Clivinina Rafinesque, 1815

Balkenohl, Michael, 2022, On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae), Zootaxa 5190 (1), pp. 99-126 : 105

Clivinina Rafinesque, 1815

The subtribe includes presently thirty-one genera worldwide. The many species listed for the twenty Oriental and African genera exhibit four very different basic morphologies (A, B, C, D). Moreover, three variations of the falciform shape A are represented (A1, A4, A5) ( Tab. 1 ).

The falciform Shape A1 ( Fig. 1 ) is found in approximately 70% of the genera and species. In this group, a morphology is found which seems to represent a close shape to the plesiomorphic dimeric form constituting the ‘nebridian configuration’ (according to Deuve 1993). Depending on the genus, the gonocoxite-1 and gonocoxite-2 are more or less fused with a more or less wide membranous area in between.

Significant differences between the subgenera Clivina s. str. and Leucocara Bousquet, 2009 are not observed.

The most notable variations are as follows: The subgenus Dacca shows more elongated, slender and sharp coxites similar to members of the C. helferi -species group. In the C. mekongensis species-group, gonocoxite-1 and gonocoxite-2 are not as intensely fused as in all other species and they are still a bit movable ( Fig. 12 ). That shape is considered as the most plesiomorphic form. This find also justifies highlighting this group by assigning it herewith as C. mekongensis -species group.

Rugiluclivina seems to be somewhat more specialized ( Fig. 21 ). The coxites are slightly curved, gonocoxite-2 is distinctly ensiform widened, and the gonocoxites-1 are more strongly medially connected to each other ( Fig. 22 ). It somewhat resembles the shape found in Forcipatorina.

The following groups have been revised recently, all including figures of the coxites: Ancus ( Balkenohl 2016) , Clivina castanea -species group ( Balkenohl 2021a), Clypeospinus ( Balkenohl 2021b), Leucocara with the C. semicarinata -species group ( Dostal & Bulirsch 2016), Rugiluclivina ( Balkenohl 1996, 1999 a, 2015, 2018a), Dacca ( Balkenohl 2020) . For Leleuporella , figures of coxites have been provided for two species ( Fedorenko 2012, Bulirsch & Magrini 2019).

Shape A1 seems to be distributed over very wide geographic ranges, probably worldwide.

The variations with one or two short macrotrichia (shapes A4 and A5) are considered to represent derived forms where the large seta is replaced by one or two short macrotrichia. The groups with shape A4 with slender coxites have been revised recently with figures of the coxites provided: Orictites and its subgenus Semictites ( Balkenohl 2017a, b), and Paracoryza ( Balkenohl 2000, 2005). The groups with shape A5 with monomeric and conspicuously curved and short coxites were revised as well and comprise Syleter ( Balkenohl 2021b) , Parasyleter ( Balkenohl 2022) , Trilophus ( Balkenohl 1999b) , and Trilophidius ( Balkenohl 2001) .

Another basic morphology is represented by the small and unusual monomeric foliform gonocoxite and semirectangular glabrous epipleurite without central articulation (shape B, Fig. 2 ). On a group level, Thliboclivina and Physoclivina have been revised and keyed out recently ( Balkenohl 2001, 2018c, Dostal 2015). Eoclivina is characterised and keyed out by Kult (1959) but without considering characters of the genitalia. The Oriental C. attenuata -species group is supposed to belong to Eoclivina as indicated in Lorenz (2005, 2022). However, this assumption is investigated here the first time. The C. attenuata -species group has not been revised in more than ninety years, with the last available treatment on a species level by Andrewes (1929).

It has been found that, apart from the conspicuous differences of characters in the female genitalia, all members of these four species groups share several unusual external characters and combinations of characters not present in the other Clivinini . Based on the striking differences, the sixteen species have been integrated into a new subtribe characterised in this contribution (see below).

