Bothynoptera Schaum, 1863

Shi, Hongliang & Liang, Hongbin, 2023, Taxonomic revision of the genus Parena Motschulsky, 1860 (Coleoptera, Carabidae, Lebiini, Metallicina), Zootaxa 5286 (1), pp. 1-144 : 45-47

Subgenus Bothynoptera Schaum, 1863

Type-species: Bothynoptera dorsigera Schaum , fixed by monotypy.

Bothynoptera Schaum, 1863: 75 ; Chaudoir, 1872: 181; Jakobson, 1907: 395; Jedlička, 1951: 59; Jedlička, 1963: 445; Habu, 1967: 149 (synonymized with Parena Motschulsky ).

Bothynoptera was described by Schaum (1863) for B. dorsigera . Later, some Parena species with relatively narrow pronotum were described under this genus ( Bates, 1873, Andrewes, 1947, Jedlička, 1951). In Jedlička's monograph ( Jedlička, 1963), Bothynoptera was regarded as a separate genus including six species, and distinguished from Parena by the differences in tempora length and pronotum shape. However, the generic characters mentioned are not in accord with all of these six species. Habu (1967) pointed out that Bothynoptera has no generic differences from Parena , and synonymized these two genera.

In the present study, we find the following important diagnostic characters supporting a new concept of subgenus Bothynoptera : (1) first setigerous pore on elytral interval 3 distant from elytra base, on a level much behind scutellar apex ( Figs 4D, 4E, 4F ); (2) male mesotarsomere 1 without adhesive setae ventrally ( Fig. 7 ) in most species; (3) antennomeres 1–3 with well-developed accessory setae ( Fig. 1B ). (4) mandibles rounded-triangular, less widened than in the other two subgenera ( Fig. 2 ); (5) retinacular ridge of right mandible shortened but not reduced to a retinacular tooth ( Fig. 2 ); (6) apex of mentum epilobes extending slightly beyond lateral lobes ( Figs 3C, 3D ). The above character states differentiate subgenus Bothynoptera from other members of Parena and are thought to be synapomorphies supporting the monophyly of the subgenus. In addition to the above characters, the following characters present in subgenus Bothynoptera probably have taxonomical importance as well: dorsum not metallic; pronotum relatively narrow; elytra discal depressions very shallow or absent; apex of abdominal sternite VII straight in both sexes; male genitalia in stout form in most species.

Most of these above character states are only present in subgenus Bothynoptera and distinguish this subgenus. However, the second character state (adhesive setae absent from male mesotarsomere 1) is also found in five species ( P. latecincta , P. monticola , P. politissima , P. fulva sp. n., and P. plagiata ) of subgenus Parena . Based on several other important differences, including the pronotum relatively wide, postgenae without a long seta, terminal ventrite slightly emarginate, and the first pore on elytral interval 3 very close to elytra base, these species are considered not to be related to species of Bothynoptera .

Subgenus Bothynoptera contains fourteen species, mainly distributed in East Asia, and many of them have relatively wide distributional ranges. China has the most diverse fauna of the subgenus, with twelve species recorded: five of them are endemic to China, five have ranges extended to the south slope of the Himalayan Range and the north part of Southeast Asian counties, and four species range Japan, Korea, and the Russian Far East. The subgenus Bothynoptera is not found in Africa, Australia, South India, or the Malay Archipelago.

Five species groups are recognized in the subgenus, based mainly on differences in male genitalia (except for P. heteronycha for which male genitalia are unknown). The P. tripunctata species group has the highest species diversity, including eight species with very similar stout male genitalia. The P. taiwana group (containing only one species) is the most atypical in the subgenus for the presence of ventral adhesive setae on male mesotarsomere 1 and rather slender male genitalia; but based on the character states mentioned above, it should be assigned to the subgenus Bothynoptera .

