Clivina paris, Balkenohl, 2025

Balkenohl, Michael, 2025, On Asian Clivinini. Revision of the westwoodi-species group of the genus Clivina Latreille, 1802 (Coleoptera, Carabidae, Clivinini), Zootaxa 5592 (1), pp. 1-70 : 8-11

Clivina westwoodi -species group

Recognition of the species group. Pronotum and elytra more or less flattened (exception C. paris sp. nov.). Female gonocoxites dimer, with coxite one and two fully movable to each other, with gonocoxite two scapiform, more or less elongated, less sclerotized, with one or two seta(e) at apex. Epipleurite formed by hyaline velum ( Figs 108 , 111 ). All setae on the reproductive tract nematiform. Labrum five-, six or seven-setose, clypeus with median part separated from clypeal-wings by more or less distinct but always clearly visible notches. Elytron with striae one to four ending free at base, interval three of elytron with four or five setigerous punctures, interval eight more or less carinate posterior humerus and apically. Abdomen with last visible abdominal sternite (segment VII) with two marginal setigerous punctures widely separated. Intermediate tibia with distinct protuberance at apical third, tarsomeres of front and intermediate leg widened in most of the species. Basal tarsomere of front leg elongated, with carina baso-laterally, laterally with acute tubercle furnished with short setae. Hind wing fully developed, specimens from New Guinea / Papua also with reduced alae. Colour piceous, fuscous or fulvous, surface with extended parts of isodiametric reticulation of different patterns.

Identification key to the species of the Clivina westwoodi- species group

1 Pronotum sturdy, disk distinctly convex, outline conspicuously square, basal part of lateral margin slanted flatly with angle less than or about 20°, base slightly produced posteriorly ( Figs 2 , 38–43 ); labrum 7-setose.............................. 2

- Pronotum with outline peltate, cordate, or subcordate, basal part of lateral margin slanted steeply with angle of about 30° or more, base distinctly produced or elongated posteriorly (examples in Figs 1 , 48 , 56 ); labrum 5-, 6- or 7-setose........... 7

2 Large sized species (more than 7.2 mm); pronotum square, slightly wider than long, disk somewhat flattened, lateral margin nearly parallel, distinctly crenate, posterior angles sharp, conspicuously protruding laterally; head, pronotum and elytron glossy; third interval of elytron with 4 setigerous punctures, first stria with small apical setigerous puncture; body length 7.24–8.96 mm; Borneo : Sabah, Brunei....................................................... monticola Andrewes, 1931

- Small to medium sized species (4.85–7.19 mm); pronotum wider than long, disk moderately or distinctly convex, lateral margin convex............................................................................................. 3

3 Pronotum with disk somewhat flattened at middle, with coreaceate-like distinctly engraved wrinkles, lateral margin distinctly crenate; piceous species; pronotum with basal part of lateral margin and basal carina forming a slight but distinct angle; body length 6.05–6.14 mm; S-Vietnam........................................................ collaricrenata sp. nov.

- Pronotum with disk distinctly convex, nearly free of rough wrinkles, lateral margin nearly smooth; testaceous species; pronotum with basal part of lateral margin and basal carina nearly continuously rounded off or with tooth at posterior angle distinctly projecting laterally.................................................................................... 4

4 Pronotum distinctly wider than long, with anterior two thirds distinctly narrowed, with teeth of posterior angles always distinct, more or less projecting laterally, basal transverse channel slightly wider.......................................... 5

- Pronotum slightly wider than long, quadratic, with teeth of posterior angles nearly rounded off, basal transverse channel narrow.............................................................................................. 6

5 Pronotum with anterior angles narrowed in anterior third, teeth of posterior angles sharp and projecting laterally; elytron with interval 8 and half of 7 with isodiametric reticulation, with end of apical tip reticulated, with bright spot at apical tip; interval 8 at apex with short carina, interval 7 convex at apex; body length 6.31–7.19 mm; Borneo : Sabah, Brunei, Sarawak......................................................................................... convexaquadrata sp. nov.

