Physodera Eschscholtz, 1829

Shi, Hongliang, Zhou, Hongzhang & Liang, Hongbin, 2013, Taxonomic synopsis of the subtribe Physoderina (Coleoptera, Carabidae, Lebiini), with species revisions of eight genera, ZooKeys 284, pp. 1-129 : 35-38

Genus Physodera Eschscholtz, 1829 Habitus: Figs 11, 12 male genitalia: Figs 76 100 female genitalia: Figs 115, 116 130, 131

Physodera dejeani Eschscholtz 1829: 8; Schmidt-Göbel 1846: 46; Lacordaire 1854: 130; Van de Poll 1889: 251; Heller 1923: 304 (key to species); Jedlička 1963: 300.

Nepalotarsus Mateu 1990: 156. Nomen nudum. [Synonym]

Type-species:

Physodera dejeani Eschscholtz, 1829, by monotypy.

Diagnosis.

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.

Generic characters.

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.

Distribution.

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

Monophyly and relationships.

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.

Taxonomic comments.

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.

Figures

Figure 76. Median lobe of male genitalia, right-lateral, ventral, left-lateral, dorsal views of Physodera dejeani Eschscholtz, a specimen from Yunnan.

Figures 7 - 12. Habitus, scale bars = 2.0 mm: 7 Anchista fenestrata subpubescens Chaudoir, a specimen from Pakistan 8 Anchista nubila Andrewes, a specimen from India 9 Metallanchista perlaeta (Kirschenhofer), a specimen from Sumatra 10 Metallanchista perlaeta (Kirschenhofer), a specimen from Java 11 Physodera dejeani Eschscholtz, a specimen from Yunnan 12 Physodera eschscholtzii Parry, a specimen from Yunnan.

Figures 95 - 110. Left and right parameres of male genitalia, scale bars = 0.2 mm: 95 Paraphaea binotata (Dejean), a specimen from Guangxi 96 Paraphaea minor sp. n., holotype 97 Anchista brunnea (Wiedemann), a paralectotype of picea Chaudoir 98 Anchista pilosa sp. n., holotype 99 Metallanchista laticollis sp. n., holotype 100 Physodera amplicollis van de Poll, a specimen from Taiwan 101 Diamella cupreomicans (Oberthuer), a specimen from Yunnan 102 Diamella arrowi (Jedlicka), holotype 103 Allocota aurata (Bates), a specimen from Tonkin 104 Allocota aurata (Bates), a specimen from Shaanxi 105 Allocota bicolor sp. n., holotype 106 Lachnoderma asperum Bates, a specimen from Taiwan 107 Dasiosoma ivorense Basilewsky, a paratype from Cote d'Ivoire 108 Dasiosoma quadraticolle sp. n., holotype 109 Orionella kathmanduensis (Kirschenhofer), a specimen from Nepal 110 Endynomena pradieri (Fairmaire), a specimen from Fujian.

Figures 111 - 125. Left ovipositors of females, ventral view, scale bars = 0.2 mm: 111 Paraphaea binotata (Dejean), a specimen from Guangxi 112 Paraphaea minor sp. n., a paratype from Hainan 113 Paraphaea philippinensis (Jedlicka), holotype 114 Anchista fenestrata fenestrata (Schmidt-Goebel), a specimen from Nepal 115 Physodera dejeani Eschscholtz, a specimen from Yunnan 116 Physodera eschscholtzii Parry, a specimen from Yunnan 117 Diamella cupreomicans (Oberthuer), a specimen from Yunnan 118 Allocota aurata (Bates), a specimen from Shaanxi 119 Allocota bicolor sp. n., a paratype from Guangdong 120 Lachnoderma asperum Bates, a specimen from Zhejiang 121 Dasiosoma bellum (Habu), a specimen from Guangdong 122 Dasiosoma maindroni (Tian & Deuve), a specimen from Tonkin 123 Dasiosoma quadraticolle sp. n., a paratype from Yunnan 124 Orionella lewisii (Bates), a specimen from Zhejiang 125 Endynomena pradieri (Fairmaire), a specimen from Taiwan.

Figures 126 - 131. Internal reproductive system of females, scale bars = 0.2 mm: 126 Paraphaea binotata (Dejean), a specimen from Guangxi 127 Paraphaea minor sp. n., a paratype from Hainan 128 Paraphaea philippinensis (Jedlicka), holotype 129 Anchista fenestrata fenestrata (Schmidt-Goebel), a specimen from Nepal 130 Physodera dejeani Eschscholtz, a specimen from Yunnan 131 Physodera eschscholtzii Parry, a specimen from Hainan.

Figures 138 - 157. Characters of Physoderina. 138 - 139 Internal reproductive system of females, scale bars = 0.2 mm: 138 Dasiosoma quadraticolle sp. n., a paratype from Yunnan 139 Orionella lewisii (Bates), a specimen from Zhejiang. 140 - 142 Right protibia, ventral view, showing the reduction of cleaning spur, scale bars = 0.5 mm: 140 Allocota aurata (Bates) (cleaning spur absent) 141 Allocota bicolor sp. n. (cleaning spur fine) 142 Diamella cupreomicans (Oberthuer) (cleaning spur well developed). 143 - 146 Terminal sternum, showing the setae and male emargination, scale bars = 0.5 mm: 143 Paraphaea binotata (Dejean), male (deeply emarginate, one seta on each side) 144 Paraphaea binotata (Dejean) (straight, two setae on each side), female 145 Allocota aurata (Bates), male (moderately emarginate, two setae on each side) 146 Allocota aurata (Bates), female (straight, two setae on each side, right side unusually with an additional seta). 147 - 148 Umbilical series of 9 th interval, right elytron, scale bars = 0.5 mm: 147 Paraphaea binotata (Dejean) (umbilical series placed in one row) 148 Metallanchista laticollis sp. n. (umbilical series placed in two rows). 149 - 150 Head, showing the different shape of mandibles, scale bars = 0.5 mm: 149 Paraphaea formosana (Jedlicka) (mandibles moderately widen) 150 Allocota bicolor sp. n. (mandibles strongly widen). 151 - 153 Pronotum, showing the different length of setae on front angle, scale bars = 0.5 mm: 151 Allocota viridipennis Motschulsky, (setae short and fine) 152 Allocota cyanipennis Heller, (setae relative long) 153 Allocota bicolor sp. n., (setae very long). 154 - 155 Pronotum, showing the difference on lateral margin, scale bars = 0.5 mm: 154 Anchista fenestrata fenestrata (Schmidt-Goebel), (lateral margin slightly angulated in middle) 155 Paraphaea formosana (Jedlicka), (lateral margin completely rounded in middle). 156 - 157 Pronotal hind angle, showing the geographical variation in Allocota bicolor sp. n., scale bars = 0.2 mm: 156 Allocota bicolor sp. n., a specimen from Guangdong, (hind angle sharp) 157 Allocota bicolor sp. n., holotype from Yunnan, (hind angle slightly rounded).