/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label]. 1 male (CRS), “Thailand bor. occ. / Pai / Soppong / 28.5-5.6. 1997 / Lgt. M. Snizek”; “PARATYPE ♂/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label]. Myanmar: 1 male (NHML), “Toungoo”; “Anchista / sp. n.”; “H. E. Andrewes Coll. / B. M. 1945-97”; “PARATYPE ♂/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label]. Cambodia: 1 female (MNHN), “Cambodge / Rég. De Chiehreng / G. Thomas 1912.”; “Muséum Paris / Coll. R. Oberthür / 1952”; “PARATYPE ♀/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label]. China: 1 female (HBUM), “2006-XI-15-16 / 海南昌江县尖峰岭 / 任国栋 / 河北大学博物馆” [2006-XI-15-16, Hainan, Changjiang, Jianfengling, Ren Guodong leg., Hebei University Museum]; “PARATYPE ♀/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label][Figs 112, 127]. 1 female (IZAS), “采集地：海南五指山 / 海拔 日期 2008-IV-03 / 方法 灯诱 采集人：杨玉霞” [Hainan, Wuzhishan, 2008.IV.03. by light trap, Yang Yuxia leg.]; “PARATYPE ♀/ Paraphaea minor / new species / Des. SHI H. L. 2011” [red label][Fig. 3].

Pronotum glabrous, widest at middle, lateral margins sinuate before hind angles; elytra reddish yellow with dark lateral and apical margins; two setigerous pores present on 3rd interval and one setigerous pore on base of 5th interval; median lobe of aedeagus with apical third slightly expanded, internal sac with sinuous main flagellum (Fig. 69).

The new species is most similar to Paraphaea formosana in male genitalia. It differs as follows: (1) 3rd interval with two setigerous pores in Paraphaea minor, but with three or four pores in Paraphaea formosana; (2) elytral patterns of these two species are different; (3) Paraphaea minor with median lobe of aedeagus less expanded subapically, left-lateral margin only slightly sinuate in dorsal view (Fig. 69); Paraphaea formosana with left-lateral margin of aedeagal median lobe more distinctly sinuate in dorsal view (Fig. 68); (4) Paraphaea minor with apical lamella completely rounded, very short, apex wide; Paraphaea formosana with apical lamella somewhat triangular, much longer, apex narrower.

This new species maybe confused with Paraphaea binotata, but can be distinguished by different elytral pattern, and pronotum widest at middle, not at apical third.

Body length usually 5.8–6.9 mm (specimens from Hainan length 7.6–7.9 mm); head and pronotum reddish yellow to brown, lateral explanate areas of pronotum paler; mouthparts, antennae and legs yellowish brown or reddish brown; elytra reddish yellow, with dark marginal fascia occupying 7th–9th intervals, fascia conjoined on elytral apex but not on base, elytral lateral margins yellowish brown; ventral side yellowish brown to dark brown. Head glabrous, without punctures or microsculpture. Pronotum cordiform, widest near middle; ratio PW/PL 1.38 to 1.52; pronotal base briefly but distinctly lobed; disc glabrous, microsculpture indistinct, without punctures; front angles with a few short setae; lateral margins completely rounded in middle, sinuate before hind angles; front angles wide and round, not projected; hind angles sharp, rectangular or nearly so, slightly pointed; disc moderately convex; basal foveae wide and even, with a few fine punctures. Elytra with striae distinct, finely punctate; intervals slightly convex, without punctures or microsculpture; 3rd interval with two setigerous pores: basal one placed approximately basal one-third, adjacent to 3rd stria; apical one placed approximately apical one-eighth, adjacent to 2nd stria or in middle of interval; 5th interval with one setigerous pore near base, adjacent to 5th stria; 15-17 umbilical pores on 9th interval. Male genitalia. Median lobe of aedeagus with apical third slightly expanded; in dorsal view, left-lateral margin moderately sinuate medially, and then gradually narrowed to apex; apical lamella placed on left-ventral side, broadly rounded, not prolongate; internal sac with main flagellum slightly sinuous; basal part of main flagellum moderately bent, so trumpet-form expansion adjacent to left margin, apical margin of trumpet-form expansion not crenulate (Fig. 69). Female genitalia. Spermatheca slender and strongly bent near middle, but not so elongate as in Paraphaea binotata; spermathecal gland inserted near apical one-third of spermatheca, distinctly expanded at basal part, not bent; spermatheca with ring-sculpture between apex and the gland insertion; spermatheca strongly bent at apical one-third (Fig. 127). Apical segment of ovipositor scimitar-shaped, inner margin slightly angulate at apical one-third; length about three times basal width; inner margin setose in apical half; apex slightly widened, with membranous extension long and slender (Fig. 112).

(Map 2). China (Hainan), Vietnam, Thailand, Myanmar, Cambodia.

This species is the smallest one of the genus, so we name it “minor”, a word from Latin meaning “small”.

Two females from Hainan (China) (Fig. 3) are distinctly larger than the other specimens from South East Asia (7.6–7.9 mm contrasting with 5.8–6.9 mm). But there is no other difference between them and other paratypes from Tonkin in external and female genitalia characters, so we include these two females in type series.

Holotype of Coptodera philippinensis Jedlička, by monotypy (NHML): female, body length = 7.6 mm, board mounted, with antennae missing except scapes, “Type” [round label with red ringed]; “Philippine Is. / Coll. Bottcher. / B. M. 1929-201.”; “217”; “Coptodera / philippinen- / type sis sp. n. / Det. ING. JEDLIČKA” [pink label]; “loan from / Brit. Mus.”; “Coptodera / philippi- / ♂ ensis / det. Jedl/ 97 / George E. Ball”; “Anchista / philippinensis / det. Jedlicka / D. Shpeley 1993”[Figs 36, 113, 128].

Shpeley and Ball (1993) mentioned the holotype is a male, but actually it is a female (Fig. 36).

Pronotum sparsely pubescent, widest at apical two-fifths, lateral margins slightly sinuate before hind angles; head and pronotum yellowish to reddish brown, elytra metallic blue; elytra with two or three primary setigerous pores on 3rd interval and one pore on base of 5th interval, intervals with a few very fine secondary setigerous pores.

By the metallic blue elytra and sparse secondary setae on pronotum and elytra, this species can be readily distinguished from other allied species.

Body length 7.6 mm; head reddish brown, pronotum, mouthparts and legs yellowish brown, apices of mandibles darker; elytra metallic blue with 1st and 2nd intervals slightly reddish, lateral margins, elytral suture and epipleurae reddish brown; ventral side brownish. Head nearly glabrous (only two very fine accessory setae visible), without punctures or microsculpture. Pronotum cordiform, widest at apical two-fifths; ratio PW/PL 1.52; pronotal base weakly lobed; disc sparsely and finely setose, lateral explanate areas and area along median line glabrous, microsculpture indistinct, without punctures; front angles with a few setae, distinctly longer than in Paraphaea binotata; lateral margins completely rounded in the widest part, slightly sinuate before hind angles; front angles widened and rounded, not projected; hind angles obtuse; disc moderately convex; basal foveae wide and even, with a few punctures. Elytra with striae very shallow, almost completely composed of rows of fine punctures; intervals flat, without punctures or microsculpture, very fine and sparse accessory setae present on all intervals; 3rd interval with two or three primary setigerous pores (in the holotype, two pores on left elytron, three pores on right one), first one placed at basal one-fourth, adjacent to 3rd stria, the last one placed at apical one-eighth, adjacent to 2nd stria, the middle one, if present, just behind the basal one, at basal one-third approximately; 5th interval with one pore near base, adjacent to 5th stria; 17-18 umbilical pores in 9th interval. Male genitalia unknown. Female genitalia. Spermatheca not distinctly bent in middle; spermathecal gland inserted near basal one-third of spermatheca, its base slightly expanded, with a very short branch; apical part of spermatheca after the gland insertion with distinct ring-sculpture, slightly expanded to fusiform; basal part before the gland insertion without sculpture, slightly bent (Fig. 128). Apical segment of ovipositor scimitar-form, inner margin slightly angulate at apical fourth; length 4 times basal width; inner margin setose in apical fourth; apex narrow, with membranous extension long and slender (Fig. 113).

(Map 2). Only known from the type locality, the Philippines.

This species is only known from the holotype. It was originally described in genus Coptodera and later moved to Allocota by Jedlička (1935b, 1963). Shpeley and Ball (1993) indicated that it is an Anchista species. Kirschenhofer (1996) overlooked Shpeley and Ball’s (1993) comments, and kept it in Allocota. Based on the pronotal shape and female genitalia characters, this species is most similar to Paraphaea minor sp. n., so we herein move it into Paraphaea.

Anchista modesta Nietner, 1856 [= Anchista brunnea (Wiedemann)], by monotypy.

Mandibles moderately widened (Fig. 149); pronotum with lateral margins slightly angulate in middle (Fig. 154), mid-lateral setae present; elytral 5th interval with one to four setigerous pores near base; terminal sternum of males moderately emarginate, with one pair of setae; median lobe of aedeagus usually strongly setose around apical orifice, internal sac without flagellum.

This genus is most closely related to Paraphaea Bates and Metallanchista gen. n. Comparison of these genera is presented in the key to genera and in the diagnosis of Paraphaea and Metallanchista.

Dorsal side generally reddish brown to dark brown; elytra unicolored or bicolored. Head glabrous or sparsely pubescent; eyes hemispherical and strongly prominent; tempora shorter than half length of eyes, strongly narrowed behind eyes; vertex flat. Antennae extended to elytral base; 1st antennomere gradually narrowed to base, 3rd slightly longer than 4th. Labrum smooth, without secondary setae; mandibles moderately widened, outer margin nearly straight (Fig. 149), glabrous on outer scrobe and dorsal ridge; terminal maxillary palpomeres fusiform in both sexes; terminal labial palpomeres strongly securiform, apex truncate in males, less widened in females; ligula with apex slightly projected, with four long setae; paraglossae membranous, not longer than ligula, adnate; mentum tooth simple, with two setae near base; submentum with two long setae; genae glabrous or sparsely pubescent beneath eyes. Pronotum slightly wider than head, disc glabrous or pubescent; mid-lateral setae present; front angles more or less setose, hind angles generally with a few additional short setae; pronotal base briefly but distinctly lobed; lateral margins slightly angulate in middle (Fig. 154), distinctly sinuate before hind angles; hind angles sharp, rectangular or nearly so. Elytra wide, apex truncate, sutural angles not projected, outer angles evenly rounded; laterally slightly depressed in basal one-third, disc with an indistinct depression near basal two-fifths; umbilical pores of 9th interval placed in one row (Fig. 147); basal margination nearly complete or only reaching to 3rd interval; basal pores well developed; 3rd interval with two to four primary setigerous pores, 5th interval with one to four primary setigerous pores near base; elytra glabrous or pubescent; 7th and 8th intervals slightly tumid near apex. Ventral side nearly glabrous; males with apex of terminal sternum moderately emarginate, with one pair of setae; females with apex of terminal sternum straight or slightly emarginate, with two pairs of setae. Legs short; protibiae with cleaning spur well developed, quite distant from inner margin; tarsi widened, 4th tarsomere bifid, claws pectinate; males with adhesive hairs well developed (two whole rows) on 1st to 3rd protarsomeres; well developed or rudimentary (two rows but very weakly present near apex) on first three mesotarsomeres; absent on metatarsomeres. Male genitalia with median lobe of aedeagus nearly straight, not twisted; apical orifice opened dorsally, basal margin of apical orifice setose or glabrous; internal sac without flagellum or apical bursa, with weakly sclerotized sclerites and spined areas on internal sac. Female genitalia. Spermatheca tubular, not bent, with indistinct ring-sculpture, inserted on bursa copulatrix; spermathecal gland slender and long, inserted near apex of spermatheca. Apical segment of ovipositor scimitar-form, curved to outer side, inner margin slightly angulate near apex; with fine fluff near apex; apex with elongate membranous extension.

(Map 3). This genus includes four species. Anchista pilosa is known only from the type locality (Bangalore, south India). The other three species are found in many localities in Indian Subcontinent, with Anchista fenestrata also occurring in Myanmar.

The relationship between Anchista and Paraphaea is discussed under Paraphaea. The monophyly of Anchista is suggested by the following apomorphic character states: (1) pronotum slightly angulate in middle; (2) median lobe of aedeagus with main flagellum reduced, usually setose around apical orifice; (3) spermathecal gland inserted near apex of spermatheca.

Based on setose aedeagal apical orifice and mandibles moderately widened, Anchista could be closely related to Paraphaea Bates, although they have a different aedeagal internal sac, male secondary sexual character and distribution center. Anchista, together with Metallanchista gen. n., is unique in Physoderina in having an aedeagal internal sac without flagellum. The slightly sclerotized area near the median lobe base shows a reduced trumpet-form expansion (Fig. 71), so we believe these three genera are closely allied.

Key to species of Anchista Nietner

Lectotype of Lebia brunnea Wiedemann, designated herein (ZMUC): female, body length = 7.4 mm, pin mounted, “Mus. / Westerm.”; “Type” [red label]; “Bengal. / Mai 1809. / D: brunneus. /Lebia brun / nea Wied. “; “LECTOTYPE ♀/ Lebia brunnea / Wiedemann, 1823 / Des. SHI H. L. 2011” [red label][Fig. 37]. Lectotype of Anchista picea Chaudoir, designated herein (MNHN), male, body length = 7.2 mm, pin mounted, “Ex Musaeo, Chaudoir” [red letter]; “TYPE” [red label]; “MUSEUM PARIS / 1952 / COLL. R. OBERTHÜR”; “picea Chaudoir, Deccan, Stevens” [large box label but pinned under specimen]; “LECTOTYPE ♂/ Anchista picea / Chaudoir, 1877 / Des. SHI H. L. 2011” [red label][Fig. 38]. Paralectotype of Anchista picea Chaudoir (MNHN): 4 males, “Ex Musaeo, Chaudoir” [red letter]; “PARATYPE” [red label]; “MUSEUM PARIS / 1952 / COLL. R. OBERTHÜR”; “PARALECTOTYPE ♂/ Anchista picea / Chaudoir, 1877 / Des. SHI H. L. 2011” [red label] [Figs 4, 70, 97].

Lebia brunnea Wiedemann: In the original literature, Wiedemann didn’t mention how many specimens belonged to the type series. We borrowed one female (Fig. 37) from ZMUC with a red label “Type” and Wiedemann’s hand-written label. We herein designate this specimen as lectotype for taxonomic purpose of fixing the name to unique name-bearing type.