The third basic pattern shows a monomeric hyaline musaceous gonocoxite and a glabrous rectangular epipleurite without central articulation (shape C, Fig. 47 ), also representing a different line. The genus Lophocoryza has been revised recently ( Balkenohl 2018b). Halocoryza was treated by Whitehead (1966, 1972) and revisited by Erwin (2011). Coryza is an unrevised genus with partly unclear synonymies, undescribed species, and partly unclear distribution.

The fourth basic pattern shows monomeric hyaline elongated foliform gonocoxites and semi-square glabrous epipleurites with complete articulation (shape D, Fig. 55 ). The shape is found in Afrosyleter , a genus treated by Bulirsch & Magrini (2018) with a figure of the gonocoxites of one species. This shape seems to represent also a different line. However, the data published and the examples at hand are poor and need further insight.

Figures

FIGURE 1. Clivina (subgenus Leucocara Bousquet, 2009) sacra Putzeys, 1875. External parts of the female reproductive tract in ventro-lateral delaminated view, highlighting some of the terms used in the descriptions.

FIGURE 2. Sulciclivina attenuata (Herbst, 1806). External parts of the female reproductive tract in ventro-lateral delaminated view, highlighting some of the terms used in the descriptions.

FIGURE 55. Afrosyleter ituricus Basilewsky, 1959, specimen from Parc du Banco, Ivory Coast. Gonocoxites and epipleurite, ventro-lateral view. Shape D.

FIGURES 7-25. Clivinini species, gonocoxites and epipleurites, ventro-lateral view. Shape A1. 7. Clivina (Clivina) fossor sachalinica Nakane, 1952, specimen from Sakhalin. 8. C. (C.) agona Putzeys, 1866, specimen from Banteay, Cambodia. 9. C. (C.) molucca Balkenohl, 2021, PT from Seram, Maluku. 10. C. (C.) helferi Putzeys, 1866, type spec. from Calcutta. 11. C. (C.) angularis Putzeys, 1867, type spec. from Calcutta. 12. C. (C.) mekongensis Lesne, 1896, specimen from Luang Prabang, Laos. 13. C. (L.) traquebarica Bonelli, 1813, type spec. from Indo China. 14. C. (L.) tutancamon Schatzmayr, 1936, specimen from Barrage, Egypt. 15. C. (Dacca) forcipata (Putzeys, 1861), specimen from Calcutta. 16. Ancus carniceps Andrewes, 1936, HT from Balor, Java. 17. Ancus excavaticeps Putzeys, 1867, specimen from Prek Toal, Cambodia. 18. Basilewskyana villiersi Basilewsky, 1948, specimen from Touba, Ivory Coast. 19. Clypeospinus validus (Andrewes, 1936), HT from Battaramulla, Sri Lanka. 20. Leleuporella mandibularis (Burgeon, 1935), specimen from Thuapa, Congo. 21. Rugiluclivina puncticollis Balkenohl, 1996, PT from Hanoi, Vietnam. 22. R. puncticollis, unmounted, ventral view. 23. Schizogenius freyi Baehr, 1983, HT from Vilu Levu, Fiji Islands. 24. S. sulcifrons Putzeys, 1846, specimen from Abercorn, USA. 25. S. neovalidus Whitehead, 1972, specimes from Aravaipa, Arizona.

FIGURES 47-54. Clivinini species, gonocoxites and epipleurites, ventro-lateral view. Shape C. 47, 48. Coryza maculata Nietner, 1856, specimens from Nepal in mounted and unmounted view. 49. Halocoryza maindoniAlluaud, 1919, specimen from Aden, Yemen in unmounted view. 50. H. arenaria (Darlington, 1939), specimen from Barbados. 51. Lophocoryza sechellensis Basilewsky, 1973, PT from Ile Praslin, Seychelles, unmounted view. 52, 53. L. araticeps omanica Balkenohl, 2018, HT in enlarged detailed and delaminated view. 54. L. sciakyi Balkenohl, 2018, PT from Saint-Augustin, Madagascar.