- Pronotum with anterior two thirds narrowed, trapezoid, teeth of posterior angles distinct but not sharp; elytron with interval 6-8 with isodiametric reticulation, with apical third reticulated, with lateral margin at tip slightly brightened; interval 8 with narrow sharp carina, interval 7 apically with step–like ridge; body length 6.21 mm; Philippines: Mindanao.... trapezicollaris sp. nov.

6 Elytron with interval 6-8 with isodiametric reticulation, with apical quarter reticulated, base of interval 3 with small tubercle; eyes semicircular protruding laterally, genae indistinct; body length 6.07–6.94 mm; Nepal to SW China......................................................................................... quadrataconvexa Balkenohl, 2023

- Elytron with interval 7 and 8 with isodiametric reticulation, with apical tip reticulated, base of interval 3 with conspicuous tubercle; eyes convex, moderately protruding laterally, genae distinct; body length 4.85–4.96 mm; Nepal.......... tuberculifer sp. nov.

7 Pronotum with outline peltate, basal part of lateral margin always convex, slanted steeply with angle of about 35°; with tooth at posterior angle inconspicuous......................................................................... 8

- Pronotum with outline cordate or subcordate, basal part of lateral margin straight or slightly convex, tooth at posterior angle usually distinct...................................................................................... 13

8 Elytron with 5 setigerous punctures on interval 3; pronotum longer than wide; body length 6.08 mm; Taiwan......................................................................................... depressa depressa Kult, 1951 o Specimens from the Philippine Islands, Samar: Elytron with intervals 6-8 with isodiametric reticulation; eye posteriorly enclosed by gena; notches at clypeus more distinct; body length 5.48 mm; Philippines: Samar ................................................................................................. depressa samarensis ssp. nov.

- Elytron with 4 setigerous punctures on interval 3; pronotum as long as wide....................................... 9

9 Labrum 7-setose, bilobed, deeply excised at middle; protibia with lateral spines conspicuously acute; body length 5.77 mm; Malaysia: Sabah .......................................................................... bilobata sp. nov.

- Labrum 6-setose, convex, straight, pentagonal, or slightly excised; protibia with lateral spines developed not extraordinarily acute.............................................................................................. 10

10 Elytron with interval 7 and 8 isodiametrically reticulated; head with frons slightly hollowed out, clypeo-frontal furrow nearly invisible; body length 4.6-5.19 mm; Philippines: Leyte ...................................... caputdepressa sp. nov.

- Elytron with all intervals isodiametrically reticulated or reticulation distinct on lateral intervals and fading up to interval 1; head with frons convex or flattened, clypeo-frontal furrow flat but traceable.......................................... 11

11 Elytron with all intervals distinctly isodiametrically reticulated; labrum pentagonal with tip at middle, 6-setose; pronotum with posterior angle distinctly rounded off with tooth indistinct; neck constriction closed with step and band of punctures; body length 5.31–5.47 mm; Indonesia: Papua .................................................... opacielytra sp. nov.

- Elytron with lateral intervals distinctly and regularly isodiametrically reticulated, reticulation fading up to interval 1; labrum with different form, 5-, 6- or 7-setose; pronotum with posterior angle less distinctly rounded off with tooth well visible; neck constriction closed or open............................................................................. 12

12 Small and slender species; head with neck constriction open at middle; labrum excised, angled laterally; pronotum on flattened disk covered with larger group of moderately sized punctures and wrinkles; body length 4.35 mm; Philippines: Palawan ............................................................................................. bauri sp. nov.

- Medium sized species; head with neck constriction closed, with distinct step also at middle; labrum straight, rounded laterally; pronotum on slightly convex disk covered with densely scattered fine punctures, wrinkled laterally; body length 5.72–6.04 mm; Philippines: Mindoro............................................................... dilutureticulata sp. nov.