Anchista picea Chaudoir: Five specimens were mentioned in the original description. In the collection of MNHN, these specimens used to be in Chaudoir’s box, but were moved to Lebiini sorted boxes by J. Mateu. Obviously, Mateu selected the first one, and put Chaudoir’s box label and a red “type” label under it. We herein designate this specimen (Fig. 38) with such labels as lectotype for this species for taxonomic purpose of fixing the name to unique name-bearing type.

Non-type material examined (Total 7 specimens from India). 1 specimen (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, XI-XII 1896”. 1 specimen (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, VIII-IX 1896”. 2 specimens (NHML), “India Nevinson Coll. 1918–14.”. 1 specimen (NHML), “Fyzabad, Unt. Prov., India. R. W. G. Hingston.”; “Anchista brunnea Wied (= picea Chaud.), H. E. Andrewes det.”. 1 specimen (NHML), “Nagpur., C. P. India, 1000ft.”; “Anchista brunnea Wied, = picea Chaud., Compared with 2 types H. E. A.”. 1 female (NMPC), “Dabhalwala, Dehra Dun, India”.

Head and pronotum glabrous; pronotum widest at middle, lateral margins slightly angulate in middle; elytra uniform brownish; 3rd interval of elytra with two pores, 5th interval with one pore near base.

Based on the glabrous surface and number of setigerous pores on 3rd and 5th intervals, this species can be readily distinguished from all the other species of Anchista. Other species with the same pore number all belong to different genera (Paraphaea, Metallanchista gen. n.). They are different in elytral pattern and pronotum shape and have other generic differences.

Body length 7.2–7.8 mm; head and pronotum reddish brown; antennae and mouthparts yellowish brown or reddish brown, 1st antennomere and apices of terminal maxillary and labial palpomeres slightly paler; pronotum with lateral explanate area yellowish brown; elytra dark brown, without distinct pattern, but usually with basal two-thirds of disc slightly paler; ventral side same color as dorsal side; femora yellow, tarsi and tibiae yellowish brown. Head glabrous, without punctures or microsculpture. Pronotum glabrous, cordiform, widest slightly before middle; ratio PW/PL 1.38 to 1.41; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles widened and rounded, not projected, with a few short setae; lateral margins slightly angulate in middle, strongly sinuate before hind angles; lateral explanate areas wide and even, with a few coarse punctures; hind angles sharp, rectangular or slightly acute, with a few very short accessory setae; basal foveae moderately deep, with a few punctures; median line distinct, strongly depressed and punctate anteriorly and subbasally. Elytra with striae shallow, with fine punctures; intervals hardly convex, without accessory setae, sparsely and finely punctate; microsculpture distinct, isodiametric; 3rd interval with two setigerous pores, basal one placed at basal one-third approximately, adjacent to 3rd interval, apical one placed at apical one-eighth approximately, adjacent to 2nd stria; 5th interval with one setigerous pore near base, adjacent to 5th stria; basal margination nearly complete. Males with adhesive hairs well developed (two whole rows) on 1st to 3rd mesotarsomeres. Male genitalia with median lobe of aedeagus straight and flat, slightly expanded to apex; left-lateral margin curved in middle; apical orifice opened dorsally, basal margin of apical orifice setose; apical lamella placed on right side, flat, short and rounded in dorsal view, slightly curved upward dorsally in lateral view; internal sac with two elongate and sinuate weakly sclerotized sclerites near apical orifice, basal part of internal sac finely scaled (Fig. 70). Female genitalia not studied.

(Map 3). India, Bengal, Sri Lanka.

Andrewes (1924a, 1927b) indicated that Anchista modesta Nietner is a synonym of Lebia brunnea Wiedemann. We didn’t examine the type of Anchista modesta, which should be in the Warsaw Museum, nor any material from Sri Lanka. We follow Andrewes’ treatment in the present paper.

Holotype of Plochionus fenestratus Schmidt-Göbel, monotypy (NMPC): male, body length = 7.7 mm, board mounted, “MUS. PRAGENSE / COLL. HELFER”; “Birma / Helfer.”; “Typus! teste / Dr. J. Obenberger” [red label]; “fenestratus / Sch. g. /COL. HELFER”[Fig. 39]. Holotype of Anchista glabra Chaudoir, monotypy (MNHN): female, body length = 8.2 mm, pin mounted, “Ex Musaeo / Chaudoir” [red letter]; “TYPE” [red label]; “Museum Paris / 1952 / Coll. R. Oberthür”; “glabra Chaudoir/ Coromandel / Pondichéry / Guerin” [box label but pinned under specimen]; “HOLOTYPE ♀/ Anchista glabra / Chaudoir, 1877 / Des. SHI H. L. 2011” [red label][Fig. 40].

Plochionus fenestratus Schmidt-Göbel: One female was mentioned in the original description. In NMPC boxes of Schmidt-Göbel’s collection, we only found a male determined by him (Fig. 39). This male is the holotype, and Schmidt-Göbel determined the sex erroneously. In the label or original description, no precise locality for this specimen was mentioned. From the record of Helfer’s expedition (Reichardt 1880), it can be inferred that this specimen was collected by Helfer from Tenasserim in Burma (Myanmar).

Anchista glabraChaudoir: Only one specimen was mentioned in the original description. Therefore the one in Chaudoir’s collection is the holotype (Fig. 40).

(Total 25 specimens). India: 1 male and 1 female (MNHN), “S. India, Nedungadu, P. S. Nathan, 1936”. 1 male (MNHN), “Ramnad, Hindonstan”. 17 specimens (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, XI-XII 1896”[Fig. 71]. 2 specimens (NHMB), “India: Orissa state, Similipal N. P. Lulung 21°56'N, 86°32'E, 25.v.–13.vi.1998, Karel & Simon Majer leg”. Nepal: 2 males and 1 female (NHMB), “E-Nepal, Koshi, M. Brancucci”; “Simraghat, 500m, 13.VI.85”[Figs 5, 6, 114, 129].

Head and pronotum glabrous; pronotum widest at middle, lateral margins slightly angulate in middle; elytra distinctly bicolored; 3rd interval of elytra with three to five setigerous pores, 5th interval with two to four setigerous pores near base.

By the glabrous surface and number of setigerous pores on 3rd and 5th intervals, this species can be readily distinguished from most allied species, except Anchista nubila. Diagnosis between these two species is presented in the key.

Male genitalia characters show this species to be very similar to Anchista brunnea, but the latter has the apical lamella wider, internal sac without strongly scaled area near middle, and small sclerite near to the base. From the external characters, these two species can be readily distinguished by the different number of setigerous pores on 3rd and 5th intervals.

Body length 7.7–8.5 mm; dorsal side yellowish to reddish yellow, pronotum lateral explanate area paler; antennae yellowish brown to reddish brown, 1st antennomere slightly paler. Elytra with variable bicolored pattern: darker color brownish to black while the paler parts yellowish to reddish yellow; sometimes elytra mostly dark, but left lateral margin and outer part of apex paler, 2nd to 6th intervals with an elongate pale patch occupying basal three-fifths as in the holotype of Anchista glabra Chaudoir (Fig. 40); or elytra mostly pale, with a small ovaloid or sub-diamondoid dark patch near apex as in some specimens from Nepal (Fig. 6); but in most specimens, elytra with pattern between these two extremities (Fig. 5). Ventral side yellowish to reddish yellow; femora yellow, tarsi and tibiae yellowish brown. Head glabrous, without microsculpture, vertex with a few fine punctures. Pronotum glabrous, cordiform, widest slightly before middle; ratio PW/PL 1.41 to 1.51; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with a few moderately long setae; lateral margins slightly angulate in middle, strongly sinuate before hind angles; lateral explanate area wide and even, with a few coarse punctures; hind angles sharp, rectangular or slightly acute, with a few very short accessory setae; basal foveae moderately deep, with a few punctures; median line distinct, strongly depressed and punctate anteriorly and subbasally. Elytra with striae shallow but distinct, finely punctate; intervals slightly convex, without accessory setae, finely and sparsely punctate; microsculpture distinct, isodiametric; 3rd interval with three to five setigerous pores, the basal one usually placed at basal one-third approximately, adjacent to 3rd stria or on middle of interval, the rest adjacent to 2nd stria; 5th interval with two to four setigerous pores, the basal one near interval base, the last one usually at middle, all pores usually in center of 5th interval or slightly adjacent to 5th stria; basal margination nearly complete. Male with adhesive hairs rudimentary (two rows, weakly present near apex) on first three mesotarsomeres. Male genitalia with median lobe of aedeagus straight and flat, slightly expanded to apex; left lateral margin slightly curved in middle; apical orifice opened dorsally, basal margin of apical orifice setose; apical lamella placed on right side, flat, slightly elongate and rounded in dorsal view, slightly curved up dorsally in lateral view; internal sac with a small strongly sclerotized sclerite near base, two weakly sclerotized elongate and sinuate sclerites near apical orifice, a small and strongly scaled area present near middle of median lobe, internal sac in basal half sparsely and finely setose ventrally (Fig. 71). Female genitalia. Spermathecal gland inserted near apex of spermatheca, strongly expanded but not bent at base; spermatheca gradually expanded to apex, apex globular with a small papilla; ring-sculpture very faint, only present on subapical part of spermatheca (Fig. 129). Apical segment of ovipositor scimitar-form, inner margin gradually curved; length nearly three times basal width; inner margin setose in apical half; apex rounded and slightly projected to outer side, with membranous extension slightly widened.

(Map 3). India, Myanmar, Nepal.

We dissected males from Nedungadu (close to type locality of Anchista glabra), Chota Nagpore, Nepal Koshi (type locality of Anchista nepalensis), and the holotype of Anchista fenestrata. There is no significant difference among these specimens from different localities, except for elytral pattern variation. We therefore herein synonymize Anchista glabra and Anchista nepalensis with Anchista fenestrata.

In this species, specimens from one locality may be obviously different in elytral pattern. Sometimes, intrapopulational variation ranges even wider than interpopulational. But statistically interpopulational pattern variation exists: individuals from south India usually with dark pattern occupying more area on elytra (e. g. in the holotype of Anchista glabra Chaudoir, Fig. 40) than those from north India; while some specimens from Nepal have the paler color occupying the largest area, with only a small dark patch near apex (Fig. 6).

Lectotype of Anchista subpubescens Chaudoir, designated herein (MNHN): female, body length = 7.8 mm, pin mounted, “Ex Musaeo / Chaudoir” [red letter]; “TYPE” [red label]; “subpubescens / Chaudoir / Indes orient bor / Bacon” [Chaudoir’s box label, but pinned under specimen]; “Museum Paris / 1952 / Coll. R. Oberthür”; “LECTOTYPE ♀/ Anchista subpubescens / Chaudoir, 1877 / Des. SHI H. L. 2011” [red label][Fig. 41].

Anchista subpubescensChaudoir: We found a female in the general Lebiini collection in MNHN, it was moved from the collection of Chaudoir. Except this one, we found no other specimen that accorded with the original literature in MNHN. The original literature didn’t indicate or imply this species was described on a single specimen, so we herein designate this specimen (Fig. 41) as lectotype for the purpose of fixing the name to unique name-bearing type.

(Total 33 specimens). Pakistan: 4 females (ZSM), “West Pakistan, Rawalpindi Umg., 25 km No., 600-, 700m, 20.XII.55, Chr. Lindemann leg.”. 1 female (ZSM), “West Pakistan, Rawalpindi Umg., Kanatti Chak, Salt Range, 9–14.I.56, Chr. Lindemann leg.”. 15 females, 8 males (ZSM), “West Pakistan, 11-, Rawalpindi Umg., Kanatti Chak, Salt Range, 15.II.56, Chr. Lindemann leg.” [4 of them with the label: “Anchista subpubescens, det. Ing Jedlička”][Figs 7, 72]. 3 females (NMPC), “West Pakistan 11.-, Rawalpindi Umg., Kanatti Chak, Salt Range 15.II.56, Chr. Lindemann leg.”. India: 2 females (CMB), “India, Rajasthan, Bharatpur 40km, w Agra N. P., Keoladeo leg., 11/07 Roppei”.

Dorsal side evenly pubescent; pronotum widest near middle, lateral margins slightly angulate; elytra bicolored; 3rd and 5th interval of elytra with primary pores small, indistinct, usually more than five pores on each interval.

Body size, form, and color the same as the nominotypical subspecies, also with similar elytral pattern variation. This subspecies differs from the nominotypical subspecies by: dorsal side finely and equally pubescent; microsculpture indistinct; head with pubescence sparse on vertex; elytra with intervals evenly pubescent; 3rd interval with five to eight setigerous pores, some pores small and indistinct, the basal one usually placed at basal one-fourth approximately, apical one adjacent to 2nd stria; 5th interval with five to seven setigerous pores at basal half, pores gradually decreasing to apex, sternum with dense pubescence. Male genitalia without significant difference from the nominotypical subspecies (Figs 71, 72).

(Map 3). India (Rajasthan), Pakistan.

Anchista subpubescens Chaudoir was originally described from north India. According to specimens examined, it is distributed in northeast Pakistan and northwest India, allopatric with Anchista fenestrata from east India (Map 3). All specimens of Anchista subpubescens have no significant difference in male genitalia and external structure exceptdistinct dorsal pubescence. We therefore rank it as a subspecies of Anchista fenestrata (Schmidt-Göbel).

Lectotype of Anchista nubila Andrewes, designated herein (NHML): female, body length = 8.5 mm, board mounted, “Dehra Dun / Rajpur, 3404 ft.”; “Bought from / Staudinger & / Bang-Haas, 1930”; “Type” [red label]; “Anchista / nubila / Type Andr. / H. E. Andrewes det.”; “H. E. Andrewes Coll. / B. M. 1945–97”; “LECTOTYPE ♀ / Anchista nubila / Andrewes, 1931 / des. SHI H. L. 2011” [Fig. 42]. Paralectotypes of Anchista nubila Andrewes: 1 male and 2 females (NHML), “Dehra Dun / Rajpur, 3404 ft.”; “Bought from / Staudinger & / Bang-Haas, 1930”; “Cotype” [round label, with green circle]; “H. E. Andrewes Coll. / B. M.1945–97”; “PARALECTOTYPE / Anchista nubila / Andrewes, 1931 / det. SHI H. L. 2011”.

The original description mentioned five specimens from Dehra Dun in Andrewes’ collection, but did not designate a holotype. In the collection of NHML, we found only four specimens. Among them, a female specimen bears a determination label of Andrewes and a red label “Type”, and this is herein designated as lectotype (Fig. 42) for taxonomic purpose of fixing the name to unique name-bearing type. The other three specimens have only a round label “co-type”, but no Andrewes’ determination label.

