Paraphaea Bates, 1873

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 : 10-12

Genus Paraphaea Bates, 1873 status resurrected

Paraphaea Bates 1873: 312; Chaudoir 1877: 236 (synonymized with Anchista Nietner); Jedlička 1963: 449 (in part, as a synonym of Anchista Nietner); Habu 1967: 137 (in part, as a synonym of Anchista Nietner); Habu 1982: 102 (in part, as a synonym of Anchista Nietner).

Type-species:

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

Diagnosis.

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.

Generic characters.

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.

Distribution

( 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.

Monophyly and relationships.

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.

Taxonomic comments.

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.

Key to species of Paraphaea Bates

1	Pronotum more or less pubescent; pronotum reddish brown, elytra metallic blue; the Philippines ( Fig. 36 )	Paraphaea philippinensis ( Jedlička)
-	Pronotum glabrous; elytra uniform brown or with bicolored pattern, not metallic	2
2	Third interval with more than three setigerous pores, 3rd 5th and 7th intervals with some secondary setigerous pores in some specimens; Taiwan ( Figs 2 , 34 )	Paraphaea formosana ( Jedlička)
-	Third interval with two setigerous pores, intervals without secondary setigerous pores	3
3	Pronotum widest at apical third, lateral margins slightly sinuate before hind angles; elytra with background dark, each side with an elongate pale patch before middle ( Fig. 1 ); widely distributed in southeast Asia	Paraphaea binotata (Dejean)
-	Pronotum widest near middle, lateral margins distinctly sinuate before hind angles; elytra with disc yellowish, lateral margins and apex dark ( Fig. 3 ); Hainan and Indo-China	Paraphaea minor sp. n.

Figures

Map 1. Known distribution of genus Paraphaea I: ● Paraphaea binotata (Dejean).

Map 2. Known distribution of genus Paraphaea II: ▲ Paraphaea minor sp. n. ◆ Paraphaea philippinensis (Jedlicka) ★ Paraphaea formosana (Jedlicka).

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).

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 31 - 36. Type materials, scale bars = 2.0 mm: 31 Lectotype of Plochionus binotatus Dejean [= Paraphaea binotata (Dejean)] 32 Lectotype of Calleida discophora Chaudoir [= Paraphaea binotata (Dejean)] 33 Lectotype of Paraphaea signifera Bates [= Paraphaea binotata (Dejean)] 34 Holotype of Parena formosana Jedlicka [= Paraphaea formosana (Jedlicka)] 35 Holotype of Paraphaea minor sp. n. 36 Holotype of Coptodera philippinensis Jedlicka [= Paraphaea philippinensis (Jedlicka)].

Figures 1 - 6. Habitus, scale bars = 2.0 mm: 1 Paraphaea binotata (Dejean), a specimen from Yunnan 2 Paraphaea formosana (Jedlicka), a specimen from Taiwan 3 Paraphaea minor sp. n., a paratype from Hainan 4 Anchista brunnea (Wiedemann), a paralectotype of picea Chaudoir from India 5 Anchista fenestrata fenestrata (Schmidt-Goebel), a specimen from Nepal 6 Anchista fenestrata fenestrata (Schmidt-Goebel), a specimen from Nepal.