Cistugo Seabrae Thomas, 1912 and C

An updated distribution map and additional museum voucher specimens of the genus Cistugo (Chiroptera: Cistugonidae), with a review of the distinction between Cistugo seabrae Thomas, 1912 and C. lesueuri Roberts, 1919, and aspects concerning their allopatric distribution Ernest C. J. Seamark1*, Teresa C. Kearney2 and Johan Watson3 1AfricanBats.org, 357 Botha Ave, Kloofsig, 0157 South Africa 2Vertebrate Department, Ditsong National Museum of Natural History (formerly Transvaal Museum), P.O. Box 413, Pretoria, 0001 South Africa 3Free State Department of Economic Development, Tourism and Environmental Affairs, Biodiversity Research, Private Bag X20801, Bloemfontein, 9300 South Africa SEAMARK, E. C. J., KEARNEY, T. C. and WATSON J., 2012. An updated distribution map and additional museum voucher specimens of the genus Cistugo (Chiroptera: Cistugonidae), with a review of the distinction between Cistugo seabrae Thomas, 1912 and C. lesueuri Roberts, 1919, and aspects concerning their allopatric distribution. Annals of the Ditsong National Museum of Natural History 2: 111–119. While recent work focused on resolution at the family level, and led to the suggestion of a new family for the genus Cistugo, the validity of Cistugo lesueuri still remains unresolved.We provide a review of the classification of C.lesueuri and C. seabrae, and an updated distribution map for the genus including new localities for each species. Two measurements initially provided to distinguish the species are revisited, and suggest the distributions of C. lesueuri and C. seabrae may not be allopatric, as had previously been assumed. Keywords: Cistugo lesueuri, Cistugo seabrae, Species Classification, Distribution. INTRODUCTION basis of diploid and fundamental chromosome num- The results of recent molecular work by Lack et al. ber (Rautenbach et al., 1993) and mitochondrial (2010) led them to propose that Cistugo Thomas, cytochrome-b data (Bickham et al., 2004; Stadel- 1912 should be placed in its own family, the mann et al., 2004, Rautian et al., 2006), Cistugo was Cistugidae. The African Chiroptera Report (ACR) recognized as a valid genus in the species lists of (2011), however, points out the correct form of the Bronner et al. (2003), Simmons (2005), Monadjem new family name is Cistugonidae. Combining mito- et al. (2010), and the ACR (2011). chondrial and nuclear DNA information with more The recorded differences of Cistugo relative to complete taxonomic sampling enabled Lack et al. other vespertilionids are that they have an ancestral (2010) to confirm earlier karyology (Rautenbach diploid number (2n = 50) with an all-acrocentric et al., 1993) and molecular results (Bickham et al., autosomal complement, and a fundamental chro- 2004; Stadelmann et al., 2004; Eick et al., 2005) that mosome number of FN = 48 (Rautenbach et al., had indicated the genus Cistugo occurs basally to 1993). Based on mitochondrial cytochrome-b data, all other vespertilionids and is separated from other Stadelmann et al. (2004: 185) found Cistugo geneti- vespertilionid genera by a primitive karyotype and a cally distinct from other Myotis at >20% uncor- high genetic distance. Historically, Cistugo has rected genetic distance, while Bickham et al. (2004: been synonymized with Myotis Kaup, 1829, albeit in 335) found an average p-distance between Cistugo some instances as a separate subgenus to Myotis, and Myotis of 21%. Using nuclear genes Eick et al. in the following species checklists: Ellerman and (2005) identified two distinctions between Cistugo Morrison-Scott (1951), Ellerman et al. (1953), and all the other vespertilionids they examined; Ellerman (1954), Ellerman and Morrison-Scott Cistugo has a unique insertion in PRKC1 and lacks a (1966), Hayman and Hill (1971), Honacki et al. unique deletion in the SPTBN intron, neither of (1978), Honacki et al. (1982), Meester et al. (1986), which were found in the other vespertilionids they Corbet and Hill (1991), Koopman (1993), examined. Lack et al. (2010) also found a single Herselman and Norton (1985), Koopman (1994), 18-bp indel in exon 6 of the nuclear exon, DMP1, and Taylor (2000). However, more recently, on the gene in Cistugo that was not found in any other *Author for correspondence. E-mail: [email protected] Vespertilionidae sequenced. 