13 Pronotum with outline distinctly cordate with anterior angles distinctly rounded off, basal part of lateral margin straight, lateral margin with concave gap before posterior tooth, posterior lateral setigerous puncture conspicuously removed from margin towards middle, flange conspicuously elongated, with widened basal transverse channel and flat basal carina ( Figs 51–54 )......... 14

- Pronotum with outline subcordate or subcordate to square with anterior angles slightly rounded or angled, basal part of lateral margin usually convex, in some species bisinuate, lateral margin without concave gap before posterior tooth, posterior lateral setigerous puncture more or less located at margin, flange regularly developed, in some cases with moderately widened basal transverse channel, with acute or convex basal transverse carina (examples in Figs 56, 63 , 69 )....................... 17

14 Labrum 5-setose; pronotum with basal lateral margin slightly convex, proepisternum slightly visible basally in dorsal view; elytron with short carina posterior humerus; body length 4.78–5.15 mm; Philippines: Pulobatu and Mindanao.............................................................................................. subdepressa Kult, 1951

- Labrum 7-setose; pronotum with basal lateral margin straight, proepisternum not visible in dorsal view; elytron without carina posterior humerus.................................................................................... 15

15 Pronotum slender, longer than wide, flange with basal transverse carina inconspicuous; eye flattened in dorsal view and oriented ventrally, enclosed posteriorly to one third by gena; body length 4.74–5.75 mm; Brunei.............. gracilicollis sp. nov.

- Pronotum as long as wide or wider than long, flange with basal transverse carina indicated by smooth convexity; eye regularly developed or subglobose, slightly enclosed by gena......................................................... 16

16 Pronotum distinctly wider than long, covered nearly completely with coreaceate-like structure and medium sized slightly elongated punctures, somewhat opaque, tooth at posterior angle acute, pointing laterally; elytron with interval 8 completely reticulated; body length 5.23–6.02 mm; Borneo : Sarawak, Sabah ............................. depressiforma sp. nov.

- Pronotum as long as wide, covered basally with some coreaceate-like structure and in basal half with circular medium sized punctures, glossy, tooth at posterior angle somewhat rounded, pointing anteriorly; elytron with interval 8 reticulated at lateral marginal channel with rest of interval glossy; body length 4.73– 4.49 mm; Borneo : Sarawak............... geiseri sp. nov.

17 Clypeus, frons, supraantennal plates, neck, pronotum, and intervals of elytron with scattered fine punctures, labrum bilobed; elytron with intervals 6-8 isodiametrically reticulated; body length 5.35 mm; Philippines: Samar ... passimpunctata sp. nov.

- Dorsal surface with all body parts not covered with fine scattered punctures; labrum straight or excised; elytron with inner intervals either smooth and glossy or with different pattern of reticulation........................................ 18

18 Elytron with all intervals isodiametrically reticulated, semi-opaque; and flange of pronotum with basal transverse channel widened, labrum 7-setose.............................................................................. 19

- Elytron with at least intervals 1–4 on disk smooth, glossy; flange of pronotum regularly developed (but see C. vigil with widened basal transverse channel), labrum 6- or 7-setose..................................................... 21

19 Pronotum with lateral margin distinctly crenulate, lateral channel distinctly widened and engraved, median line widened, basal lateral margin straight; disk with pattern of punctures resembling to an indistinct letter Y; body length 5.92–6.16 mm; Borneo : Sarawak.......................................................................... margineserrata sp. nov.

- Pronotum with lateral margin smooth, lateral channel regularly developed, median line narrow, disk with few punctures and slight line-like impression basally (microscopic punctures not considered)....................................... 20

20 Medium sized species; head with frons and neck forms continuous surface, without neck constriction, without clypeo-frontal furrow; elytron with base retracted; clypeus straight, without laterally projecting teeth; body length 5.86–5.91 mm; Indonesia: Irian Jaya ............................................................................ hartmanni sp. nov.