Diagnostic characters. Dorsal surface unicolorous or elytra with light or dark patches, without metallic hue. Tempora abruptly or gradually narrowed behind eyes, length of tempora plus neck-constriction usually more or less longer than one-third of diameter of eye; postgenae with a pair of suborbital setae, generally in similar length as supraorbital setae (but very short in P. heteronycha ); antennomeres 1–3 with distinct accessory setae, one or two of them on the antennomere 2 in similar length as two primary ones. Mandibles moderately widened, rounded-triangular, retinacular ridge of right mandible present, one-third to two-thirds length of terebral ridge; mentum with a pair of setae, setae normally longer than terminal labial palpomere (but very short in P. tesari , P. obsucra and P. heteronycha ); lateral lobes short and narrow, apex rounded-angulate, inner margins strongly oblique, outer margins nearly straight or slightly arcuate, epilobes relatively wide, apex extending beyond lateral lobes. Pronotum usually relatively narrow, PW/PL 1.19–1.43, narrower or slightly wider than head, PW/HW = 0.85–1.12; lateral explanations narrow; lateral margins clearly sinuate before posterior angles in most species; posterior angles forming indistinct right angles in most species. Elytral interval 3 with three setigerous pores in most species (with four or more in P. dorsigera ); first pore distant from elytra base, on a level much behind scutellar apex; disc without depression or very faintly depressed near middle of interval 5, lateral sides slightly depressed near anterior third. Abdominal sternite VII with two setae on each side in most species (with three in P. tesari , P. obsucra and P. kurosai ), apex straight in both sexes. Male mesotarsomere 1 without adhesive setae ventrally (except P. taiwana ), mesotarsomere 2 and 3 with adhesive setae ventrally. Male genitalia with median lobe very stout (AL/AW = 3–4) in most species, but slightly slender in P tesari group (AL/AW = 4.5–5), very slender in P. taiwana group (AL/AW = 5.5–6); endophallus with primary sclerite well chitinized, flared basal expansion well-developed, apical flagellum present, more or less extended to apex; apical sclerite V-shaped. Female ovipositor with gonocoxite II nearly rectangular in most species, apex with five to seven ensiform setae, subequally arranged or grouped at angles.

Key to species of subgenus Bothynoptera Schaum

1. Tarsal claws asymmetric ( Fig. 6F ): inner claw elongate with pectinations only present on basal half, outer claw normal (opposite for prolegs); elytra with two distinct small tubercles at basal quarter and middle of interval 6; elytra background color of a gradient from basal dark brown to apical light yellow, elytral setigerous pores forming black umbilicular spots ( Fig. 26 ). (4. P. heteronycha group)................................................................ [10] P. heteronycha sp. n.

- Tarsal claws symmetric ( Fig. 6E ); elytra without tubercles; elytra of different color, setigerous pores same color as background.......................................................................................... 2

2. Elytra with apical truncation distinct, bisinuate, straight or concave inward; outer apical angles prominent ( Figs 5G–5J ). (5. P. dorsigera group)...................................................................................... 3

- Elytra with apical truncation indistinct, evenly rounded, outer apical angles completely rounded ( Fig. 5E )............... 4

3. Elytral interval 3 with four setigerous pores, five or six in some specimens; elytra depressed at pores, forming fovea slightly wider than interval width.......................................................... [11] P. dorsigera (Schaum)

- Elytral interval 3 with three setigerous pores, elytra not depressed at pores........................ [12] P. kurosai Habu

4. Tempora gradually narrowed behind eyes, length of tempora plus neck-constriction two-thirds of diameter of eye or more (6. P. tesari group) ....................................................................................... 5

- Tempora abruptly narrowed behind eyes, length of tempora plus neck-constriction at most slightly greater than one-third of diameter of eye....................................................................................... 6

5. Tempora shorter, length of tempora plus neck-constriction approximately two-thirds of diameter of eye; apical lamella of male genitalia longer, LL subequal to LW; China, Vietnam....................................... [13] P. tesari (Jedlička)

