CREOBIINA JEANNEL, 1941

ROIG-JUÑENT, SERGIO, 2000, The Subtribes And Genera Of The Tribe Broscini (Coleoptera: Carabidae): Cladistic Analysis, Taxonomic Treatment, And Biogeographical Considerations, Bulletin of the American Museum of Natural History 2000 (1), pp. 1-90 : 52-53

CREOBIINA JEANNEL, 1941

Cnemacanthini Lacordaire, 1854 : ( Cnemacanthus Gray, 1832 = Promecoderus Dejean, 1829 , non Cnemacanthus sensu Brullé, 1834 ).

Creobitae Jeannel, 1941: 287 (as a subfamily).

Creobiina : Ball, 1956: 45. Townsend, 1971: 180. Roig-Juñent, 1995a: 52.

TYPE GENUS: Creobius Guérin-Ménéville, 1838 .

REMARKS: The suprageneric name Cnemacanthini has strict priority over the name Creobiina . Nevertheless, use of Cnemacanthini would create confusion, since it was used as the tribal name for the genus Cnemalobus , a harpaline group (Roig-Juñent, 1993).

TAXA INCLUDED: The subtribe Creobiina remains as defined by Ball (1956) (table 3). It represents the largest group of Broscini in number of species (appendix 1). Jeannel (1941) considered Creobiina to include the austral genera with the basal orifice of median lobe open, and the genera Miscodera and Broscodera . Ball (1956) excluded these two genera from Creobiina , including them in Broscina . This is the viewpoint followed in this study.

The subtribe Creobiina comprises 11 genera with 97 species (only two genera are monospecific) (appendix 1).

Creobiina shares with the sister subtribe Barypina losses of sclerites affecting the male genitalia (without internal sclerites) and the female tract (spermatheca without vaginal apophysis). Creobiina shows nine synapomorphies, with only one unique derived feature: enlarged left paramere (532). The other structures suffer reversions within Creobiina or changes to another character state: the shape of mentum tooth (102); male foretarsomeres 1–4 with adhesive vestiture ventrally (441); rami of gonocoxite 9 absent (630). Several features are reversions to the plesiotropic condition: paraglossas short (161), median lobe with basal orifice completely open (540), dorsal surface membranous (550), and with basal expansions (561), or parallelism with other groups, like nematiform setae of subapical setose organ absent (641).

Within this subtribe, there are two clear monophyletic groups, also recognized by Ball (1956). One group includes four Australian genera, and is characterized by nine synapomorphies, five of them unique structures within the tribe: mentum without tooth (100), mesoepimera with carina (291), antecoxal suture of metasternum not visible (321), presence of a large tooth in the internal sac, that is seen in the apical orifice when the internal sac is not everted (621), and accessory gland in the spermatheca (701). Some genera of this group have large species, and some species have adaptations associated with fossorial habits, like fore and middle tibiae with lateral expansions, characters also present in Brullea ( Nothobroscina ).

The other group is a large clade, including the genus Promecoderus with 41 species. This clade is supported by three synapomorphies: glossal sclerite without carina (140), antennal pubescence from half of the third antennomere (220), and presence of four setae on the glossal sclerite (152) as exclusive character within Broscini . The cladograms (figs. 14, 15) show that the relationships among the different genera are not resolved. Results of this study (fig. 15) show that the genus Promecoderus is not monophyletic and the taxonomic revision of this genus is necessary in order to clear up the different groups of species within it.

The two main groups of creobiines inhabit different habitats: one group, Cerotalis and related genera, are big to medium-size species and occur principally in dry habitats. The second group, Promecoderus , Creobius , Cascellius , and related genera, are small to medium-size, bright-green or brown species, and occur in Nothofagus forests.