(Total 16 specimens from India). 9 specimens (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, VIII-XI 1896”[Fig. 73]. 3 specimens (MNHN), “Chota Nagpore, Nowatoli, R. P. Cardon, V.VI.1896”. 1 specimen (MNHN), “Inde Angl, Ghates, R. P. F. Tabourel, VII-IX 1898”. 1 specimen (MNHN), “S. India, Nedungadu, P. S. Nathan, 1936”[Fig. 8]. 1 specimen (MNHN), “Chota Nagpore”; “Ex Musaeo, H. W. Bates, 1892”. 2 specimens (NHML), “India mer. Tanjore Distr. Nedungadu”.

Head and pronotum glabrous; pronotum widest near middle, lateral margins distinctly angulate at middle; elytra uniform brownish; 3rd interval of elytra with three or four pores, 5th interval with two or three pores near base.

The glabrous surface and the number of setigerous pores on the 3rd and 5th intervals readily distinguish this species from most allied species, except Anchista fenestrata fenestrata. Diagnosis between these two species is presented in the key. In general appearance, this species looks like Anchista brunnea, but can be distinguished from the latter by the wider and more angulate pronotum, and the different number of elytral setigerous pores.

Body length 7.5–8.5 mm; dorsal side uniform reddish brown, pronotum lateral explanate areas somewhat paler; elytra usually with margins darker, but not forming central patch; antennae reddish brown, 1st antennomere paler; femora yellowish, tibiae and tarsi darker; ventral side reddish brown. Head glabrous, without or with very fine isodiametric microsculpture. Pronotum glabrous, cordiform, widest slightly before middle; ratio PW/PL 1.30 to 1.41; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with few short setae (distinctly shorter than in Anchista fenestrata fenestrata); lateral margins distinctly angulate in middle, strongly sinuate before hind angles; lateral explanate areas wide and flat, with few coarse punctures; hind angles sharp, rectangular or slightly acute, with few very short accessory setae; basal foveae moderately deep, with few punctures; median line distinct, strongly depressed and punctate anteriorly and subbasally. Elytra with striae shallow, distinct, finely punctate; intervals slightly convex, without accessory setae, finely and sparsely punctate; microsculpture distinct, isodiametric; 3rd interval usually with three setigerous pores, sometimes with four, all adjacent to 2nd interval, basal one usually placed at basal one-fourth approximately; 5th interval usually with two setigerous pores, sometimes with three, basal one near to interval base, all pores adjacent to 5th stria; basal margination nearly complete. Males with adhesive hairs rudimentary (two rows weakly present near apex) on 1st mesotarsomere, well developed (two whole rows) on 2nd and 3rd mesotarsomeres. Male genitalia with median lobe of aedeagus stout, bent to right side near apex; in lateral view, distinctly expanded in middle, gradually narrowed to apex; left lateral margin not curved in middle; apical orifice opened dorsally, basal margin of apical orifice with long setae; apical lamella placed on right side, flat, wide and rounded in dorsal view, not curved up dorsally; internal sac with a weakly sclerotized sclerite near middle, two weakly sclerotized elongate and sinuate sclerites near apical orifice, some parts of internal sac irregularly scaled (Fig. 73). Female genitalia not studied.

(Map 3). India.

Holotype (MNHN): male, body length = 7.8 mm, board mounted, genitalia dissected and deposited in microvial pinned under specimen, “Bangalore / Chikkangalur / Tabourel 1900”; “MUSEUM PARIS / 1952 / COLL. R. OBERTHÜR”; “HOLOTYPE ♂/ Anchista pilosa / new species / Des. SHI H. L. 2011” [red label][Figs 43, 74, 98].

Pronotum pubescent, lateral margins hardly angulate near middle, sinuate before hind angles; elytra with bicolored pattern; elytra pubescent, primary setigerous pores not distinct; elytral basal margination extended only to 3rd interval; median lobe of aedeagus without setae around apical orifice.

Based on pubescent surface and elytral pattern, the new species resembles the subspecies Anchista fenestrata subpubescens Chaudoir, but can be distinguished in having the following characters: (1) elytral basal margination extended only to 3rd interval, but in Anchista fenestrata subpubescens basal margination nearly complete; (2) pronotum with lateral margins less angulate and hind angles less distinct in the new species; (3) males with tarsi adhesive hairs well developed (two whole rows) on first three mesotarsomeres, but in Anchista fenestrata subpubescens adhesive hairs rudimentary (weakly present near apex) on first three mesotarsomeres in males; (4) males of the new species with terminal sternum more deeply emarginate than in Anchista fenestrata subpubescens; (5) median lobe of aedeagus without setae around apical orifice, but in Anchista fenestrata subpubescens median lobe of aedeagus with long setae around apical orifice.

Body length 7.8 mm; dorsum yellow to reddish yellow, pronotum lateral explanate areas pale yellow; 1st antennomere yellow, remaining antennomeres reddish brown. Elytra bicolored, forming faint pattern, background nearly brownish, with lateral part slightly paler; large yellow patch occupying inner five intervals from base to apical third; lateral margins and epipleura yellow. Ventral side yellow, sterna darker toward apex; femora and tibiae yellowish, tarsi yellowish brown. Head sparsely pubescent, nearly glabrous on front, without microsculpture or punctures. Pronotum with disc pubescent, rather sparser along median line, nearly glabrous on lateral explanate areas;pronotumcordiform, widest slightly before middle; ratio PW/PL 1.53; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with few moderately long setae; lateral margins hardly angulate in middle, strongly sinuate before hind angles; lateral explanate areas wide and even; hind angles slightly obtuse, with few accessory setae; basal foveae moderately deep, with few punctures; median line indistinct, slightly depressed subbasally. Elytra with striae shallow, distinct, slightly punctate along striae; intervals slightly convex, with dense and regular accessory setae; microsculpture indistinct; primary setigerous pores on 3rd and 5th intervals hardly visible, five pores on 3rd interval, two on the 5th; basal margination extended only to 3rd interval. Males with adhesive hairs well developed (two whole rows) on first three mesotarsomeres; males with terminal sternum more deeply emarginate than other members of the genus (as in Fig. 145), but less deeply than in Paraphaea (Fig. 143). Male genitalia with median lobe of aedeagus straight and flat, gently expanded to apex; left lateral margin slightly curved in middle; apical orifice opened dorsally, margin around apical orifice not setose; apical lamella placed on right side, flat and rounded, width subequal to length in dorsal view, not curved upwards dorsally in lateral view; internal sac with two moderately sclerotized flagella near apical orifice, internal sac irregularly scaled and sclerotized, basal part more distinctly sclerotized (Fig. 74). Female genitalia unknown.

(Map 3). Only known from type locality Bangalore (India).

The name “pilosa” comes from Latin, meaning pubescent, referring to the pubescent surface of this species.

Based on median lobe of aedeagus without setae, elytral basal margination incomplete and male terminal sternum more deeply emarginate than other species, the new species seems to be isolated in the genus. But, male genital internal sac characters and distribution of male adhesive hairs on tarsi indicated that this species is a member of Anchista.

Metallanchista laticollis Shi & Liang, sp. n.

Surface glabrous, with distinct metallic reflection; mandibles moderately widened (Fig. 149); umbilical pores of 9th interval placed in two rows (Fig. 148); males with terminal sternum deeply emarginate, bisetose on each side; males without adhesive hairs on metatarsomeres; median lobe of aedeagus without setae around apical orifice; internal sac with main flagellum reduced. These characters readily distinguish the new genus from all other genera of Physoderina.

The gen. n. is most closely allied to Anchista, but can be distinguished by: (1) median lobe with apical lamella longer and bent dorsally; (2) males with two pairs of setae on terminal sternum; (3) umbilical pores of 9th interval placed in two rows. Metallanchista new genus also resembles some species of Physodera, but differs by the following characters: (1) pronotum with mid-lateral setae; (2) basal margination of elytra nearly complete; (3) elytral intervals more or less convex; (4) 3rd to 5th intervals slightly depressed at basal two-fifths; (5) mandibles moderately widened.

Body rather flat, dorsal side with distinct metallic reflections; microsculpture indistinct. Head glabrous; eyes hemispherical, prominent; tempora short, nearly half as long as eyes, abruptly narrowed behind eyes; vertex flat. Antennae hardly extended to elytral base; 1st antennomere stout, slightly expanded in middle, slightly narrowed to base, 3rd as long as 4th. Labrum smooth, without any secondary setae; mandibles moderately widened, outer margin nearly straight, glabrous on outer scrobe and dorsal ridge; terminal maxillary palpomeres fusiform in both sexes; terminal labial palpomeres moderately expanded but not strongly securiform in both sexes; ligula with apex nearly truncate, with four long setae; paraglossae membranous, slightly longer than ligula, adnate; mentum tooth wide and rounded, with two long setae near base, a few additional short setae present on central area of mentum; submentum with four long setae, distant from lateral margin; genae and gula with a few short seta at anterior part. Pronotum wider than head, disc glabrous; mid-lateral setae present; front angles more or less setose, hind angles generally with a few additional short setae; pronotal base briefly but distinctly lobed; lateral margins rounded or slightly angulate, strongly sinuate before hind angles; hind angles sharp. Elytra wide, apex truncate, sutural angles not projected, outer angles completely rounded; lateral margins slightly depressed in basal one-third, disc with an indistinct depression near basal two-fifths; intervals glabrous, without additional setae; umbilical pores of 9th interval placed in two rows (Fig. 148), the outer row (the primary umbilical pore series) adjacent to lateral expansion usually composed of 18 pores, the inner row (secondary umbilical pore series) adjacent to 8th stria usually composed of 10 pores; basal margination nearly complete; basal pores well developed; 3rd interval with two setigerous pores, basal one located at basal one-third approximately, adjacent to 3rd stria, apical one at apical one-eighth approximately, adjacent to 2nd stria; 5th interval slightly widened at base, with one setigerous pore near base, adjacent to 5th stria; 7th and 8th intervals slightly tumid near apex. Ventral side nearly glabrous; males with terminal sternum deeply emarginate apically (Fig. 143), bisetose on each side; females with terminal sternum straight apically, bisetose on each side. Legs short; protibiae with cleaning spur well developed, distant from inner margin; tarsi widened, 4th tarsomere bifid, claws pectinate; males with adhesive hairs well developed (two whole rows) on 1st to 3rd pro-, and 2nd to 3rd mesotarsomeres, rudimentary (two rows weakly present near apex) on 1st mesotarsomere. Male genitalia with median lobe of aedeagus not twisted; apical orifice opened dorsally, apical lamella without long setae; apical lamella strongly bent dorsally; internal sac with main flagellum fully reduced, without any distinct sclerite, some parts of internal sac strongly scaled. Female genitalia. Apical segment of ovipositor straight, glabrous; membranous extension short, slightly sclerotized. Internal reproductive system not studied.

(Map 4). Malay Peninsula, Sumatra, Java.

The genus name “Metallanchista” is combined from the Greek “metall”, a reference to metal, and the genus name “Anchista”, meaning that it is closely related to genus Anchista, but strongly metallic. The gender of this genus name is feminine.

The new genus is most closely allied with Anchista, as both genera have the main flagella of the aedeagal median lobe fully reduced. Both are also allied with Paraphaea and Endynomena in sharing: (1) median lobe of aedeagus usually strongly setose around apical orifice; (2) mandibles usually moderately widened; (3) basal margination of elytra usually complete. These character states may be synapomorphies for this lineage in the subtribe.

We establish this new genus based on the following apomorphic character states: (1) umbilical pores of 9th interval placed in two rows; (2) median lobe of aedeagus with apex strongly bent dorsally, internal sac without distinct sclerite, but scaled in certain areas; (3) submentum quadrisetose, ventral side of head with some accessory setae; (4) terminal sternum of males with two pairs of setae on each side.

So far as we know, there is no other genus of Lebiini with two rows of umbilical pores as in Metallanchista gen. n. The pore series in Metallanchista gen. n. contains more pores than in other allied genera, 28 pores contrasting with usually 15–20 pores in other genera. So the peculiar pore series is not the result of a different arrangement, but due to the presence of the unique secondary pore series adjacent to the 8th stria. The outer pore series, composed of 18 pores adjacent to the lateral expansion, is the primary pore series and homologous with the umbilical pore series in other genera.

Holotype (NHMB): male, body length = 7.8 mm, board mounted, genitalia dissected and deposited in microvial pinned under specimen, “Thailand, 1.-20.iii.1996 / Chumphon prov. / Pha To env. 9°48’ 98°47’ / K. Majer leg.”; “HOLOTYPE ♂/ Metallanchista laticollis / new species / Des. SHI H. L. 2011” [red label][Figs 44, 75, 99].

Elytra uniform metallic piceous–green, intervals slightly convex; pronotal width 1.65 times length, lateral margins slightly angulate in middle, front angles with only a few very short setae. This species is similar to Metallanchista perlaeta, but can be distinguished from the latter by pronotal lateral margins angulate in middle and dorsal side with less metallic color.

Body length 7.8 mm; dorsum piceous, head and pronotum with moderate reflection, pronotum slightly brownish, elytra dark green, with strong metallic reflection; antennae, mouthparts yellowish brown, apices of mandibles darker, apices of terminal labial and maxillary palpomeres yellow; femora dark brown, tibiae and tarsomeres reddish brown; ventrum brown. Head glabrous, without microsculpture or punctures. Pronotum strongly transverse, widest slightly before middle; lateral explanate areas very wide and even, with a few punctures; ratio PW/PL 1.65; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures or pubescence; front angles with a few very short setae; lateral margins slightly angulate in middle, strongly sinuate before hind angles; hind angles obtuse, sharp, not projected, with a few accessory setae; basal foveae shallow, with a few punctures; median line distinct, short, not reaching to base. Elytra with striae shallow, distinct, finely punctate; intervals slightly convex; microsculpture absent; 3rd interval with two setigerous pores, the basal one placed at basal one-third approximately, adjacent to 3rd stria, the apical one placed at apical one-eighth approximately, adjacent to 2nd stria; 5th interval with one setigerous pore near base. Male genitalia with median lobe of aedeagus tubular, slightly compressed; left lateral margin not curved, right lateral margin gently expanded in dorsal view; apical orifice opened dorsally, margin around apical orifice without long setae; apical lamella placed on right side, flat and elongate, apex rounded, length twice width; apical lamella strongly bent dorsally in lateral view; subapical area finely setose on right side. Internal sac without distinct sclerite; two separate areas near to dorsal margin strongly and coarsely scaled, basal one smaller and more coarsely scaled; main flagellum and trumpet-form expansion reduced to a hardly visible semi-sclerotized sclerite near base (Fig. 75). Female genitalia unknown.

(Map 4). Only known from the type locality PhaTo (Thailand).

The name “laticollis” is from the Latin “lat-” meaning wide and “coll” meaning neck, referring to the pronotum. This species is named for its rather wide pronotum.

The holotype, a female, came from Langkawi Island. We have not examined it.

