Western Hemisphere Zuphiini: descriptions of Coarazuphium whiteheadi, new species, and Zuphioides, new genus, and classification of the genera (Coleoptera, Carabidae) Author Ball, George E. Author Shpeley, Danny text ZooKeys 2013 315 17 54 http://dx.doi.org/10.3897/zookeys.315.5293 journal article http://dx.doi.org/10.3897/zookeys.315.5293 1313-2970-315-17 Coarazuphium Gnaspini,Vanin & Godoy, 1998 Parazuphium ; Godoy and Vanin 1990 : 795 (not Jeannel 1942 ). Zuphium ; Mateu 1993 : 486 (not Latreille 1805 ). Coarazuphium Gnaspini, Vanin & Godoy, 1998: 298. Alvares and Ferreira 2002 : 41. Lorenz 2005 : 507. Pellegrini and Ferreira 2011a : 39. Pellegrini and Ferreira 2011b : 47. Type species. Parazuphium tessai Godoy & Vanin, 1990 (designated by Gnaspini et al. 1998 ). Generic name. As explained by Gnaspini et al. (1998 : 299), the name Coarazuphium is a compound Latinized neuter noun derived from "coara" , meaning hole or cave in the Tupi language (Brazilian, native tongue), plus Zuphium . The name refers to the troglophilic or troglobitic habits of the species of this genus, and their basic nominotypical Zuphium -like features. Recognition. With character states of Western Hemisphere Zuphium genus-group, restrictedas follows: size small; pronotum and dorsal surface of elytra moderately densely setose, setae decumbent; body markedly depressed; integument pale; head narrow, oviform, with posteriolateral margins broadly rounded; antennae elongate, antennomere 1 as long or longer than antennomeres 2-4; humeri narrowed, slightly or markedly so; metasternum short, metepisternum quadrate. Brachypterous or apterous. Male genitalia (Figs 10 A- 10C): phallus without dorsal paraostial sclerites. Female genital tract (Fig. 12A, 12B): without secondary spermathecal gland. Description. None required here. See Gnaspini et al. (1998 : 298-299), Pellegrini and Ferreira (2011b) , and description of Coarazuphium whiteheadi below. Geographical distribution. Confined to the Neotropical Region, the seven species of this genus are known only from southeastern Brazil ( Pellegrini and Ferreria 2011b : 57, fig. 10) and southern Mexico (Map 1). This is an example of the "Paleo-American distribution pattern" (see Halffter (1987) and Liebherr (1994 : 845)). Way of life. The previously described species of genus Coarazuphium inhabit caves in one of two types of substrate ( Pellegrini and Ferreira 2011b : 55): limestone, occupied by Coarazuphium bezerra Gnaspini, Vanin & Godoy; Coarazuphium cessaima Gnaspini, Vanin & Godoy; Coarazuphium formoso Pellegrini & Ferreira; Coarazuphium pains Alvares & Ferreira; and Coarazuphium tessai (Godoy & Vanin); and iron ore, occupied by Coarazuphium tapiaguassu Pellegrini & Ferreira. The caves in iron ore are described as "shallow" . Members of those species occupying the caves in limestone substrate are described as freely walking over the soil, and presumably resting exposed, rather than resting (hiding?) under rock cover. In contrast, all of the specimens collected of Coarazuphium tapiaguassu were found under rocks, on the cave floor. In contrast, known members of the Mexican species, Coarazuphium whiteheadi , new species, seem to be surface inhabitants (see below) Parasites. Some specimens of Coarazuphium tapiaguassu were infested by laboulbenelian fungi. The fungi were not identified further. See this topic for Coarazuphium whiteheadi , below. Evolutionary considerations. The ultrastructural features (i.e., principally sensillar) observed with scanning electron microscopy by Pellegrini and Ferreria (2011a , 2011b ) differ only slightly between the two species of Coarazuphium that they studied, and between that genus and Zuphioides (cf. accompanying SEM illustrations of Zuphioides mexicanum ). So, it seems to us unlikely that such features are or will be evolutionarily informative. In contrast, the standard structural troglobitic features of Coarazuphium (lengthening of antennae and legs, depigmentation, micro- or