112 ANNALS OF THE DITSONG NATIONAL MUSEUM OF NATURAL HISTORY, VOLUME 2, 2012 Observed morphological differences of Cistugo 1985; Lynch, 1994; Watson, 1998; Taylor, 2000; relative to Myotis include their usually having glands Seamark and Kearney, 2006; Monadjem et al., present on the wings (Meester et al., 1986; 2010) the known distributions of the species have Simmons, 2005). Roberts (1919: 112), however, did expanded, and ranges of measurements have been not observe glands on the holotype of C. lesueuri, introduced to the key characters to encompass the and since the skin of the holotype is dried with the variation introduced by additional specimens. Thus wings folded against the body it has subsequently Hayman and Hill (1971) identified C. seabrae as not been possible to re-assess this character from being distributed from Angola, Namibia and north- the specimen. Shortridge (1942: 79) also mentioned western Cape Province, and having a smaller fore- in the account of individuals of C. lesueuri caught at arm (32.5–32.9 mm); while C. lesueuri was recorded the Hex River Valley Estate near Citrusdal that ‘In dry as being distributed in the southwestern Cape skins these glands become absorbed and lost to Province (also from Ceres), with a larger forearm of sight’. In Cistugo the anterior pair of premolars is 34.5 mm. The aspects presented in the classification smaller than in other Myotis, and the large upper of Meester et al. (1986) were the same as those premolar has a better developed antero-internal in Hayman and Hill (1971), with the distribution for cusp than in other Myotis (Thomas, 1912). C. seabrae given as the northwestern Cape Prov- Stadelmann et al. (2004: 178), however, incorrectly ince, western Namibia and southwestern Angola, reported this character as relating to the lower fourth while that of C. lesueuri was recorded as the south- premolar, rather than the upper fourth premolar. western Cape Province, from the Franschhoek area Two species are described within the genus north to Citrusdal, and northeast to Beaufort West Cistugo: C. seabrae Thomas, 1912 and C. lesueuri and Mazelsfontein. Taylor (2000) repeated the Roberts, 1919. When Roberts described C. lesueuri same forearm range for C. seabrae as previous au- no other records of C. seabrae, other than the thors (Hayman and Hill, 1971; Meester et al., 1986), holotype, were known. Roberts (1919: 112) described but reduced the minimum size for C. lesueuri to C. lesueuri as differing ‘most markedly from Cistugo 34 mm, and introduced colour differences as indi- seabrae in size, the cranial and external characters cated in the original species descriptions, the dorsal seemingly differing but little; there appear to be no colour of C. seabrae being described as a dull grey- glands on the wing, however, and the colouration is ish-yellow, relative to the honey-yellow dorsal colour somewhat different, both of which may eventually in C. lesueuri. The distribution reported by Taylor prove to be of little importance’. Cistugo seabrae (2000) for C. seabrae was similar to that given by was described from Mossamedes in Angola Hayman and Hill (1971) and Meester et al. (1986), (Thomas, 1912), whereas C. lesueuri was described i.e., Northern Cape, Namibia, southwestern Angola, from Lormarins [= L’Ormarins, Franschhoek but was more extensive for C. lesueuri, i.e., besides Valley], Paarl District, in the Western Cape of the distribution in the Western and Northern Cape, South Africa. Corbet and Hill (1980) synonymized the species was also reported to extend into the C. lesueuri with C. seabrae, given the limited distinc- Free State in South Africa and Lesotho. tion between the two taxa other than size and colour. Shortridge (1942: 37, 79) and Herselman and Hayman and Hill (1971) and subsequent authors Norton (1985: 94, 96) comment on differences (Honacki et al., 1982; Meester et al., 1986; Taylor, between C. lesueuri and C. seabrae in the number, 2000; Simmons, 2005; Monadjem et al., 2010), shape, size and position of the glands, those in however, retained C. lesueuri as a separate species C. lesueuri being narrower, less thick and a shorter from C. seabrae, although they usually indicated distance to the middle of the radius than those in some reservation about the validity of C. lesueuri, C. seabrae, and in C. lesueuri they may only occur pointing out the taxonomic relationship between on one wing, whereas in C. seabrae they are present these two species requires revision. on both wings and in some cases there are two The classification proposed to distinguish the two glands rather than one gland present on each side. species was initially based on measurements of However, these characters associated with the wing each of the holotype individuals, thus according to glands have not been extensively evaluated across Roberts (1954) the holotype of C. seabrae, which all specimens, possibly in part because the glands was known at the time from Mossamedes in Angola are not always visible on the specimens. to Goodhouse on the Orange River was smaller, Based on the complete mitochondrial cyto- with a forearm and greatest skull length of 32.5 mm chrome-b sequence Stadelmann et al. (2004: 185) and 13.2 mm, respectively; by contrast, the reported the ‘two closely related species differed by holotype of the larger C. lesueuri measured less than 30 substitutions from each other’, and that 34.5 mm and 14.4 mm. As new specimens of these within each species two individuals of C. lesueuri relatively rare species have been collected and showed no substitutions, while two individuals of reported (Smithers, 1983; Herselman and Norton, C. seabrae differed by two substitutions. Bickham SEAMARK ET AL.: TAXONOMY AND DISTRIBUTION OF CISTUGO SEABRAE AND C.

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