(Total 2 females from Indonesia) 1 female (CRS), “Sumatra C., Padang 3. 97, Panjang”[Fig. 9]. 1 female (NHML), “Java”; “Bowring., 63 47*”[Fig. 10].

Elytral color various, metallic violaceous, green, or cupreous, but always with distinct bluish color along lateral margins; pronotal width 1.50 times length, lateral margins completely rounded in middle, front angles with moderately long setae; elytral intervals hardly convex. This species is close to Metallanchista laticollis sp. n., but can be distinguished from it by different color and pronotal shape.

Body length 9.1–9.9 mm; head nearly black, with luster, antennae dark brown; mouthparts reddish brown, apices of terminal palpomeres yellow; pronotum black, with luster, slightly bluish in lateral expansion; elytra strongly metallic, disc violaceous, green, or cupreous, sometimes with basal and lateral areas bluish, but at least lateral margins distinctly bluish; epipleurae yellowish brown with metallic blue; ventral side dark brown to black, slightly lustrous; legs black, tarsomeres brown. Head glabrous, without microsculpture or punctures. Pronotum rounded, widest slightly before middle; lateral explanate areas very wide, flat, with a few punctures; ratio PW/PL 1.50; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures or pubescence; front angles with some moderately long setae; lateral margins completely rounded in middle, strongly sinuate before hind angles; hind angles slightly acute, distinctly but slightly projected, with a few accessory setae; basal foveae shallow, with a few punctures; median line shallow, not reaching to apical or basal margin. Elytra with striae barely sulcate, composed of fine punctures, but slightly deepened in middle of 6th stria; intervals hardly convex, without punctures or microsculpture; 3rd interval with two setigerous pores, basal one placed at basal one-third approximately, adjacent to 3rd stria, apical one placed at apical one-eighth approximately, adjacent to 2nd stria; 5th interval with one setigerous pore near base. Male genitalia unknown. Female genitalia with apical segment of ovipositor slender and straight, without setae, length about four times basal width; widest at base, gradually narrowed to apex, apex sharp; inner and outer margin nearly straight; extended part on apex slightly sclerotized, very short. Spermathecal characters not studied.

(Map 4). Malay Peninsula, Sumatra, Java.

So far only the three females of this species from different localities are known to us. They are different in color from each other, probably representing geographical variation. The holotype from Malay Peninsula has the elytra cupreous, apex violaceous, lateral margin slightly blue (by original description). The second female is from Sumatra and has the elytra violaceous on most areas, laterally with bluish band occupying 7th to 9th intervals, and the two bands joined at base. The last one is from Java and has elytra that are fully metallic green, but slightly bluish along lateral margin. More specimens, especially males, are needed for further study.

Physodera dejeani Eschscholtz, 1829, by monotypy.

This genus can be readily recognized by the combination of the following character states: dorsal side usually glabrous, at most elytra with a little secondary pubescence; mandibles strongly widened; pronotum wide, ratio PW/PL more than 1.4; primary mid-lateral setae of pronotum usually absent, if present, lateral margin with numerous accessory setae nearly along full length; protibiae with cleaning spur well developed; elytra with one or two setae near base of 5th interval; median lobe of aedeagus with apical orifice opened apically.

This genus is closest to Allocota, but can be distinguished in having: (1) pronotum and elytra much wider, ratio PW/PL more than 1.4 (usually more than 1.6, but narrower in Physodera eschscholtzii and allied species), but much narrower in Allocota, ratio PW/PL less than 1.4; (2) cleaning spur present on protibia (as in Fig. 142), but usually absent (Fig. 140) in Allocota, if present, very fine (Fig. 141); (3) 5th interval of elytra only with one or two primary pores near base, if additional small pores present, all pores placed before middle, but in Allocota, 5th interval of elytra with four to ten setigerous pores, equally distributed; (4) median lobe of aedeagus strongly bent to right side in dorsal view, but only slightly bent in Allocota; (5) apical segment of female ovipositor strongly elongate, but only a little longer than width in Allocota.

Some species of Anchista, Paraphaea or Metallanchista could be confused with members of this genus, but can be distinguished from Physodera by having the mandibles narrower, primary lateral seta of pronotum present and different shape of the aedeagus.

Body length 8.0 to 12.5 mm, stout, pronotum usually wide, elytra wide and distinctly convex. Dorsal side usually metallic, usually with pattern composed of vivid color, or with ivory callosities on pronotum or elytra. Head and pronotum glabrous, elytra glabrous or with a few accessory setae; microsculpture indistinct. Head glabrous; eyes hemispherical, strongly prominent; tempora shorter or slightly longer than half length of eyes; tempora abruptly or gradually narrowed behind eyes; vertex flat. Antennae extended to basal one-fifth of elytra approximately; 1st antennomere slightly narrowed to base, 3rd slightly longer than 4th. Labrum smooth, without or with a few very short secondary setae; mandibles strongly widened, outer margins rounded, glabrous on outer scrobe, sometimes along dorsal ridge with a few fine and short setae; terminal maxillary palpomeres fusiform in both sexes; terminal labial palpomeres not widened in both sexes, or strongly securiform and truncate apically in males and less widened in females; ligula with apex truncate or slightly projected, with four long setae; paraglossae membranous, not longer than ligula, adnate; mentum tooth simple, apex rounded or truncate, with two setae or without seta near base; submentum with two long setae; genae glabrous beneath eyes. Pronotum distinctly wider than head, disc glabrous; front angles more or less setose, setae usually restricted to front angle, or nearly distributed along full length of lateral margins; mid-lateral seta usually absent; pronotal base more or less lobed; lateral margins usually completely rounded in middle; hind angles sharp or rounded. Elytra wide, strongly convex, apex truncate or slightly curved, sutural angles not projected, outer angles rounded; lateral margins more or less depressed at basal one-third, disc without distinct depression; intervals flat, glabrous or with a little pubescence or additional setae; basal margination reaching only to 3rd or 4th interval; basal pores well developed; primary setigerous pores small, 3rd interval with two to four setigerous pores, sometimes the apical one present on 2nd interval, 5th interval with one or two primary setigerous pores near base; 7th and 8th intervals slightly to strongly tumid near apex. Ventral side nearly glabrous; males with terminal sternum moderately emarginate, with one or two pairs of setae; females with terminal sternum apex straight or slightly curved, with two pairs of setae. Legs short; protibiae with cleaning spur well developed, distant from inner margin; tarsi widened, 4th tarsomere bifid, claws pectinate; males with adhesive hairs on 1st to 3rd protarsomeres well developed (two whole rows), those on 1st to 3rd mesotarsomeres rudimentary (two rows, weakly present near apex) or well developed (two whole rows) on some of these tarsomeres. Male genitalia with median lobe of aedeagus not twisted, strongly bent to right side in dorsal view; apical orifice opened apically; apical lamella usually long; dorsal surface with a few fine setae subapically; internal sac with main flagellum fine, apex nearly reaching to apical orifice, trumpet-form expansion usually small; apical bursa present or absent; secondary flagellum distinct or not. Female genitalia. Spermatheca tubular, with ring-sculpture, inserted at the joining of bursa copulatrix and common oviduct; spermathecal gland much longer than spermatheca, inserted near middle of spermatheca; spermatheca not distinctly bent. Apical segment of ovipositor tapered, apex pointed, base gradually widened, apex with membranous extension fine and long.

Oriental Region: India, Sri Lanka, Nepal, South China, Indochina Peninsula, Malay Peninsula, Malay Archipelago, New Guinea, the Philippines. Not discovered in Japan or Australia.

Physodera appears to be closely related to Allocota, based on the similar aedeagal characters, generally glabrous surface, and pronotum usually without primary mid-lateral seta. Female internal reproductive system of Physodera resembles that of Paraphaea philippinensis (Jedlička), and may suggest a close relationship between these two genera.

Species of Physodera are more morphologically diverse than those in other genera of the subtribe. Physodera dejeani and related species seem to be more distant from other species of Physodera, according to the differences in the terminal labial palpomeres, setae on the terminal sternum and male genital characters. Further study may divide this genus into other subgenera or even genera.

Nevertheless, we suppose species of Physodera, or Physodera + Allocota, form a monophyletic lineage, such as those species previously arranged in Anchista (Anchista + Paraphaea + Metallanchista in the present paper). Monophyly of Physodera could be inferred by: (1) elytra strongly convex and wide; (2) pronotum strongly transverse; (3) terminal sternum or tergum usually with yellowish patch; (4) median lobe of aedeagus strongly bent to right side in dorsal view, apical lamella usually long.

Heller (1923) provided a key for the seven species of Physodera known at that time. Later, Andrewes (1930b, 1930c), Darlington (1971) and Mateu (1990) added some species to the genus. To date, including one new combination (Lachnoderma andrewesi Jedlička), a total of 15 available names has been included in Physodera, with one regarded as a junior synonym and one a subspecies. However, there is no comprehensive taxonomic work on this brilliant beetle group and such a work would be useful.

Lachnoderma andrewesi Jedlička was moved to Allocota by Jedlička (1963) and followed by Kirschenhofer (1996). But, from pronotal shape and elytral setigerous pores, it is far different from all other members of Allocota. After studying the female holotype, we confirmed that it is actually a Physodera species. The supporting characters are present in the diagnosis part of Physodera.

Mateu (1990) proposed the subgenus name Nepalotarsus for Physodera bousqueti Mateu and other species with adhesive hairs present on first three protarsomeres in males, but the type species of the subgenus was not fixed in the original publication. According to the Zoological Code of Nomenclature (4th Edition), Article 13.3, the name Nepalotarsus is a nomen nudum, an unavailable name.

List of species:

Physodera amplicollis van de Poll, 1889: 254. Type locality: Java. Syntype location unknown.

Physodera andrewesi (Jedlička, 1934: 120) (Original: Lachnoderma). Type locality: Mt. Makiling (Philippinens). Holotype deposited in NHML. comb. n.

Physodera bacchusi Darlington, 1971: 326. Type locality: New Guinea. Holotype deposited in NHML.

Physodera bifenestrata Heller, 1923: 304. Type locality: Sandakan (Borneo). Syntype deposited in SNSD.

Physodera bousqueti Mateu, 1990: 156. Type locality: Nepal. Holotype deposited in BRIO.

Physodera chalceres Andrewes, 1930b: 135. Type locality: Penang (Malaya). Holotype deposited in NHML.

Physodera cyanipennis van de Poll, 1889: 253. Type locality: Celebes. Syntype location unknown.

Physodera dejeani Eschscholtz, 1829: 8. Type locality: Manilla (Philippines). Syntype deposited in MNHN.

Physodera diglena Andrewes, 1930c: 202. Type locality: Selangor (Malaya). Holotype deposited in NHML.

Physodera eburata Heller, 1923: 304. Type locality: Los Banos (Luzon). Syntype deposited in SNSD.

Physodera eschscholtzii Parry, 1849: 179. Type locality: Ceylan and Philippines. Syntypes deposited in NHML (probably) and MNHN.

Physodera davidis Fairmaire, 1887: 92. Type locality: Fokien (China). Syntype deposited in MNHN.

Physodera eschscholtzii sumatrensis (Kirschenhofer, 1996: 761) (Original: Allocota). Type locality: Sumatra. Holotype deposited in NHMW

Physodera noctiluca Mohnike, 1875: 154. Type locality: Java. Syntype deposited in MNHN (probably).

Physodera parvicollis van de Poll, 1889: 252. Type locality: Hong Kong. Syntype location unknown.

Diamella kaszabi Jedlička, 1952, by monotypy.

Head and pronotum more or less pubescent; elytra with accessory setae at odd intervals or along striae, even intervals glabrous; vertex distinctly tumid, posterior supraorbital setae distant from eyes, insertions more or less pointed; internal sac of male genitalia with main flagellum extraordinarily thick, apical part of trumpet-form expansion strongly expanded.

In general appearance, this genus resembles Allocota, but the latter is different from Diamella in having: (1) head and pronotum always glabrous; (2) vertex less tumid and posterior supraorbital setae not distant from eyes; (3) lateral setae on pronotal margin absent; (4) males with terminal sternum bisetose on each side; (5) main flagellum of aedeagus finer.

Dorsal side generally brownish to dark brown, elytra usually with metallic reflections; elytra unicolored. Head more or less pubescent, clypeus with some pubescence; eyes hemispherical, strongly prominent; tempora gradually narrowed behind eyes, longer than half length of eye; vertex distinctly tumid, posterior supraorbital setae distant from eyes, insertions more or less prominent, sometimes forming a horn-like hump. Antennae extended to basal one-sixth of elytra approximately; 1st antennomere slightly curved, 3rd slightly longer than 4th. Labrum with faint microsculpture, glabrous or with a few very fine short secondary setae, apex straight or emarginate; mandibles distinctly widened, outer margins rounded (as in Fig. 150), glabrous on outer scrobe, sometimes with a few short setae along dorsal ridge; terminal maxillary palpomeres fusiform in both sexes; terminal labial palpomeres strongly securiform, truncate apically in males, less widened in females; ligula with apex slightly projected, with four long setae; paraglossae membranous, not longer than ligula, narrow, adnate; mentum tooth simple, with two setae near base; submentum with two long setae; genae glabrous beneath eyes. Pronotum slightly wider than head, disc more or less pubescent; lateral margins with setae along full length, sometimes sparser at middle; primary mid-lateral setae present; pronotal base more or less lobed; lateral margins rounded in middle, sinuate before hind angles; hind angles sharp, rectangular or acute. Elytra wide, apex truncate, sutural angles not projected, outer angles completely rounded; anterior one third of sides slightly depressed or not, disc with or without a depression; odd intervals with accessory setae, even intervals glabrous; striae with or without setae; umbilical pores of 9th interval placed in one row; basal margination reaching to 3rd interval; basal pores present; primary setigerous pores indistinct among accessory ones, 3rd and 5th intervals with some larger pores; 7th and 8th intervals more or less tumid near apex. Ventral side with long and dense pubescence except proepisterna; males with apex of terminal sternum moderately emarginate, with 1one pair of setae; females with apex of terminal sternum straight or slightly curved, with two pairs of setae. Legs short; protibiae with cleaning spur well developed, distant from inner margin; tarsi widened, 4th tarsomere bifid, claws pectinate; males with adhesive hairs well developed (two whole rows) on 1st to 3rd protarsomeres, rudimentary (two rows, weakly present near apex) on 1st to 3rd mesotarsomeres. Male genitalia with median lobe of aedeagus not twisted; apical orifice opened apically; internal sac with main flagellum extraordinarily thick, apex nearly reaching to apical orifice, trumpet-form expansion strongly expanded; apical bursa absent; secondary flagellum short and indistinct. Female genitalia. Spermatheca very short, tubular, with distinct ring-sculpture, inserted on bursa copulatrix; spermathecal gland much longer than spermatheca, inserted distal to the middle of spermatheca; spermatheca indistinctly bent. Apical segment of ovipositor scimitar-shaped, curved to outer side; apical half setose; apex with elongate sclerotized extension.