anophthalmy, and reduction of elytral length, elytral humeri, metathorax and metathoracic wings) plus details of male and female genitalia are evolutionarily informative. Based on eye loss and more elongate appendages, Gnaspini et al. (1998 : 303) proposed that Coarazuphium cessaima showed the more modified features, compared to Coarazuphium tessai and Coarazuphium bezerra , the only other species of Coarazuphium known at that time. Alvares and Ferreira (2002 : 43) proposed that, based on the features noted above, their newly described Coarazuphium pains would occupy a position intermediate between Coarazuphium cessaima and Coarazuphium tessai + Coarazuphium bezerra . The external features of Coarazuphium evidently evolved in parallel with, and independent of, four other zuphiine troglobite taxa: the remarkable Spanish Ildobates neboti Espanol , 1966; the Canary Islands Parazuphium feloi Machado (1997: 163); the Australian Nullarbor Speozuphium poulteri Moore (1995: 159) and Speothalpius grayi Moore (1995: 160). Another Canary Islands Parazuphium ( Parazuphium damascenum canariense Machado, 1992: 580) exhibits these same reductive (though less developed) features, but it is evidently conspecific with the continental Palaearctic Parazuphium damascenum damascenum (Fairmaire, 1897). For a more detailed discussion of the matter see Machado (1992 : 581-582). Gnaspini et al. (1998 : 308-309) proposed for the Brazilian species of Coarazuphium that "..... these highly derived troglobitic features are due to a long-term isolation inside the subterranean environments, which took place under the drier climate to which the region was in the past and is still submitted in the present. It is largely accepted in the literature that cave arthropods are related to litter epigean and/or endogean ancestors, which already inhabited humid habitats. Therefore,... ... the ancestral species [of Coarazuphium ] should have been epigean and lived in forested (or at least humid) areas, and occurred at least in part of the region where the genus occurs nowadays. From [such ancestral stock] several lineages invaded the caves from the northern Bambui Speleological Province, where they became isolated with the progressive shrinkage of humid environments. Thence, the origin of the genus takes back to a time when the area was not drying yet, which is probably the Tertiary". Discovery of the Mexican species adds important details to the story of evolution of Coarazuphium . First, the marked geographical range extension (from southeastern South America to the southern part of the North American continent) shows that this genus was not confined to eastern Brazil. Further, the distribution pattern lends support to the hypothesis of Gnaspini et. al. (see above) that Coarazuphium originated in early Tertiary time (i.e., before the beginning of the drying trend that extended through much of the Cenozoic era). Second, the extra-speleal humid forest existence of Coarazuphium whiteheadi suggests that the basic troglobitic features of Coarazuphium (microphthalmy, brachyptery, depigmentation, etc.) evolved in surface habitats, though probably in forested deep leaf litter locations, and were in effect preadaptations for cave life. An additional observation relating to evolutionary history of the Zuphium genus groupis that shared features of Zuphioides and Coarazuphium indicate that these two genera may be adelphotaxa, with Zuphioides retaining mostly ancestral features including life in lowland hygrophilous or mesophilous situations. Key to species of genus Coarazuphium Gnaspini, Vanin & Godoy

Gnaspini et al. 1998 Gnaspini et al. 1998	Coarazuphium cessaima
Coarazuphium tapiaguassu
Alvares and Ferreira 2002 Alvares and Ferreira 2002	Coarazuphium pains
Fig. 5AFig. 9BFig. 10C	Coarazuphium whiteheadi
Pellegrini and Ferreira 2011a	Coarazuphium formoso
Gnaspini et al. 1998 Gnaspini et al. 1998	Coarazuphium bezerra
Godoy and Vanin 1990 Godoy and Vanin 1990	Coarazuphium tessai