- Slender species; head with flat clypeo-frontal furrow, with neck constriction formed by row of punctures and distinct step; elytron with base more straight; clypeus slightly excised with laterally slightly projecting teeth; body length 4.91–5.05 mm; Borneo : Sabah, Brunei................................................................. spatulifigura sp. nov.

21 Elytron with isodiametric reticulation on interval 8 and 7 or 8, 7, and 6 with lateral strip............................ 24

- Elytron with interval 1–4 or 1–5 smooth at middle on disk, other intervals including parts of base and apex reticulated... 22

22 Elytron with intervals 1–4 smooth at middle on disk, interval 7 with step-like carina at apex; head glossy, with micro-punctures.......................................................................................... 23

- Elytron with intervals 1–5 smooth at middle on disk, interval 7 completely flat at apex; head completely covered with isodiametric reticulation; notches at clypeus indistinct; labrum 7-setose, slightly excised; body length 3.84–4.54 mm; Borneo : Sabah ............................................................................... caputopaca sp. nov.

23 Labrum 7-setose, excised at middle; head 1/7 narrower than pronotum; clypeus and frons with continuous surface, without clypeo-frontal furrow; clypeus with notches distinct; pronotum with lateral margin parallel; body length 5.17–6.38 mm; Borneo : Sabah, Sarawak, Brunei............................................................. circumreticulata sp. nov.

- Labrum 6-setose, straight; head 1/4 narrower than pronotum; with wide distinct clypeo-frontal furrow, clypeus with blunt carina; clypeus with notches small; pronotum with lateral margin converging anteriorly; body length 5.21–5.39 mm; Sulawesi ................................................................................... circumopaca sp. nov.

24 Elytron with interval 8, 7, and 6 with lateral strip of reticulation............................................... 25

- Elytron with interval 7 and 8 completely reticulated or interval 8 completely and interval 7 with lateral strip reticulated... 27

25 Pronotum with basal transverse channel distinctly widened, lateral margin distinctly convex, disk convex; head with neck constriction open at middle; elytron with interval 5 at humerus with short sharp carina; labrum 7-setose; body length 5.83–6.46 mm, Indonesia: Irian Jaya .............................................................. vigil Darlington, 1962

- Pronotum with basal transverse channel narrow, lateral margin slightly to moderately convex, disk more or less convex; head with neck constriction open or closed; elytron with interval 5 at humerus connected to lateral margin, labrum 6- or 7-setose.. .................................................................................................. 26

26 Labrum 7-setose; clypeus slightly concave in between lateral teeth; neck constriction closed by punctures; pronotum with lateral margin more or less parallel but still indistinctly convex; elytron with interval 8, 7, and wide stripe of interval 6 isodiametrically reticulated, apically reticulated by a quarter; eyes slightly less prominent protruding as in C. wallacei ; female epipleurite with the 2 setiferous punctures close together; body length 5.72 mm; New Guinea, Japan............ westwoodi Putzeys, 1867 ,

- Labrum 6-setose; clypeus straight in between lateral teeth; neck constriction open at middle; pronotum with lateral margin more or less convex; elytron with interval 8, 7, and indistinct stripe of interval 6 isodiametrically reticulated, apically reticulated by a third; eyes more prominent protruding as in C. westwoodi ; female epipleurite with the 2 setiferous punctures widely distant; body length 5.39–5.74 mm; Sulawesi............................................ wallacei wallacei Putzeys, 1867 o Specimens from the Philippine Islands: piceous; head with neck more closed; pronotum attenuated in anterior half; male aedeagus more prolonged and narrowed anterior oroficium, not distorted at apex; body length 5.29–6.09 mm; Philippines: Mindanao, Luzon, Marinduque, Leyte ........................................ wallacei philippinensis Kult, 1951 o Specimens from the Solomon Islands: fuscous; legs testaceous; eye more globose; clypeo-frontal furrow slight; pronotum with slight depression at level of posterior setigerous puncture; male aedeagus narrowed towards apex, distorted and fractuate at apex; body length 4.71–5.21 mm; Solomon Islands..................... wallacei solomonensis ssp. nov.