(Map 5). South China, Indo-China Peninsula, Malay Peninsula, the Philippines, Borneo, Sumatra, Java. Not discovered in South Asia or islands east of Wallace’s Line.

Jedlička proposed the name Diamella for this genus. Unfortunately, it was preoccupied by a genus of snails in Hydrobiidae. We propose a new name “Diamella” for this genus herein, respecting Jedlička’s notion, with only one letter changed. The gender of this genus name is feminine.

The nearest allied genus to Diamella is unclear. From general appearance and setigerous pores on odd intervals, this genus may be related to Allocota. Detailed discussion is presented in the taxonomic comments below.

Monophyly of Diamella is suggested by the following apomorphic character states: (1) vertex distinctly tumid, posterior supraorbital setae distant from eyes, insertions more or less pointed; (2) males with one pair of setae on terminal sternum; (3) internal sac of aedeagus with main flagellum extraordinarily thick; (4) apical segment of ovipositor with apical extension sclerotized.

The first described species of this genus, Diamella cupreomicans (Oberthür), has been placed in Calleida, Physodera and Allocota by different authors. Obviously, Diamella cupreomicans is much different from members of Calleida or Physodera. In general appearance, Diamella cupreomicans resembles some species of Allocota, but the concept of Allocota has never been clearly defined before the present work.

Among Physoderina, under the present system, Diamella cupreomicans could be more related to Allocota than any other genera both from external and genital characters (Allocota also has the main flagellum slightly thick). But Diamella cupreomicans should be excluded from Allocota based on two important characters: (1) males with one pair of setae on terminal sternum; (2) apical segment of ovipositor long and with apical extension sclerotized.

The mysterious species Diamella kaszabi has never been mentioned after Jedlička (1952, 1963). So far, it is only known from the unique female holotype. We examined the holotype and confirmed that this species is congeneric with Diamella cupreomicans. The most important characters shared by them are tumid vertex and uneven pubescence on elytral intervals. We therefore redefine the concept of genus Diamella herein to include three species.

Holotype of Diamella kaszabi Jedlička, by original designation (MTMB): female, body length = 8.1 mm, board mounted, “Formosa / Sauter”; “Kosempo / 908.”; “TYPUS” [red label]; “Diamella / Kaszabi sp. n. / det. ING. JEDLIČKA” [pink label][Fig. 45].

The original literature clearly indicated that this unique specimen is holotype. The type locality “Kosempo” is old name for “Jiasianpu”, a small hill near township of Jiasian, Kaohsiung County. The locality = 23.07088°N, 120.59386°E, altitude about 380m.

This species can be readily distinguished from the other two species of the genus by: (1) dorsal side brownish, elytra without metallic reflection; (2) elytral base strongly narrowed, hind wings reduced; (3) striae with punctures very coarse. Furthermore, the species also differs from Diamella cupreomicans by: vertex less tumid, front angles of pronotum not narrowed; and from Diamella arrowi by: pronotum narrower, elytra with some setae along striae.

Body length 8.1 mm; dorsal side uniform brown, lateral explanate area of pronotum paler, pronotum and elytra with luster but not metallic; mouthparts and antennae reddish brown, apices of mandibles dark brown; legs yellowish brown; ventral side brownish. Head without microsculpture, with punctures and pubescence; tempora slightly shorter than length of eyes; vertex distinctly tumid, but not forming horn-like hump at posterior supraorbital seta insertion. Pronotum slightly wider than head, widest slightly before middle; ratio PW/PL 1.25; pronotal base strongly lobed; front angles not narrowed, slightly projected forward; lateral margins slightly expanded in middle, slightly rounded but not angulate, strongly sinuate before hind angles; hind angles subrectangular, distinctly projected; disc convex, microsculpture indistinct; lateral explanate areas wide and even; basal foveae slightly deep; disc nearly impunctate, with a few punctures along median line and in basal foveae; disc with a few long setae on basal and lateral areas, lateral explanate area glabrous; lateral margins with long accessory setae equidistant along full length; median line distinct, not reaching to apical and basal margins; disc slightly rugose beside median line. Elytra wider than pronotum, shoulder strongly narrowed, hind wings reduced, nearly as long as elytra; elytral lateral border wide, somewhat reflexed up; striae very deep, coarsely and irregularly punctate, with some setae, especially abundant on 5th to 7th striae; intervals convex, odd intervals with secondary pores, primary pores not distinct among the secondary pores, 3rd and 5th intervals with some larger pores; umbilical series of 9th interval composed of 22 pores; disc without distinct depression; 7th and 8th intervals slightly tumid near apex; elytral lateral margins with dense fine setae; epipleura pubescent. Ventral side. Prosternum with long hairs, proepisterna glabrous; mesosternum nearly glabrous; abdomen hairy. Male genitalia unknown. Female genitalia not studied.

(Map 5). Only known from the type locality, Taiwan.

Lectotype of Calleida cupreomicans Oberthür, designated herein (NNML): male, body length = 6.9 mm, board mounted, “Dr. B. Hagen. / Tandjong Morawa. / Serdang / (N. O. Sumatra)”; “type” [green label]; “Calleida / cupreomicans / R. Obr type”; “Museum Leiden. / Physodera / cupreomicans / Oberth / Det:”; “type” [red label]; “LECTOTYPE ♂ / Calleida cupreomicans / Oberthür, 1883 / des. SHI H. L. 2011” [red label][Fig. 46]. Paralectotype of Calleida cupreomicans Oberthür: 1 female (NNML), “Dr. B. Hagen. / Tandjong Morawa. / Serdang / (N. O. Sumatra)”; “Physodera/ cupreomicans / Oberth. / H. E. Andrewes det.”; “Museum Leiden. / Physodera / cupreomicans / Oberth / Det:”; “Cupreomicans / sp. n. R. Oberth.”; “PARALECTOTYPE ♀ / Calleida cupreomicans / Oberthür, 1883 / det. SHI H. L. 2011” [red label]. Paralectotype of Calleida cupreomicans Oberthür: 1 male (MNHN), “Type” [red label]; “Dr. B. Hagen. / Tandjong Morawa. / Serdang / (N. O. Sumatra)”; “Museum Paris / 1952 / Coll. R. Oberthür”; “Calleida / cupreomicans / R. Oberthür, Type / Notes Leyden Museum / Vol. V. 1883 p. 218”; “PARALECTOTYPE ♂ / Calleida cupreomicans / Oberthür, 1883 / det. SHI H. L. 2011” [red label]. Lectotype of Allocota aerata Bates, designated herein (MSNG): female, body length = 7.5 mm, board mounted, “Bhamò / Birmania / Fea VI 1885”; “TYPUS” [red letter]; “Allocota / aerata / Bates”; “aerata / Bates”; “Allocota / aerata / typus! Bates” [yellow label]; “SYNTYPUS / Allocota / aerata / Bates, 1892” [red label]; “Physodera / cupreomicans / Oberth. / H. E. Andrewes det. 1933”; “Museo Civico / di Genova”; “Museo Civico / di Genova”; “LECTOTYPE ♀ / Allocota aurata / Bates, 1892 / des. SHI H. L. 2011” [red label] [Fig. 47].

Calleida cupreomicans Oberthür. A total three specimens was mentioned in the original literature. We found one of them in MNHN, the general collection of Lebiini, and the other two in NNML. All of them have Oberthür’s determination labels, and agree with original description. We herein designate the male specimen (Fig. 46) in NNML as lectotype for taxonomic purpose of fixing the name to unique name-bearing type.

Allocota aerata Bates. The original literature didn’t mention how many specimens were studied. We found only one female in MSNG with Bates’ determination and type label. It seems that this specimen is the only syntype. We herein designate this female as lectotype (Fig. 47) for taxonomic purpose of fixing the name to unique name-bearing type.

(Total 47 specimens). China: 1 female (HBUM), “Guangxi, Leye, Yachang Forestry Center, 2004.VII.24, Yu Yang & Gao Chao leg.”. 1 male (IZAS), “Yunnan, Fugong county, Lumadeng, Yaping, Yamuhe, 27.13561°N, 98.84166°E, 1612m, 2007.X.7, beating, Shi Hongliang & Liang Hongbin leg.”. 2 males (CAS), “Yunnan, Fugong county, Maji, Majimi, 27.40301°N, 98.82446°E, 1505m, 2005.VIII.26, beating, Liang Hongbin, Zhang Jinfeng leg.”. 1 specimen (IZAS), “Yunnan, Fugong county, Pihe, Wawa, 2005.VIII.24, Liang Hongbin leg.”. 2 females (IZAS), “Yunnan, Fugong county, Pihe, Wawa, 26.58548°N, 98.90467°E, 1120m, 2005.VIII.24, beating, Liang Hongbin, Zhang Jinfeng leg.”[Figs 117, 132]. 1 male (CAS), “Yunnan, Fugong county, Pihe, Bajiao, 26.54816°N, 98.89576°E, 1120m, 2005.VIII.23, beating, Liang Hongbin & Zhang Jinfeng leg.”. 1 female (IZAS), “Yunnan, Longyang, Mangkuan, Baihualing, 25.30367°N, 98.80031°E, 1625m, 2007.X.10, on shrubs, Liang Hongbin leg.”. 2 males (IZAS), “Yunnan, Tengchong county, Hehua, Nangyan, 24.94046°N, 98.38541°E, 1150m, 2006.VI.2, on vegetation, Liang Hongbin leg.”[Fig. 101]. 2 males, 1 female (CAS), “Yunnan, Tengchong county, Mangbang, Longwenqiao, 25.02396°N, 98.67675°E, 1285m, 2006.VI.5, on shrubs, D. Kavanaugh & R. Brett leg.”. 1 female (CAS), “Yunnan, Tengchong county, Qingshui, Rehai, 24.94919°N, 98.44921°E, 1450m, 2006.VI.1, on vegetation, D. Kavanaugh & R. Brett leg.”. 1 specimen (IZAS), “Yunnan, Tengchong county, Mangbang, Longwenqiao, 2006.VI.5, Beating, Liang Hongbin leg.”. 1 male (HBUM), “Yunnan, Yangbi county, Shunbi, 2004.V.21, Yang Xiujuan & Liu Yushuang leg.”. 1 female (IZAS), “Yunnan, Xishuangbanna, Menglun Botany Garden, 2009.XII.1, Tang G. leg.”. 1 female (HBUM), “Yunnan, Yingjiang county, 2004.V.17, Yang Xiujuan & Liu Yushuang leg.”. 2 males (HBUM), “Yunnan, Gengma county, Mengding, 2004.VIII.6-7, Li Jing, Yuan Caixia leg.”. 1 male (IZAS), “Yunnan, Xishuangbanna, Mengla, 620-650m, 1959.V.29, Zhang Yiran leg”. 1 female (IZAS), “Yunnan, Xishuangbanna, Damenglong, 650m, 1958.V.3, Hong Chunpei leg.”. 1 female (IZAS), “Hainan, Changjiang county, Bawangling, Est. 2 station, 19.09901°N, 109.17540°E, 1015m, 2007.V.9, light trap, Liang Hongbin leg.”. 2 males (IZAS), “Hainan, Baisha county, Nankai, 19.07926°N, 109.41133°E, 262m, 2009.XI.22, Liang Hongbin leg.”. 1 male (IZAS), “Hainan, Yinggeling, Hongkan Reservoir, 19.08121°N, 109.49839°E, 525m, 2009.XI.24, Liang Hongbin leg.”[Figs 13, 77]. Vietnam: 5 males 1 female (IZAS), “Tonkin, Hoa-Binh, 1939.VII, leg. A. de Cooman”. 1 female (IZAS)”Tonkin, Hoa-Binh, leg. A. de Cooman”. 1 female (MNHU), “Annam, Phuc-Son, Nov. Dez., H. Fruhstorfer”. 1 specimen (MNHN), “Tan Hoa, Indochine, VI-43”. Laos: 1 specimen (CMB), “Laos, 34 km nnw., Lanau Nam Pho, 5.5.2005, leg. Yokoi”. 1 specimen (NHML), “Laos, nam mia, 31-IV.1918, R. Vitalis de Salvaza”. Malaya: 1 specimen (NHML), “Malaya, Kuala Lumpur., at night, Jan. 9.1924”. Borneo: 1 specimen (MNHN), “Nord Borneo, Mont Kina Balu, 5-8-1903, John Waterstradt”. 1 female (NHML), “Sarawak:, foot of Mt. Dulit, Junction of rivers, Tinjar & Lejok., 25.ix, 1932”. 1 female (NHML), “Pagat, Borneo, 8/82, Grabowth”. Sumatra: 1 specimen (NHML), “Sumatra, Manna 1901, M. Knappert.”. 3 specimens (MNHN), “Paggar Alam, Sumatra, J. Bouchard”. Java: 1 specimen (NHML), “Java merid., Palabuan 1892, H. Fruhstorfer”.

Vertex strongly tumid, forming horn-like humps at posterior supraorbital seta insertions; pronotal front angles strongly narrowed; elytral base wide, hind wings developed; elytra copperish to dark green, strongly metallic; elytra with some setae along striae. Comparison with the other two species of this genus is presented in the key above and diagnosis under those species.