- Tempora longer, length of tempora plus neck-constriction subequal to diameter of eye; apical lamella of male genitalia very short, LL about half as LW; Bhutan, Nepal................................................ [14] P. obscura Mateu

6. Elytra interval strongly punctate, distance between interval punctures one to two times as that between stria punctures; antennae longer, exceeding level of first discal pore on elytra; dorsal surface yellow, elytra with a pair of dark spots behind middle, sometimes without these spots or with a pair of additional spots near base ( Fig. 36 ); median lobe of aedeagus slender. (7. P. taiwana group)........................................................................ [15] P. taiwana Hua

- Elytra interval without puncture or with punctures much sparser than stria punctures; antennae shorter, hardly reaching elytra base; elytra pattern not as above; median lobe of aedeagus stout. (8. P. tripunctata group)............................ 7

7. Elytra unicolor or disc slightly paler, not forming well-defined pattern........................................... 8

- Elytra with well-defined pattern.......................................................................... 9

8. Elytra unicolor, reddish brown to dark brown........................................... [16] P. tripunctata (Bates)

- Elytra yellowish brown, disc gradually turn to pale yellow from base, forming an obscure large pale patch............................................................................................[17] P. shapingensis Xie & Yu

9. Elytra reddish brown, with large pale serrate band near middle........................... [19] P. malaisei (Andrewes)

- Elytra light yellow to reddish brown, with one to four dark patches............................................. 10

10. Elytra with one central black patch behind middle, reaching third or fourth interval outward; Japan, Korea, Russia (Fareast) and east China...................................................................... [18] P. monostigma (Bates)

- Elytra with three or four dark patches; southwest China, Laos................................................. 11

11. Elytra dark reddish brown, with four black patches, the basal group close to elytra humeri, reaching seventh or eighth interval outward....................................................................... [20] P. quadrisignata Mateu

- Elytra light yellow to yellowish brown, basal black patches distant from humeri, at most reaching sixth interval outward.. 12

12. Apical margin of labrum strongly emarginate ( Fig. 46C ); dorsum pale yellow; pronotum disc with a pair of dark stripes; elytra with four black patches: a pair of basal stripes and a pair of well separate subapical spots; Sichuan (Muli), Xizang (Cona)................................................................................... [21] P. emarginata sp. n.

- Apical margin of labrum straight; dorsum reddish yellow; pronotum without dark patch; elytra subapical patches more or less continuous at elytral suture forming one large middle patch................................................... 12

13. Elytra darker; basal patches distant from elytral base (not covering first discal pore); subapical patch concave at anterior margin; in dorsal view, median lobe of aedeagus strongly narrowed at apex, LW subequal to LL; Sichuan (Luding).......................................................................................... [22] P. gonggaica sp. n.

- Elytra lighter, basal patches close to elytral base (covering first discal pore); subapical patch protruded at anterior margin; in dorsal view, median lobe of aedeagus slightly narrowed at apex, LW about twice as LL; Sichuan (Kangding, Ebian)..........................................................................................[23] P. triguttata sp. n.

Figures

FIGURE 1. Antennae and mouthparts of Parena spp. A–D Line drawings of basal three antennomeres, view of lateral surface: A P. (Crossoglossa) cavipennis (Bates). B P. (Bothynoptera) malaisei (Andrewes). C P. (Parena) africana (Alluaud). D P. (Parena) ruficornis sp. n. E Antennomeres 7–11 of P. (Parena) nigrolineata (Chaudoir), showing the sensory pits on dorsal surface. F–G Labrum of P. (Crossoglossa) cavipennis (Bates), Beijing: F dorsal surface. G ventral surface. H Maxilla of P. (Crossoglossa) cavipennis (Bates), Beijing. Scale bars = 0.2mm.