27 Elytron with interval 7 and 8 completely reticulated; carina at apex of interval 8 narrowed and elongated; labrum 7-setose; lateral margin of pronotum nearly parallel and straight; clypeus with distinctly reflexed margin, body length 6.91 mm; Philippines: Luzon.............................................................................. berndjaegeri sp. nov.

- Elytron with interval 8 completely and interval 7 with lateral strip reticulated; carina at apex of interval 8 of moderate length; labrum 6- or 7-setose; lateral margin of pronotum more or less convex; clypeus with moderately or finely reflexed margin... 28

28 Melaenous, small species; pronotum with nearly straight lateral margin, covered densely with medium sized punctures, more dense at middle; labrum convex, 7-setose; elytron with interval 8 apically with step-like carina, interval 7 flattened at apex; body length 4.03–4.55 mm; Borneo : Sarawak, Brunei....................................... punctaticollis sp. nov.

- Fuscous, medium sized species; pronotum with moderately convex lateral margin, with puncture pattern bilaterally resembling often to the letter Y; labrum obtuse angled, straight, or excised, 6- or 7-setose; elytron with interval 8 apically carinate, interval 7 with step-like carina................................................................................ 29

29 Head with distinct closed step at neck constriction and row of confluent punctures; labrum 6-setose, obtuse angled; clypeus with notches indistinct, lateral teeth hardly protruding; pronotum in anterior two thirds trapezoid, distinctly wider than long; hind body subcylindrical, intervals moderately convex; aedeagus developed as elongate spatula; body length 5.67 mm; Philippines: Manila .................................................................................... paris sp. nov.

- Head without or with small step at neck constriction, constriction open or closed; labrum 7-setose, straight or excised; clypeus with notches distinct or indistinct, lateral teeth distinctly or slightly protruding; pronotum more or less attenuating in anterior half, slightly wider than long; hind body regularly developed, intervals slightly convex; aedeagus widened apically...... 30

30 Pronotum with outline square, lateral margin slightly crenulated; elytron with interval 7 covered by half of width with isodiametric reticulation (beside the completely covered interval 8); male aedeagus with apical lamella in dorsal view slightly bent at apex, in lateral view duck-bill like; body length 4.07–5.32 mm; Sumatra; Nias; Perak; Malaysia, Pahang ........................................................................................... integra Andrewes, 1929

- Pronotum with outline trapezoid or with angles distinctly rounded, lateral margin smooth; elytron with interval 7 covered laterally with a small just traceable stripe of isodiametric reticulation (beside the completely covered interval 8)......... 31

31 Pronotum with outline attenuating in anterior half; neck constriction developed as slight step with punctures posteriorly; eye prominent; elytron with interval 8 convex; male aedeagus in dorsal view with apical lamella straight to apex, in lateral view spoon-like widened; body length 4.58–5.42 mm; Sumatra, S Malaysia............................... similata sp. nov.

- Pronotum with outline subsquare with all angles rounded (resembling to sub-globosity in dorsal view); neck constriction developed as step, flat and indistinct at middle; eye flattened; elytron with interval 8 flattened; male aedeagus in dorsal view conspicuously arcuate in apical fifth including lamella, lamella in lateral view duck-bill shaped; body length 4.44 mm; N Sumatra ................................................................................ neglecta sp. nov.

Species taxonomic account

Figures

FIGURES 105–109. Clivina species, female external reproductive tract with gonocoxites and epipleurite. (105) C. monticola Andrewes, 1931, lectotype, Borneo, Kina Balu. (106) C. monticola, spec. from Brunei, Tembourong. (107) C. convexaquadrata sp. nov., paratype from Borneo, Sabah, Danum Valley. (108) C. quadrataconvexa Balkenohl, 2023, spec. from N-India, Rishikesh. (109) C. tuberculifer sp. nov., paratype from Nepal, Chitwan NP.