Body length 6.9–8.0 mm; head and pronotum dark brown, lateral explanate areas of pronotum paler; elytra copperish to dark green, strongly metallic; mouthparts and antennae brown, apices of terminal labial palpomeres yellow; legs dark brown, with faint metallic reflection, tarsomeres brown; ventral side brownish with luster. Head without punctures or microsculpture, vertex with a few setae; tempora slightly shorter than length of eyes; vertex strongly tumid, forming horn-like humps at posterior supraorbital seta insertions, posterior supraorbital setae distant from eyes; labrum slightly widened to apex, apical margin straight or slightly curved. Pronotum slightly wider than head, widest at apical one-third; ratio PW/PL 1.27–1.35; pronotal base strongly lobed; front angles distinctly narrowed, not projected forward; lateral margins slightly expanded in middle, slightly rounded but not angulate, strongly sinuate before hind angles; hind angles acute or subrectangular, distinctly projected; disc convex, microsculpture indistinct; lateral explanate areas wide and even; basal foveae slightly deepened; disc nearly impunctate, with a few punctures only along median line and in basal foveae; disc and basal area with a few long setae, lateral explanate areas glabrous; lateral margins with long accessory setae equidistant along full length, sometimes sparser on middle area; median line distinct, not reaching apical or basal margins; disc slightly rugose beside median line. Elytra wider than pronotum, shoulder not narrowed, hind wings well developed; elytral lateral borders narrow, not reflexed up; striae distinct, coarsely and irregularly punctate; intervals slightly convex, odd intervals with secondary pores, primary setigerous pores indistinct among the secondary pores, only two pores distinguishable: one pore on 5th interval base adjacent to 5th stria and one apically on 3rd interval adjacent to 2nd stria; striae with some setae, more abundant in 5th to 7th striae; umbilical series of 9th interval composed of 17–18 pores; 3rd to 6th intervals slightly depressed subapically; 7th and 8th intervals distinctly tumid near apex; elytral lateral margins with dense fine setae; epipleura pubescent. Ventral side. Prosternum with long pubescence, proepisterna glabrous; mesosternum nearly glabrous; abdomen hairy. Male genitalia with median lobe of aedeagus straight and flat, gently expanded to apex; dorsal side slightly pubescent subapically; left lateral margin slightly curved in middle; apical orifice opened apically; apical lamella placed on right side, triangular, width subequal to length in dorsal view; internal sac with main flagellum thick and slightly sinuous, distinctly grooved ventrally, nearly reaching apical orifice; trumpet-form expansion strongly expanded, nearly half length of main flagellum (Fig. 77). Female genitalia. Spermatheca straight, short; apical fourth slightly widened, forming a weak globular expansion, with distinct ring-sculpture; subbasal part of spermatheca with faint ring-sculpture; spermathecal gland inserted near apical fourth of spermatheca, not branched, base slightly thickened and curved, slender and long, much longer than spermatheca (Fig. 132). Apical segment of ovipositor scimitar-shaped, slightly curved outwards after middle, gradually narrowed to apex; length four times basal width approximately; inner and outer margins finely setose in apical half; apex very sharp and fine, with very narrow and long sclerotized extension (Fig. 117).

(Map 5). South China (Yunnan, Guangxi, Hainan); Myanmar, Laos, Vietnam, Malaysia, Indonesia (Sumatra, Borneo, Java).

Two females from Borneo differ from typical specimens by dark blue elytra, but other characters are identical with this species.

We dissected a paralectotype of Calleida cupreomicans Oberthür and a male from Yunnan, very close to the type locality of Allocota aerata Bates, and found that there were slight differences: specimen from Yunnan with median lobe of aedeagus a little more expanded to right side, and apical lamella slightly slenderer than that in the paralectotype of Calleida cupreomicans.

After examining types of both species, Andrewes (1936) indicated that Allocota aerata Bates is a synonym of Calleida cupreomicans Oberthür. We also examined both types and confirmed the synonymy. Types of these two species come from far distant localities, but they are identical. The slight difference of genitalia between specimens is regarded as geographical variation.

Holotype of Lachnoderma arrowi Jedlička, by original designation (NHML): male, body length = 7.3 mm, board mounted, “Philippine Is. / Coll. Bottcher. / B. M. 1929-201”; “96”; “Type” [round label with red ring]; “Lachnoderma / Arrowi / Jedl. sp. n. / det. ING. JEDLIČKA” [pink label] [Figs 48, 78, 102].

The original literature clearly indicated that this specimen (Fig. 48) is the holotype. But precise locality was not provided.

Vertex moderately tumid, not forming horn-like humps at posterior supraorbital seta insertions; pronotum front angles not narrowed, lateral margins completely rounded in middle; elytra dark blue, strongly metallic; elytra with striae very shallow, with accessory setae only on odd intervals. Male genitalia of this species resemble Diamella cupreomicans, but: (1) apical lamella very short and wide, apex rounded; (2) in lateral view, median lobe of aedeagus slender, ventral margin less expanded near base.

Body length 7.3 mm; head and pronotum reddish brown, with luster but not metallic, lateral explanate areas of pronotum paler; elytra metallic blue with disc somewhat reddish brown; mouthparts and antennae reddish brown, apices of terminal labial palpomeres yellow; legs brown, tarsomeres slightly paler; ventral side brownish. Head without microsculpture or punctures, sparsely pubescent on vertex; tempora slightly longer than half length of eyes; vertex moderately tumid, posterior supraorbital setae distant from eyes, seta insertions slightly humped, but not horn-like; labrum slightly widened to apex, apical margin deeply emarginate. Pronotum slightly wider than head, widest slightly before middle; ratio PW/PL 1.45; pronotal base briefly but distinctly lobed; front angles wide; lateral margins strongly expanded in middle, completely rounded, slightly sinuate before hind angles; hind angles subrectangular, hardly projected; disc slightly convex, microsculpture indistinct; lateral explanate areas wide and even; basal foveae shallow; disc nearly impunctate, with a few punctures along median line and in basal foveae; disc with long setae except area along median line, some setae distributed to middle of lateral explanate areas; lateral margins with long accessory setae through their full lengths, sparser after middle; median line distinct, nearly reaching basal and apical margins; disc not rugose. Elytra wider than pronotum, shoulders not narrowed, hind wings well developed; elytral lateral borders narrow, not reflexed up; striae very shallow, finely punctate; scutellary intervals with three large pores (basal pores); intervals flat, without microsculpture, odd intervals with secondary pores, more abundant on 7th interval; primary pores indistinct among secondary pores, 3rd and 5th intervals with some larger pores distinguishable; setae not present along striae; umbilical series of 9th interval composed of 15 pores; disc without distinct depression; lateral margins slightly depressed at basal third; 7th and 8th intervals slightly tumid near apex; elytral lateral margins with dense and fine setae; epipleura pubescent. Ventral side. Prosternum with long pubescence, proepisterna glabrous; mesosternum nearly glabrous; abdomen hairy. Male genitalia with median lobe of aedeagus straight and flat, gently expanded to apex; dorsal side slightly pubescent subapically; left lateral margin slightly curved in middle; apical orifice opened apically; apical lamella placed on right side, short and rounded, distinctly wider than length in dorsal view; internal sac with main flagellum thick and slightly corkscrewed, distinctly grooved ventrally, nearly reaching apical orifice; trumpet-form expansion strongly expanded, nearly half length of main flagellum (Fig. 78). Female genitalia unknown.

(Map 5). Only known from the type locality, the Philippines.

Allocota viridipennis Motschulsky, 1859, by monotypy.

Dorsal side glabrous; mandibles strongly widened; posterior supraorbital setae near eyes, insertions not tumid; pronotum narrow, ratio PW/PL less than 1.4; lateral setae of pronotum absent; elytral 3rd and 5th intervals with four or more setigerous pores, subequally placed; 7th intervals sometimes with setigerous pores; protibiae with cleaning spur reduced or absent; males with two pairs of setigerous pores on terminal sternum.

This genus is closest to Physodera; their differences are presented in the diagnosis of Physodera. In general appearance, Allocota is similar to Diamella, but can be distinguished from the latter by the glabrous surface and vertex not tumid.

Body length 6.0 to 8.7 mm, slender, pronotum narrow. Elytra strongly metallic, sometimes disc more or less reddish. Head and pronotum glabrous, elytra glabrous except primary setigerous pores; elytra usually with faint isodiametric microsculpture. Head glabrous; eyes hemispherical, strongly prominent; tempora slightly longer than half length of eyes, gradually narrowed behind eyes, not expanded; vertex not tumid. Antennae extended to about elytral basal one-third; 1st antennomere gradually narrowed to base, 3rd longer than 4th. Labrum smooth, without secondary setae, slightly widened to apex, apical margin more or less emarginate; mandibles strongly widened, outer margin rounded (Fig. 150), surface glabrous; terminal maxillary palpomeres fusiform in both sexes; terminal labial palpomeres more or less widened in males, fusiform, truncate or slightly securiform; less widened in females; ligular apex truncate, with four long setae; paraglossae membranous, as long as ligula, adnate; mentum tooth simple, apex rounded or sharp, with two setae near base; submentum with two long setae; genae glabrous beneath eyes. Pronotum slightly wider than long, nearly as wide as head; disc glabrous, front angles with a few setae, lateral margins glabrous; mid-lateral setae absent; pronotal base briefly but distinctly lobed; lateral margins slightly expanded in middle; hind angles more or less distinct; basal foveae shallow. Elytra narrow, slightly convex, slightly widened to apex; apex truncate or slightly curved, sutural angles not projected, outer angles rounded; disc without distinct depression; intervals flat, without additional pubescence; basal margination only reaching 3rd interval; basal pores present; 3rd and 5th intervals with four or more setigerous pores, sometimes 7th interval also with setigerous pores; 7th and 8th intervals strongly tumid near apex. Ventral side nearly glabrous; males with terminal sternum moderately emarginate apically, with two pairs of setae (Fig. 145); females with terminal sternum straight or slightly curved apically, with two pairs of setae, rarely one additional setae present on one side (the right side in Fig. 146). Legs short; protibiae with cleaning spur absent (Fig. 140), or very fine and short (Fig. 141), distant from inner margin; tarsi widened; 4th tarsomere bifid, claws pectinate; males with adhesive hairs on first three protarsomeres well developed (two whole rows), those on first three mesotarsomeres rudimentary (two rows, weakly present near apex) or well developed (two whole rows) on some mesotarsomeres. Male genitalia with median lobe of aedeagus not twisted, slightly bent to right side in dorsal view; apical orifice opened apically; apical lamella short; dorsal surface with some fine setae subapically; internal sac with main flagellum moderately thick, apex not reaching apical orifice, trumpet-form expansion small; apical bursa absent; secondary flagellum distinct. Female genitalia. Spermatheca tubular, with more or less ring-sculpture, inserted at the joining of the bursa copulatrix and common oviduct; spermathecal gland with basal part very fine, inserted at middle of spermatheca; spermatheca not distinctly bent. Apical segment of ovipositor very short and wide, apex rounded, with membranous extension slightly widened.

(Map 6). East Asia and Southeast Asia: Japan, China, Indo-China Peninsula, Malay Peninsula, Philippine Islands, Borneo, Sumatra, Java, Sulawesi. Not discovered in South Asia or the eastern part of the Malay Archipelago.

Monophyly of Allocota is suggested by the following apomorphic character states: (1) cleaning spur on protibiae reduced or absent; (2) setigerous pores on 5th interval not restricted to basal half; (3) apical segment of ovipositor only slightly longer than basal width.

We studied the type series and other material of Taicona aurata Bates (type-species of Taicona), and a male of Allocota viridipennis Motschulsky (type-species of Allocota) from Java (type locality) that had been compared with the type by Andrewes. Except for color and body size, the only significant difference between them is that setigerous pores are present on the 7th interval in Allocota. Even the male genitalia show no obvious differences at the species level (see the detailed discussion under Allocota aurata). So it can be inferred that these two species are extremely closely related. As the pore distribution on the 7th interval is not constant even at the species-level (see discussion under Allocota viridipennis), we herein synonymize Taicona Bates with Allocota Motschulsky.

A total of seven species was included in Allocota before the present study, but four of them should be transferred to other genera. In the present paper, we combine species previously included in Taicona with Allocota, propose two new synonyms, and describe a new species. Therefore, in the present concept of Allocota, a total of four species is included. Three of these are very closely allied with each other, and strictly allopatric; while the fourth one, Allocota bicolor sp. n., is quite different from the others and sympatric with Allocota aurata.

Key to species of Allocota Motschulsky

Lectotype of Allocota caerulea Andrewes, designated herein (MNHN): male, body length = 6.8 mm, pin mounted, “Ex Musaeo, Chaudoir” [red printed]; “Museum Paris, 1952, Coll. R. Oberthür”; “viridipennis / Motsch. / Singapore / Wallace” [large box label but pinned under specimen]; “LECTOTYPE ♂/ Allocota caerulea / Andrewes, 1933 / des. SHI H. L. 2011” [red label][Fig. 49]. Paralectotype of Allocota caerulea Andrewes: a male (MNHN), “♂”; “Ex Musaeo, Chaudoir” [red printed]; “Museum Paris, 1952, Coll. R. Oberthür”; “PARALECTOTYPE ♂/ Allocota caerulea / Andrewes, 1933 / des. SHI H. L. 2011” [red label].

Allocota viridipennis Motschulsky. The type series of Allocota viridipennis Motschulsky which was mentioned by Andrewes (1933) should be in Moscow University together with Motschulsky’s main collection. Unfortunately, we have not had opportunity to examine this collection. But in the collection of NHML we found a male of Allocota viridipennis from the type locality (Java) that had been compared with the type by Andrewes (it was mentioned by Andrewes 1933, as a female). This specimen is enough for us to recognize this species.

Allocota caerulea Andrewes. Andrewes (1933) indicated that the name Allocota caerulea Andrewes was for the species mentioned by Chaudoir (1877) as Allocota viridipennis, and type material was in the collection of Oberthür. So, the two specimens from Chaudoir’s ex-collection in MNHN determined by Chaudoir as Allocota viridipennis should be the type series of Allocota caerulea Andrewes. We designate the male with Chaudoir’s box label pinned under as lectotype (Fig. 49) herein for Allocota caerulea Andrewes, for the taxonomic purpose of fixing the name to a single specimen and preventing further uncertainty.

(Total 33 specimens). Singapore: 1 specimen (MNHN), “Singapore, A. Raffray.”. 1 specimen (MNHN), “Singapore”; “Ex Musaeo, H. W. Bates, 1892”. 2 specimens (MNHN), “Singapore”; “ Ex Musaeo, L. Fairmaire, 1896”[Figs 15, 80]. Sumatra: 4 specimens (MNHN), “Sumatra, Palembang”. 1 male (NHMB), “W Sumatra prov.; Kerinci Seblat N.P.; 24km NE Tapan; Muara Sako – E env.; 2°05'S, 101°15'E; 400–550m; Dembický leg.; 4–18.iii.2003” Borneo: 1 male (MNHN), “Sarawak, Hewilt”. 1 specimen (NHML), “Quop, W. Sarawak, G. E. Bryant., 1.III.14”. 2 specimens (NHML) “N. Borneo, Bettotan, Nr. Sandakan., July 29.1927”. 1 male (NHML) “Sandakan, Borneo, Baker”; “Ex Mus. Coll. Agric. Phil. Is.”; “Allocota viridipennis Motch., H. E. Andrewes det.”. 3 specimens (MNHN), “Est Borneo, Batanbessi, Mem. W. Walsh, 1937”. 1 specimen (MNHN), “Est Borneo, Batan bessi, Me. M. E. Walsh, 1937”. 7 specimens (MNHN), “Est Borneo, Kariovang.”. 1 specimen (NHML), “Borneo”. Java: 1 specimen (MNHN), “Java (Meuwen Bay), Détr. De la Sonde, Raffray & Maindron, 1878”. 5 specimens (MNHN), “Museum Paris, Java, Deyrolle 1877”. 1 male (NHML) “23.III.1924, Depok, Karny”; “Ex. Mus. Buitenzorg”; “Allocota viridipennis Motch., Compared with type H. E. A.”[Figs 14, 79].

