Physoderina Chaudoir, 1831

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 : 7-9

Subtribe Physoderina Chaudoir

Physoderides Chaudoir 1877: 213. Type genus: Physodera Eschscholtz, 1829.

Diagnosis.

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.

Monophyly and relationships.

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.

Genera included.

Chaudoir (1877) proposed the name Physoderides 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.

Distribution.

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.

Key to genera of Physoderina

1	Head completely glabrous; pronotum usually glabrous, rarely with disc sparsely pubescent	2
-	Head at least with some pubescence on vertex; pronotum densely and equally pubescent	6
2	Mandibles moderately widened, outer margin slightly arcuate (Fig. 149); mid-lateral setae of pronotum present, lateral margins with a few accessory setae restricted to front and hind angles; median lobe of aedeagus usually with long setae (Figs 67-73), if without, main flagellum of internal sac absent (Figs 74-75)	3
-	Mandibles strongly widened, outer margin semicircular (Fig. 150); mid-lateral setae of pronotum usually absent, if present, lateral margins with numerous accessory setae nearly reaching to middle area; median lobe of aedeagus without long setae, main flagellum of internal sac well developed (Figs 76, 79)	5
3	Umbilical pores of 9th interval placed in two rows, some pores adjacent to 8th stria, others in middle of the interval (Fig. 148); males with two pairs of setae on terminal sternum; median lobe of aedeagus with apical lamella longer, bent to dorsal side (Fig. 75)	Metallanchista gen. n.
-	Umbilical pores of 9th interval placed in one row, all adjacent to 8th stria (Fig. 147); males with one pair of setae on terminal sternum; median lobe of aedeagus with apical lamella shorter, not bent	4
4	Lateral margins of pronotum completely rounded at middle (Fig. 155); males with adhesive hairs on 1st metatarsomere; males with terminal sternum deeply emarginate (Fig. 143); median lobe of aedeagus twisted to left, internal sac with main flagellum developed and sinuous (Figs 67-69)	Paraphaea Bates
-	Lateral margins of pronotum slightly angulate at middle (Fig. 154); males without adhesive hairs on metatarsomeres; males with terminal sternum moderately emarginate (as in Fig. 145); median lobe of aedeagus not twisted, internal sac with main flagellum reduced (Figs 70-73)	Anchista Nietner (part)
5	Pronotum wide, PW/PL more than 1.4; protibiae with cleaning spur developed (as in Fig. 142); 5th interval with one or two large setigerous pores near base, if additional small pores present, all pores placed before middle; aedeagus with apical part of median lobe strongly bent to right side (Fig. 76)	Physodera Eschscholtz
-	Pronotum narrow, PW/PL less than 1.4; protibiae with cleaning spur more or less reduced (Figs 140, 141); 5th interval with four to ten setigerous pores, equally placed; aedeagus with apical part of median lobe slightly bent to right side (Figs 79-84)	Allocota Motschulsky
6	Posterior supraorbital setae distant from eyes, insertions more or less tumid, forming a pair of humps (Figs 13, 45-48); elytra with fine setae along striae and on odd intervals, even intervals glabrous; internal sac with main flagel lum extraordinarily thick (Figs 77-78); apical part of trumpet-form expansion strongly expanded	Diamella nom. n.
-	Posterior supraorbital setae near eyes, insertions even, not forming humps; elytra equally pubescent on all intervals; internal sac with main flagellum fine or absent; if present, trumpet-form expansion at most with apical part moderately expanded	7
7	Mentum tooth bifid, with more than six setae; elytral striae indistinct, with very coarse punctures; mandibles with accessory setae on outer scrobe; labrum with long additional setae, as long as primary ones; internal sac with main flagellum projected beyond apical orifice (Fig. 85)	Lachnoderma Macleay
-	Mentum tooth simple, with two to four setae; elytral striae distinct, deep or shallow, at most with fine punctures; mandibles with outer scrobe glabrous, sometimes with a few fine setae along outer ridge; labrum without or with very fine additional setae, distinctly shorter than primary ones; internal sac with main flagellum at most reaching apical orifice, not projected (Figs 72, 74, 91-94)	8
8	Pronotum with basal foveae very deep, forming short grooves (Figs 24-26); elytral dorsal setigerous pores indistinct; ligula with four long setae at apex and a few short setae on dorsal surface; males with adhesive hairs reduced on all tarsomeres	Dasiosoma Britton
-	Pronotum with basal foveae shallow (Figs 7, 27-30); 5th interval with one or two distinct setigerous pores on base; ligula only with four long setae on apex; males with adhesive hairs distinct, at least on 1st, 2nd protarsomeres and 1st mesotarsomere	9
9	Mandibles moderately widened, outer margin slightly arcuate (Fig. 149); males with one pair of setae on terminal sternum; median lobe of aedeagus with apical part gradually narrowed, internal sac without main flagellum (Figs 72, 74)	Anchista Nietner (part)
-	Mandibles strongly widened, outer margin semicircular (Fig. 150); males with two pairs of setae on terminal sternum; median lobe of aedeagus with apical part equal in width, internal sac with main flagellum present but sometimes very short (Figs 91-94)	10
10	Elytral intervals slightly convex, striae distinct (Figs 27-29); 7th interval without setigerous pore; pronotal base weakly but distinctly lobed; males with adhesive hairs on first two protarsomeres only; median lobe of aedeagus with apical orifice opened apically, without seta around the orifice (Figs 91-93)	Orionella Jedlička
-	Elytral intervals flat, striae very shallow, indistinct in apical part (Fig. 30); 7th interval with some setigerous pores; pronotal base hardly lobed; males with adhesive hairs on first three protarsomeres; median lobe of aedeagus with apical orifice opened dorsally, with long setae around the orifice (Fig. 94)	Endynomena Chaudoir