FIGURES 110–128. Clivina species, female external reproductive tract with gonocoxites and epipleurite.(110) C. caputdepressa sp.nov., paratype.(111) C. opacielytra sp. nov., paratype from Papua, Palmer River.(112) C. gracilicollis sp. nov., paratype.(113) C. depressiforma sp. nov., paratype from Sabah, Gunung Alab peak. (114) C. geiseri sp. nov., paratype. (115) C. margineserrata sp. nov., paratype. (116) C. hartmanni sp. nov., paratype. (117) C. spatulifigura sp. nov., paratype. (118) C. caputopaca sp. nov., paratype from Sabah, Danum Valley. (119) C. circumreticulata sp. nov., paratype from Brunnei, Temburong. (120) C. circumopaca sp. nov., paratype. (121) C. vigil Darlington, 1962, spec. from Indonesia, Irian Jaya, Genyem. (122) C. westwoodi Putzeys, 1867, lectotype. (123) C. wallacei wallacei Putzeys, 1867, spec. from Dorey. (124) C. wallacei philippinensis Kult, 1951, spec. from Manila. (125) C. wallacei solomonensis ssp. nov., paratype. (126) C. punctaticollis sp. nov., paratype from Brunei, Temburong. (127) C. integra Andrewes, 1929, spec. from Nias Island. (128) C. similata sp. nov., paratype.

FIGURES1–2.Clivinaspecies, pronotum,dorsalview,semi-schematicview.Ontheright,some ofthefeatures anddiscriminating characters used in the descriptions and the identification key are highlighted. On the left, the measurement of the angle at the base is figured. (1) C. depressiforma sp. nov. with cordate shape. (2) C. quadrataconvexa Balkenohl, 2023, with square shape.

FIGURES 38–52. Clivina species, pronotum, dorsal view. (38) C. monticola Andrewes, 1931, lectotype. (39) C. collaricrenata sp.nov., holotype.(40) C.convexaquadrata sp.nov., holotype. (41) C.trapezicollaris sp.nov., holotype. (42) C.quadrataconvexa Balkenohl, 2023, spec. from Myanmar, Putao. (43) C. tuberculifer sp. nov., holotype. (44) C. depressa depressa Kult, 1951, holotype. (45) C. depressa samarensis ssp. nov., holotype. (46) C. bilobata sp. nov., holotype. (47) C. caputdepressa sp. nov., paratype. (48) C. opacielytra sp. nov., paratype. (49) C. bauri sp. nov., holotype. (50) C. dilutureticulata sp. nov., holotype. (51) C. subdepressa Kult, 1951, lectotype. (52) C. gracilicollis sp. nov., holotype.

FIGURES 53–67. Clivina species, pronotum, dorsal view. (53) C. depressiforma sp. nov., paratype. (54) C. geiseri sp. nov., holotype. (55) C. passimpunctata sp. nov., holotype. (56) C. margineserrata sp. nov., holotype. (57) C. hartmanni sp. nov., holotype. (58) C. spatulifigura sp. nov., holotype. (59) C. caputopaca sp. nov., holotype. (60) C. circumreticulata sp. nov., holotype. (61) C. circumopaca sp. nov., holotype. (62) C. berndjaegeri sp. nov., holotype. (63) C. vigil Darlington, 1962, holotype. (64) C. westwoodi Putzeys, 1867, lectotype. (65) C. wallacei wallacei Putzeys, 1867, lectotype. (66) C. wallacei philippinensis Kult, 1951, holotype. (67) C. wallacei solomonensis ssp. nov., holotype.

FIGURES 68–72. Clivina species, pronotum, dorsal view. (68) C. punctaticollis sp. nov., holotype. (69) C. paris sp. nov., holotype. (70) C. integra Andrewes, 1929, spec. from Sumatra, Ketambe. (71) C. similata sp. nov., paratype. (72) C. neglecta sp. nov., holotype.