Elytra metallic blue, more or less greenish in some specimens; metasternum and abdomen much darker than prosternum; front angles of pronotum with setae very short and fine (Fig. 151); elytral 7th interval with setigerous pores in some specimens; internal sac of aedeagus with setose area divided into two parts, apex of secondary flagellum simple.

In general appearance, this species resembles Allocota cyanipennis Heller, but can be distinguished in having: (1) internal sac of aedeagus with setose area divided into two parts, apex of secondary flagellum simple (Figs 79, 80); but in Allocota cyanipennis, setose area divided into three parts, apex of secondary flagellum strongly expanded forming a large triangular sclerite (Fig. 81). (2) front angles of pronotum with setae very short and fine; but in Allocota cyanipennis, those setae much longer. (3) Setae on elytral lateral margins more distinct in Allocota cyanipennis.

Body length 6.0–6.9 mm; head and pronotum orange red to reddish brown; mouthparts earth yellow, palpomeres brownish, apex of terminal palpomeres yellow; 1st antennomere reddish yellow, the remaining antennomeres usually slightly darker; elytra metallic blue, sometimes more or less greenish, or green on lateral areas, elytral suture and lateral margins metallic; legs usually much darker than pronotum, darkest on apical half of femora; ventral side of head and prosternum the same color as dorsal side; metasternum and abdomen brownish to piceous, much darker than prosternum. Head glabrous, without punctures, microsculpture indistinct; males with terminal labial palpomeres fusiform, slightly expanded in middle. Pronotum glabrous, cordiform, widest at apical one-third; ratio PW/PL 1.22 to 1.32; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with a few fine and short setae (distinctly shorter than in Allocota cyanipennis); lateral margins rounded, slightly expanded in middle, strongly sinuate before hind angles; lateral explanate areas moderately wide, without punctures; hind angles more or less distinct, rectangular or nearly so, usually slightly pointed; basal foveae very shallow, without punctures; median line very fine, usually indistinct. Elytra with striae slightly sulcate, finely punctate; intervals nearly flat, without accessory setae, with a row of very fine punctures; microsculpture very shallow, isodiametric, or absent; 3rd and 5th intervals with four to ten setigerous pores, their position variable; 7th interval with some setigerous pores or without pore; setae on lateral margins very fine and short, hardly visible. Legs. Protibiae with cleaning spur absent (Fig. 140); males with adhesive hairs well developed (two whole rows) on first two mesotarsomeres, rudimentary (two rows, weakly present near apex) on 3rd mesotarsomere. Male genitalia with median lobe of aedeagus stout, slightly bent to right side near apex in dorsal view, right margin slightly curved before apical lamella; apical lamella placed on right side, narrow, slightly elongate; internal sac with setose area divided into two parts; apex of secondary flagellum simple, not expanded; trumpet-form expansion with ventral margin more or less expanded (Figs 79, 80). Female genitalia. Apical segment of ovipositor very short, slightly longer than width; outer margin straight; inner margin curved, with fine setae on apical half; membranous extension short and wide. Internal reproductive system not studied.

(Map 6). Singapore, Borneo, Sumatra, Java.

We examined some material from Singapore, Borneo and Sumatra identical with the lectotype of Allocota caerulea, and a male from Java determined as Allocota viridipennis and compared with type by Andrewes. Except color variation on elytra, the only other significant difference is: “Allocota viridipennis” from Java with five or six setigerous pores on elytral 7th interval (Fig. 14), but the lectotype of “Allocota caerulea” without any pores on 7th interval (Fig. 15). Setigerous pores on 7th interval are not only variable between specimens, e. g., individuals with or without pores can be found from same locality (for example in Java or Borneo), but also in same individual, e. g., one or two pores may present on 7th interval on one elytron, and no such pores on the other elytron. In addition, the aedeagi of these specimens show no significant differences (Figs 79, 80). We therefore synonymize Allocota caerulea Andrewes with Allocota viridipennis Motschulsky herein.

As mentioned above, this species is variable in color and setigerous pores on elytra. Usually, the elytra are nearly metallic blue (Figs 15, 49), and without setigerous pores on 7th interval. But in some specimens from Java elytra are distinctly greenish (Fig. 14); in some specimens from Java and Borneo, 7th interval with several setigerous pores.

Holotype of Allocota cyanipennis Heller, monotypy (SNSD): male, body length = 6.3 mm, board mounted, “Tangcolan / Bukidnon /Baker”; “cyanipennis / Typus” [red label]; “14261”; “1921 / 3” [yellow label]; “Staatl. Museum für / Tierkunde, Dresden”[Fig. 50].

Heller (1923) didn’t clearly state this species was based on a single specimen, but he only cited one specimen with a serial number “14261”. So, the holotype (Fig. 50) was originally fixed by monotypy, as the description implies a single specimen.

(Total 5 specimens). The Philippines: 1 specimen (NHML), “Philippine Is., Coll. Bottcher., B. M. 1929–201”. 1 male (NHML), “Island of Basilan, Baker”; “Ex Mus. Coll. Agric. Phil. Is.”; “Allocota cyanipennis Heller, Compared with type H. E. A.”[Figs 16, 81]. 1 specimen (NHML), “Tangcolan, Bukienon, Baker”. Sulawesi: 1 male and 1 female (NHML), “Celebes”.

Elytra metallic blue; metasternum and abdomen much darker than prosternum; front angles of pronotum with setae relative long (Fig. 152); elytral 7th interval without setigerous pores; internal sac of aedeagus with setose area divided into three parts, apex of secondary flagellum forming a large triangular sclerite.

This species is closest to Allocota viridipennis Motschulsky; differences are presented in the diagnosis part under the latter species.

Body length 6.3–7.1 mm; head and pronotum orange red or brown; mouthparts earth yellow, palpomeres brownish, apex of terminal palpomeres yellow; 1st antennomere reddish, the remaining antennomeres usually slightly darker; elytra metallic blue, elytral suture and lateral margins metallic; legs usually much darker than pronotum, darkest on apical half of femora, basal half usually the same color as pronotum; ventral side of head and prosternum the same color as dorsal side; metasternum and abdomen brownish to piceous, much darker than prosternum. Head glabrous, without punctures, microsculpture indistinct; males with terminal labial palpomeres fusiform, slightly expanded in middle. Pronotum glabrous, cordiform, widest at anterior third; ratio PW/PL 1.28 to 1.32; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with setae relative long (Fig. 152); lateral margins rounded, slightly expanded in middle, strongly sinuate before hind angles; lateral explanate areas moderately wide, without punctures; hind angles slightly acute, less distinct; basal foveae very shallow, without punctures; median line very fine, usually indistinct. Elytra with striae very shallow, finely punctate; intervals nearly flat, without accessory setae, with a row of very fine punctures; microsculpture absent; 3rd and 5th intervals with four to ten setigerous pores, their positions variable; 7th interval without setigerous pores; setae on lateral margins fine and short, but more distinct than in Allocota viridipennis. Legs. Protibiae with cleaning spur absent (Fig. 140); males with adhesive hairs well developed (two whole rows) on first two mesotarsomeres, rudimentary (two rows, weakly present near apex) on 3rd mesotarsomere. Male genitalia with median lobe of aedeagus stout, more distinctly bent to right side near apex than in Allocota viridipennis in dorsal view, right margin slightly curved before apical lamella; apical lamella placed on right side, narrow, slightly elongate; internal sac with setose area divided into three parts; apex of secondary flagellum expanded, formed a large triangular sclerite; trumpet-form expansion with ventral margin more or less expanded. Female genitalia not studied (Fig. 81).

(Map 6). The Philippines, Sulawesi.

Pronotum and head of two specimens from Sulawesi we examined are brown, much darker than those from the Philippines which are vivid orange.

We regard specimens from the Philippines and Sulawesi as a distinct species from Allocota viridipennis Motschulsky for the constant difference in internal sac of aedeagus as mentioned above in the diagnosis section under Allocota viridipennis. In external characters these two species are very similar, with only slight differences in setae on pronotal front angles (Figs 151, 152) and elytral lateral margins.

Lectotype of Taicona aurata Bates, designated herein (MNHN): male, body length = 7.2 mm, pin mounted, “NAGASAKI”; “TYPE”[red label]; “Taicona / aurata / Bates”; “Ex Musaeo / H. W. Bates / 1892”; “Museum Paris / 1952 / Coll. R. Oberthür”; “LECTOTYPE ♂ / Taicona aurata / Bates, 1873 / Des. SHI H. L., 2011” [red label][Fig. 51]. Paralectotypes of Taicona aurata Bates: 1 female (MNHN), “NAGASAKI”; “PARATYPE”[red label]; “Ex Musaeo / H. W. Bates / 1892”; “Museum Paris / 1952 / Coll. R. Oberthür”; “Taicona / Bates”; “PARALECTOTYPE ♀ / Taicona aurata / Bates, 1873 / Des. SHI H. L., 2011” [red label]; Labial removed and pinned isolated with 2 labels: “Taicona / aurata”; “Ex Musaeo / H. W. Bates / 1892”. 1 female (NHML), “Type” [round label with red circle]; “Japan. / G. Lewis / 1910-320”; “Taicona / aurata / Bates”; “PARALECTOTYPE / Taicona aurata / Bates, 1873 / det. SHI H. L. 2011” [red label]. 1 male (NHML), “NAG”; “Japan. / G. Lewis. / 1910-320”; “Ex coll. Brit. Mus.”; “Co-type” [round label with green circle]; “Taicona / aurata/ Bates”; “H. E. Andrewes Coll. / B. M. 1945-97”; “PARALECTOTYPE / Taicona aurata / Bates, 1873 / det. SHI H. L. 2011” [red label].

Taicona aurata Bates. Bates (1873) didn’t state the number of specimens in the type series. We found a total of four specimens bearing Bates’ determination labels in the collection of MNHN and NHML. Two specimens from different collections were both determined and labeled as “type”. We herein designate the male in the collection of MNHN as lectotype (Fig. 51) for taxonomic purpose of fixing the name to unique name-bearing type.

Taicona perroti Jedlička. As indicated in the original description, the holotype of Taicona perroti Jedlička should be in MNHN. But we didn’t find it in MNHN or NMPC where the collection of Jedlička is deposited. We suspect that the holotype is probably still in MNHN, but has been misplaced.

(Total 36 specimens). Japan: 2 males, 1 female (OMNH), “KASUGA NARA, 1959.V.31, K. Ueda leg.”[Fig. 82]. 1 female (OMNH), “KASUGA NARA, 1959.V.31, T. Tomiwa leg.”[Fig. 18]. 1 female (NHML), “Japan. G. Lewis. 1910-320.”; “Shiba San Chio 1883”. Shaanxi: 1 male, 4 females (IZAS), “Shaanxi, Foping county, Shangshawo, 33.59716°N, 108.01366°E, 1107m, 2007.VIII.15, beating, SHI Hongliang, YANG Ganyan leg.”[Figs 104, 118, 133]. Guangdong: 3 males, 1 female (IZAS), “Guangdong, Shixing county, Chebaling, Xianrendong village, 24.73478°N, 114.20727°E, 508m, 2008.VII.23, beating, LIANG Hongbin leg.”. Hainan: 1 female (IZAS), “Hainan, Baisha county, Yinggeling Mt., Hongkan reservoir, 19.08121°N, 109.49839°E, 525m, 2009.XI.24, beating, LIANG Hongbin leg.”. 1 male, 2 females (IZAS), “Hainan, Baisha county, Nankai, 19.07926°N, 109.41133°E, 262m, 2009.XI.22, beating, LIANG Hongbin leg.”[Fig. 19]. 1 specimen (IZAS), “Hainan, Baisha county, Nankai, beating, 2009.11.21, LIANG Hongbin leg.”. 1 male, 1 female (IZAS), “Hainan, Baisha county, Nankai, Daoyin village; on vegetation; 19.01021°N, 109.36910°E, 336m, 2010.4.15 D, LIN Meiying leg.”. Yunnan: 1 male (IZAS), “China, Yunnan Prov., Fugong, Lumadeng, Yaping vill. Plant beating, 27.13076°N, 98.87447°E; 1295m, 2005.8.25 day, Liang H. B., Zhang J. F. leg.”. 1 male (CAS), “China, Yunnan Prov., Fugong, Pihe Town, Wawa, Plant beating, 26.58548°N, 98.90467°E; 1120m, 2005.8.24 day, Liang H. B., Zhang J. F. leg.”. 2 males, 2 females (IZAS), “China, Yunnan Prov., Tengchong, Qingshui, Rehai, on vegetation; 24.94861°N, 98.45181°E; 1470m, 2006.6.1 day, Liang H. B., Hu P. leg.”[Fig. 20]. 2 males, 1 female (CAS), “China, Yunnan Prov., Tengchong, Mangbang, Longwenqiao, on shrubs; 25.02329°N, 98.67710°E; 1290m, 2006.6.5 day, Liang H. B., Hu P. leg.”. 1 female (IZAS), “China, Yunnan, Ruili, Dengga to Mafengshan. N23.95285, E97.59808 – N23.94485, E97.55647; 927–1207m; 2009.VIII.10, Shi H. L. leg. beating”. 1 female (IZAS), “Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.VI.12, WANG Shuyong leg.”. 1 male, 1 female (IZAS), “Yunnan, Jinghong, Nabanhe Reserve, Mandian, N.22.13061, E1000.67377, 718m, 2010.IX.30, LIN Meiying leg.”[Fig. 21]. 1 male, 2 females (IZAS), “Yunnan, Xishuangbanna, Menglun Botany Garden, Lyushilin; 2009.XI.17, 643m, Tang G., Yao Z. Y. leg.”. Vietnam: 1 male, 1 female (IZAS), “TONKIN, Hoa-Binh, 1939.VII, leg. A. de Cooman”. 1 male (IZAS), “TONKIN, Hoa-Binh, 1940.VII, leg. A. de Cooman”[Figs 83, 103]. 1 male (IZAS), “TONKIN, Hoa-Binh, leg. A. de Cooman”. 1 specimen (MNHN), “tonkin, Cap. Fouquet”. 1 specimen (MNHN), “Tonkin, Region de, Hoa-Binh”. 1 specimen (MNHN), “Tonkin, Reg. De Hoa-Binh, A. De Cooman, 1929”. Laos: 1 male (ZSM), “Laos, Umg. Vientiane, III.-VI.1963”. Nepal: 1 female (CRS), “W. Nepal, chitwan Distrikt/ Chitwan Nat. Park, 230m, / Leg. Probst 28.–30.5.1993”. 1 male (NHMB), “Jiri-Shivalaya (Khimti Khola) 2500–1800m, 11–12.VI.1987”; “C-Nepal, Janakpur, C. J. Rai”

Elytra metallic green or cupreous–green, disc with more or less distinct reddish patch in some specimens; metasternum and abdomen not darker than prosternum; front angles of pronotum with setae very fine and short (as in Fig. 151); elytral 7th interval without setigerous pores; internal sac of aedeagus with setose area divided into two parts, apex of secondary flagellum simple.