FIGURE 2. Mandibles of Parena spp.: left mandible, dorsum (ld); right mandible, dorsum (rd); left mandible, venter (lv); right mandible, venter (rv). A P. (Crossoglossa) cavipennis (Bates), Shaanxi. B P. (Parena) latecincta (Bates), Zhejiang. C P. (Bothynoptera) dorsigera (Schaum), Guangxi. D P. (Bothynoptera) taiwana Hua, Taiwan. E P. (Bothynoptera) tripunctata (Bates), Beijing. Scale bar = 0.5mm. Abbreviations: rr retinacular ridge of right mandible; vg ventral groove.

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 4. Elytral base (left) of Parena spp. A P. (Parena) bicolor Motschulsky, Sumatra. B P. (Parena) africana (Alluaud), Cameroon. C P. (Crossoglossa) laesipennis (Bates), Taiwan. D P. (Bothynoptera) tripunctata (Bates), Beijing. E P. (Bothynoptera) dorsigera (Schaum), Guangxi. F P. (Bothynoptera) heteronycha sp. n., Laos, paratype. Scale bars = 0.5mm (a for fig. D, b for figs A B E F, c for fig. C). Abbreviations: bp elytral basal pore; sa the apex of scutellum; d1 the first discal setigerous pore.

FIGURE 5. Elytral apex (left) of Parena spp. A P. (Parena) africana (Alluaud), Cameroon. B P. (Parena) ruficornis sp. n., Central Africa, paratype. C P. (Parena) valeriae Facchini, holotype. D P. (Parena) madagascariensis (Alluaud), Madagascar. E P. (Bothynoptera) tesari (Jedlička), Taiwan. F P. (Bothynoptera) heteronycha sp. n., Laos, paratype. G P. (Bothynoptera) kurosai Habu, Japan. H P. (Bothynoptera) dorsigera (Schaum), Yunnan. I P. (Bothynoptera) dorsigera (Schaum), Beijing. J P. (Bothynoptera) dorsigera (Schaum), Japan. Scale bars = 0.5mm

FIGURE 6. A Hind wing, P. (Crossoglossa) cavipennis (Bates), Beijing, scale bar = 1mm. Abbreviations: oc oblongum cell; wc wedge cell. B–G Tarsals claws of right hindlegs for Metallicina spp., scale bar = 0.2mm: B Pachycallida rufoplagiata Jeannel, Madagascar, Sept Lacs, female. C Metallica viridipennis Chaudoir, Cameroon, Bois des Singes, male. D P. (Crossoglossa) sciakyi sp. n., holotype. E P. (Bothynoptera) kurosai Habu, Nepal. F P. (Bothynoptera) heteronycha sp. n., Laos, paratype. G P. (Parena) latecincta (Bates), Vietnam. H Protibia of P. (Parena) nigrolineata (Chaudoir), scale bar = 0.5 mm. I Mesotarsomeres of male of P. (Parena) nigrolineata (Chaudoir), adhesive hairs present on mesotarsomere 1, scale bar = 0.2mm. J Mesotarsomeres of male for P. (Parena) latecincta (Bates), adhesive hairs absent from mesotarsomere 1, scale bar = 0.2mm.

FIGURE 7. Diagram of the ventral surface of male mesotarsomeres 1–3, white strips representing the variety of adhesive setal patterns in different species of genus Parena.

FIGURE 26. Parena (Bothynoptera) heteronycha sp. n. A Habitus of holotype. B Habitus of paratype. C Mouthparts of holotype. D Labels of holotype. E Metatarsus. F Mesotarsus. G Protarsus. Scale bars = 2 mm for A, B; 0.5mm for E, F, G.

FIGURE 36. Habitus of Parena (Bothynoptera) taiwana Hua. A Male from Guangxi. B Female from Taiwan. C Male from Taiwan. Scale bars = 2 mm.

FIGURE 46. Parena (Bothynoptera) emarginata sp. n. A Habitus of holotype. B Habitus of paratype. C Mouthparts of holotype. D Labels of holotype. Scale bars = 2 mm.