Crossoglossa Chaudoir, 1872

Shi, Hongliang & Liang, Hongbin, 2023, Taxonomic revision of the genus Parena Motschulsky, 1860 (Coleoptera, Carabidae, Lebiini, Metallicina), Zootaxa 5286 (1), pp. 1-144 : 25-27

Subgenus Crossoglossa Chaudoir, 1872

Type-species: Crossoglossa testacea Chaudoir, 1872 . Subsequently designated by Jeannel, 1949: 971.

Crossoglossa Chaudoir, 1872: 177 ; Chaudoir, 1878: 151 (misspelled as Ceroglossa ); Jakobson, 1907: 395; Alluaud, 1917: 85; Andrewes, 1919: 483 (synonymized with Phloeodromius ); Jeannel, 1949: 972; Jedlička, 1952: 210.

Crossoglossa was erected by Chaudoir (1872) to accommodate his three species without considering the earlier name Parena Motschulsky, 1860 . The name Crossoglossa was then widely used, and several species were described under that name (e.g., Bates, 1873; Alluaud, 1917). However, the type species of Crossoglossa was not fixed until Jeannel (1949) designated Crossoglossa testacea Chaudoir for it. The concept of Crossoglossa (sensu Bates, 1873; Alluaud, 1917) was similar to the genus Parena at present until Andrewes (1919, 1928) synonymized it with Phloeodromius and Parena . Thereafter, Crossoglossa was used as a subgenus name in different concepts ( Jeannel, 1949; Jedlička, 1952).

In the present study, Crossoglossa is treated as a subgenus of Parena to accommodate species related to P. testacea , which are different from members of the other two subgenera of the genus in the following unique character states: (1) pronotum much wider (PW/PL 1.50–1.82) than other species of the genus; (2) mentum without setae, lateral lobes extremely large, inner margin nearly straight, epilobes very narrow ( Fig. 3A ); (3) male genitalia rather slender, endophallus with the primary sclerite rudimentary; (4) gonocoxite II of ovipositor dichotomous.

The present concept of the subgenus Crossoglossa is similar to that of Jedlička (1952), containing nine species distributed through East Asia, all parts of the Oriental Realm, Papua New Guinea, and easternmost to the Solomon Islands. They are not known from Africa or Australia. Three species groups are recognized in the subgenus, each of them containing one or several strictly allopatric species.

The exceptional male and female genital characters suggest Crossoglossa is monophyletic. However, its relationship in the genus Parena is difficult to infer before a comprehensive phylogenetic analysis is conducted. Based on the modified male and female genitalia, Crossoglossa is considered to be a highly specialized group in the genus.

Diagnostic characters. Dorsal surface generally unicolor (except P. sciakyi sp. n.), pale yellow to reddish brown. Tempora very short, abruptly narrowed behind eyes, length of tempora plus neck-constriction approximately one-third of diameter of the eye; accessory setae on antennomeres 1–3 absent or very short and sparse; postgenae with a pair of suborbital setae; mandibles extremely widened, semicircular, retinacular ridge of right mandible reduced to an elongate retinacular tooth; mentum without a pair of setae, lateral lobes extremely large, inner margins nearly straight, outer and apical margins widely arcuate, epilobes very narrow, apex much shorter than lateral lobes. Pronotum strongly transverse, PW/PL 1.50–1.82, generally greater than 1.60; much wider than head, PW/HW = 1.05–1.26, generally greater than 1.10; lateral explanations very wide; lateral margins usually slightly sinuate before posterior angles; posterior angles obtuse-rounded. Elytral interval 3 with three setigerous pores, first one very close to elytra base, near level of scutellar apex; disc with large depressions near middle of intervals 3 to 6 (except for P. levata ), lateral sides depressed near anterior third. Apex of abdominal sternite VII with two or three setae on each side, straight or slightly notched in males, straight or projected at middle in females. Male mesotarsi with biseriate ventral adhesive setae on basal three tarsomeres. Male genitalia with median lobe slender, AL/AW 5.4–6.8; apical lamella thin and wide, apical orifice opened to left; endophallus with primary sclerite reduced to a small triangular piece near base, apical sclerite ribbon-form, squamate sheath large and heavily scaled, squamate sac simple or divided. Female ovipositor with dichotomous gonocoxite II, inner branch with two ensiform setae apically, outer branch with two or three ensiform setae apically.