This species is very close to Allocota viridipennis Motschulsky. We didn’t find significant difference between their male genitalia. These two species can be distinguished by different color on elytra and ventral side, and also by different distribution (Map 6).

Body length 6.4–8.2 mm; head and pronotum reddish yellow to dark brown; mouthparts yellowish brown, apices of terminal palpomeres paler; 1st antennomere usually slightly paler than the remaining antennomeres; elytra metallic green to cupreous green, sometimes disc with a more or less distinct reddish patch, elytral suture and lateral margins yellowish or metallic; legs reddish yellow to dark brown, usually the same color as pronotum, sometimes slightly darker on apex of femora; ventral side the same color as pronotum; metasternum and abdomen not darker than prosternum. Head glabrous, without punctures, microsculpture indistinct; males with terminal labial palpomeres fusiform, slightly expanded in middle. Pronotum glabrous, cordiform, widest at apical one-third; ratio PW/PL 1.20 to 1.35; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, slightly transverse, without punctures; front angles with a few very fine short setae (distinctly shorter than in Allocota cyanipennis); lateral margins rounded, slightly expanded in middle, strongly sinuate before hind angles; lateral explanate areas moderately wide, without punctures; hind angles usually acute and sharp; basal foveae very shallow, without punctures; median line very fine, usually indistinct. Elytra with striae slightly distinct, finely punctate; intervals slightly convex, without accessory setae, with a row of very fine punctures; usually with very faint isodiametric microsculpture; 3rd and 5th intervals with four to ten setigerous pores, their positions variable; 7th interval without setigerous pores; setae on lateral margins very fine and short, hardly visible. Legs. Protibiae with cleaning spur absent (Fig. 140); males with adhesive hairs well developed (two whole rows) on first two mesotarsomeres, rudimentary (two rows, weakly present near apex) on 3rd mesotarsomere. Male genitalia with median lobe of aedeagus stout, slightly bent to right side near apex in dorsal view, right margin slightly curved before apical lamella; apical lamella placed on right side, narrow, slightly elongate; internal sac with setose area divided into two parts; apex of secondary flagellum simple, not expanded; trumpet-form expansion with ventral margin more or less expanded (Figs 82, 83). Female genitalia. Spermatheca straight, tubiform, moderately long; apical part not distinct widened, with very faint ring-sculpture; spermathecal gland inserted near middle of spermatheca, not branched, slightly thickened at base, not longer than spermatheca (Fig. 133). Apical segment of ovipositor very short, slightly longer than width; outer margin straight; inner margin curved, with fine setae on apical half; membranous extension short and wide (Fig. 118).

(Map 6). Japan, China (Shaanxi, Guangdong, Hainan, Yunnan), Vietnam (Tonkin), Laos, Nepal.

Jedlička (1963) described Taicona perroti based mainly on the distinct reddish patch on elytral disc and bluish metallic color being different from Allocota aurata. These characters are always present in specimens from Tonkin or Hainan. But we didn’t find any difference between this “form” and the typical “Allocota aurata” from Japan in male genitalia or external characters except color. The other characters mentioned by Jedlička (1963), such as differences in pronotal shape and punctures on striae, vary among individuals, so we herein synonymize Taicona perroti Jedlička with Taicona aurata Bates.

A total of four different color forms was found in this species from different localities: (1) typical “Allocota aurata” comes from Japan, and was also found in Shaanxi and Guangdong of China; the head and pronotum are reddish brown; elytra metallic green, disc slightly reddish, not forming distinct patch (Figs 18, 51); (2) specimens from Tonkin, Hainan, Laos and Nepal, namely “Allocota perroti”, with head and pronotum vivider compared to specimens from Japan, elytra metallic green, slightly bluish, disc with large reddish patch reaching 5th or 6th interval, the patch much more distinct than in the “typical Allocota aurata” (Fig. 19); (3) specimens from the southern part of Yunnan (Xishuangbanna) with head, pronotum and legs dark brown; elytra metallic green, disc not reddish (Fig. 21); (4) specimens from the western part of Yunnan (Fugong, Tengchong, Ruili) with head and pronotum slightly darker than in the “typical form”, elytra distinctly cupreous, disc not reddish (Fig. 20). These forms were only different in color, but other characters including male genitalia are the same. So we consider them as geographical variation rather than distinct species or subspecies.

This species has male genitalia very similar with Allocota viridipennis Motschulsky implying they could be synonyms. However, their constant differences on elytral and ventral color, as well as the distribution gap make us quite doubtful to synonymize them at present. Therefore, we keep them as distinct species before more specimens are available, especially those from Indo-China.

The shape of right paramere apex is quite different among specimens of this species. (Figs 103, 104) According to examined materials of some other species in Physoderina, we speculate that in this subtribe the shape of right paramere is an intraspecifically variable character, but less variable between different genera (Figs 95–110). Therefore, this character has no taxonomic importance at both species and genus levels.

Holotype (IZAS): male, body length = 8.3 mm, pin mounted, genitalia dissected and deposited in microvial pinned under specimen, “China, Yunnan, Ruili / Dengga to Mafengshan; / 23.95285°N, 97.59808°E / 23.94485°N, 97.55647°E”; “927–1207m; 2009.VIII.10 / SHI H. L. leg.; Beating / Inst. of Zool., CAS / 瑞丽市等噶至麻风山"; “IOZ(E)1891845”; "HOLOTYPE ♂/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label][Figs 52, 84, 105]. Paratypes (10 males, 13 females): Yunnan: 2 males (IZAS), “China, Yunnan, Ruili / Dengga to Mafengshan; / 23.95285°N, 97.59808°E / 23.94485°N, 97.55647°E”; “927–1207m; 2009.VIII.10 / SHI H. L. leg.; Beating / Inst. of Zool., CAS / 瑞丽市等噶至麻风山"; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. 1 male 2 females (IZAS) [the male preserved in 100% alcohol, females pinned], “CHINA, Yunnan, Ruili, / Dengga to Sepeng bridge; / 23.95285°N, 97.59808°E / 23.97518°N, 97.56944°E”; “927–807m; 2009.VIII.11 / SHI H. L. leg., Beating; / Inst. of Zool., CAS / 瑞丽市等噶至色蓬桥”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label] [Fig. 134]. 1 female (HBUM), “2005-VIII-3 / 云南瑞丽市勐秀 / 2150m 毛本勇 / 河北大学博物馆” [2005-VIII-3 / Yunnan, Ruili, Mengxiu / 2150m Mao Benyong leg. / Hebei University Museum]; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. 1 male 1 female (CBW), “CHINA, Yunnan Prov., / Xishuangbanna, Menglun / Reserve, west part. 570m / 2009.VI.2; BI Wenxuan leg.”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. Guangdong: 1 female (IZAS), “CHINA, Guangdong Prov., / Shixing County, Mt. Che- / baling, Xianrendong; / 24.73478°N, 114.20727°E”; “508m; 2008.VII.23 / vegetation beating; / LIANG Hongbin leg. / Ins. of Zool., CAS”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label][Fig. 119]. 1 female (IZAS) “CHINA, Guangdong / Shixing, Chebaling / Chayuan (Tea Garden) / N24.72320, E114.25640”; “354m, 2008.7.27 day / Liang H. B.; vegetation; Institute of Zoology / 广东始兴县车八岭”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. Hainan: 1male, 1 female (IZAS), “CHINA, Hainan, / Ledong, Jianfengling, / 5th area; beating / 18.73263°N, 108.87023°E”; “978m; 2009.XII.3; Day / Lin Meiying coll. / Inst. of Zoology, CAS / 尖峰岭核心区五区”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label][Fig. 17]. 1 male (IZAS) [preserved in 100% alcohol], “CHINA, Hainan, Wuzhi- / shan Mt., way to the peak; / beating; / 18.90161°N, 109.68844°E”; “997m; 2009.XI.28 / LIANG Hongbin leg. / Inst. of Zool., CAS / 海南五指山主峰”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. Vietnam: 1 female (NNML), “Museum Leiden / Viet Nam (Dak Lak Prov.) Chu / Yang Sin N. P.: 6–8 km S of / head quarters: S of Dam / construction-site. 2–10.vi.2007. / Leg. C. van Achterberg, R. de / Vries & E. Gasso Miracle”; “primary evergreen forest near / stream; in malaise traps; 800- / 820m, 12°26'26.3"N / 108°19'58.5"E”. Laos: 2 males, 2 females (NHMB), “LAOS-NE, Houa Phan prov., / 20°11–13'N 103°59'–104°01'E, / Ban Saluei – Phou Pane Mt., / 9.–17.vi.2009, 1300–1900 m, / Michael Geiser leg.”; “NHMB Basel, NMPC Prague / Laos 2009 Expedition: / M. Brancucci, M. Geiser, / Z. Kraus, D. Hauck, V. Kubáň”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. 2 males, 2 females (NHMB), “LAOS-NE, Houa Phan prov., / ~20°13'N, 104°00'E, / PHOU PANE Mt., / 1.–16.vi.2009, 1350~1500 m, / M. Brancucci leg.”; “NHMB Basel, NMPC Prague / Laos 2009 Expedition: / M. Brancucci, M. Geiser, / Z. Kraus, D. Hauck, V. Kubáň”; “PARATYPE/ Allocota bicolor/ new species / Des. SHI H. L. 2011” [red label]. Thailand: 1 female (CMB), “Thailand, Prov. Nan. Bo Khua/ 19.4-7.5.2004/ Moravec Petr”.

Elytra metallic blue, disc without reddish patch; metasternum and abdomen not darker than prosternum; front angles of pronotum with long setae (Fig. 153); elytral 7th interval with setigerous pores; protibiae with cleaning spur present but very fine (Fig. 141); internal sac of aedeagus without setose area near middle, apex of secondary flagellum simple.

From dorsal color, this new species resembles Allocota viridipennis Motschulsky or Allocota cyanipennis Heller, but it is actually more isolated in the genus than it appears. The new species can be distinguished from the other three species of the genus in having: (1) protibiae with cleaning spur present, but absent in other species; (2) males with adhesive hairs rudimentary on first three mesotarsomeres, but well developed on 1st and 2nd mesotarsomeres in other species; (3) males with terminal labial palpomeres truncate, but fusiform in the other species; (4) setae on pronotal front angles longer than in other species; (5) internal sac of aedeagus without setose area near middle.

Body length 7.5–8.7 mm; head and pronotum vivid orange red; mouthparts yellowish brown, palpomeres dark brown, apex of terminal labial and maxillary palpomeres yellow; base of 1st antennomere yellowish, remaining part of 1st antennomere and 2nd to 4th antennomeres piceous, gradually turning to brownish after 5th antennomere; elytra metallic blue, elytral suture and lateral margins metallic; legs with most parts much darker than pronotum, coxae, trochanters and base of femora yellow, apical part of femora and tibiae nearly black, tarsomeres usually dark brown; ventral side of head and prosternum almost the same color as dorsal side; metasternum and abdomen yellowish, not darker than prosternum. Head glabrous, without punctures, microsculpture indistinct; males with terminal labial palpomeres expanded at apex, truncate but not strongly securiform. Pronotum glabrous, cordiform, widest at about apical one-third; ratio PW/PL 1.30 to 1.40; pronotal base briefly but distinctly lobed; disc moderately convex, microsculpture indistinct, without punctures; front angles with long setae, distinctly longer than other species of the genus; hind angles with a few secondary setae; lateral explanate areas moderately wide, without punctures; lateral margins slightly expanded in middle, usually slightly sinuate before hind angles; hind angles usually rounded or subrectangular, not sharp or projected (Fig. 157), but in two females from Guangdong, hind angles sharp, acute, distinctly projected (Fig. 156); basal foveae very shallow, without punctures; median line very fine, indistinct; disc finely transversely rugose. Elytra with striae slightly sulcate, finely punctate; 7th, 8th striae and base of the remaining striae nearly unimpressed; intervals not distinctly convex, without accessory setae; microsculpture absent; 3rd, 5th and 7th intervals with four to ten setigerous pores, their positions variable, most of them on center of interval; lateral margins with some short and fine but distinct setae. Legs. Protibiae with cleaning spur present, but fine and short (Fig. 141); males with adhesive hairs rudimentary (two rows, weakly present near apex) on first three mesotarsomeres. Male genitalia with median lobe of aedeagus stout, apex slightly bent to right side in dorsal view, right margin not curved before apical lamella; apical lamella placed on right side, triangular, length equal to its basal width, apex not sharp; internal sac without setose area near middle; secondary flagellum reaching apical orifice, apex simple; trumpet-form expansion with ventral margin not expanded (Fig. 84). Female genitalia. Spermatheca straight, tubiform, moderately long; apical half slightly widened, apical third with distinct ring-sculpture; spermathecal gland inserted near middle of spermatheca, not branched, fine, base not distinctly thickened, much longer than spermatheca (Fig. 134). Apical segment of ovipositor very short, slightly longer than width; outer margin straight; inner margin curved, with fine setae on apical half; membranous extension fine, placed on the outer side of apex (Fig. 119).

(Map 6). China (Yunnan, Guangdong, Hainan), Vietnam, Laos, Thailand.

The name “bicolor” is a Latin noun, referring to the sharp contrast between the orange head and pronotum and the metallic blue elytra.

Two female paratypes coming from Guangdong are slightly different from the holotype from Yunnan in pronotal shape. Their pronotum hind angles are sharper and more projected, and lateral margins are more strongly sinuate before hind angles (Fig. 156) than in the holotype (Fig. 157). But the other characters are the same, and we consider they are just geographical variations.

The new species is a special lineage in Allocota, and could be the most primitive one of the genus, based on some primary characters such as presence of cleaning spur on protibiae. Moreover, the female reproductive system and male secondary characters (labial palpomeres, adhesive hairs on tarsomeres, setae on terminal sternum) are similar to Physodera eschscholtzii, but they are totally different in ovipositor. Similarity between these two species may also suggest the affinity between genera Allocota and Physodera.

Lachnoderma cinctum Macleay, 1873, by monotypy.

Dorsal side densely and evenly pubescent, pubescence long and erect; labrum and mandibles with long accessory setae (as long as primary ones on labrum); mentum tooth bifid; basal foveae of pronotum wide, not forming groove; elytral striae very coarsely punctate; males with adhesive hairs absent on all tarsomeres; aedeagus with main flagellum of internal sac projected out from apical orifice. This genus is most similar to Dasiosoma; their differences are presented in the diagnosis of Dasiosoma.