Key to species of subgenus Crossoglossa Chaudoir

1. Dorsum reddish yellow to reddish brown, tarsi same as or slightly darker than dorsum............................... 2

- Dorsum pale yellow, tarsi black, distinctly darker than dorsum. (3. P. testacea group)............................... 5

2. Elytral sutural angles only faintly pointed ( Fig. 14A ), outer apical angles completely rounded ( Fig. 14I ); striae well incised, finely punctate; intervals slightly convex (1. P. cavipennis group)............................ [1] P. cavipennis (Bates)

- Elytral sutural angles distinctly pointed, extended as short spines ( Figs 14B, 14C ), outer apical angles more or less rounded-angulate ( Fig. 14J ); striae not incised, replaced by rows of fine punctures; intervals flat (2. P. laesipennis group).......... 3

3. Elytra not depressed on intervals 3 to 6; Sumatra, Borneo.................................... [3] P. levata Andrewes

- Elytra more or less depressed near middle on intervals 3 to 6; other localities...................................... 4

4. Elytral outer apical angles only weakly angulate, forming rounded obtuse angles; apical lamella of aedeagus wider, LL/LW 1.50–1.65; widely in East Asia and Southeast Asia, except the Malay Archipelago............... [2] P. laesipennis (Bates)

- Elytral outer apical angles distinctly angulate, forming sharper obtuse angles; apical lamella of aedeagus narrower, LL/LW approximately 1.75; Java ............................................................ [4] P. obenbergeri Jedlička

5. Elytral disc with large black patch; sutural angles not pointed ( Fig. 14H ); Solomon Islands........... [9] P. sciakyi sp. nov.

- Elytra uniformly yellow; sutural angles more or less pointed; other localities...................................... 6

6. Elytral sutural angles sharply pointed, forming short spines ( Figs 14D, 14E )....................................... 7

- Elytral sutural angles gently pointed, forming small denticles ( Figs 14F, 14G )..................................... 8

7. Antennomere 1 yellow, other ten antennomeres black; elytral intervals with dense punctures, similar density as punctures in striae, so that striae poorly defined; apical lamella of aedeagus shorter, LL/LW = 1.0–1.1; India, Nepal, Vietnam, China.................................................................................... [5] P. testacea (Chaudoir)

- Antennomeres 1 to 3 yellow, other eight antennomeres black; elytral intervals with sparse punctures, much sparser than punctures in striae, so that striae clearly defined; apical lamella of aedeagus longer, LL/LW approximately 1.3; Sulawesi ................................................................................ [6] P. sulawesiensis Kirschenhofer

8. Tibiae yellow; elytra disc with depressions distinct; Maluku Islands, Adonara Island, Papua New Guinea, E. Java, Sumatra, the Philippines........................................................................[7] P. mellea (Chaudoir)

- Tibiae black; elytra disc with depressions absent or very shallow; W. Java, Borneo.............. [8] P. cruralis Andrewes

Figures

FIGURE 3. A–D Labia of Parena spp.: A P. (Crossoglossa) cavipennis (Bates), Beijing. B P. (Parena) latecincta (Bates), Zhejiang. C P. (Bothynoptera) dorsigera (Schaum), Guangxi. D P. (Bothynoptera) tripunctata (Bates), Beijing. E–F Heads of Parena spp. in lateral view: E P. (Bothynoptera) taiwana Hua, Taiwan, suborbital setae present. F P. (Parena) latecincta (Bates), Yunnan, suborbital seta absent. Scale bars = 0.5mm.

FIGURE 14. Elytra apex of Parena (Crossoglossa) spp. A–H Elytra sutural angle (left), scale bar = 0.2mm. A P. cavipennis (Bates), Shaanxi. B P. laesipennis (Bates), Vietnam. C P. levata Andrewes, Borneo. D P. testacea (Chaudoir), Guangdong. E P. sulawesiensis Kirschenhofer, Sulawesi. F P. cruralis Andrewes, Borneo. G P. mellea (Chaudoir), Philippine. H P. sciakyi sp. n., holotype. I–L Elytra outer apical angle (left), scale bar = 0.5mm. I P. cavipennis (Bates), Shaanxi. J P. laesipennis (Bates), Taiwan. K P. sciakyi sp. n., holotype. L P. testacea (Chaudoir), Guangdong.