Rhinolophus cohenae, Peter J. Taylor & Samantha Stoffberg & Ara Monadjem & Martinus Corrie Schoeman & Julian Baylis & Fenton P. D. Cotterill, 2012

Peter J. Taylor, Samantha Stoffberg, Ara Monadjem, Martinus Corrie Schoeman, Julian Baylis & Fenton P. D. Cotterill, 2012, Four New Bat Species (Rhinolophus hildebrandtii Complex) Reflect Plio-Pleistocene Divergence of Dwarfs and Giants across an Afromontane Archipelago, PLoS ONE 7 (9), pp. 1-23 : 16-17

Rhinolophus cohenae new species urn:lsid:zoobank.org:act:D0E1C9DD-7D36-4C01-AE0D-60AD474960A0

Cohen’s Horseshoe Bat

Fig. 4, 4 , Fig. S1 , Table 3, 4, Table S1, Appendix S1

Holotype. DM 8626 ; adult male, alcohol skin, skull and prepared baculum ( Fig. 9f ), collected by L. Cohen, 28 September 2004. Skull and mandible in good condition.

Type locality. Barberton, Mountainland Nature Reserve, 68 km SE Sudwala, Mpumalanga Province, South Africa, 25 ° 43 ' 8 " S; 31 ° 15 ' 58 " E; elevation 690 m asl.

Diagnosis. Lowest recorded peak frequency (33.0 kHz in the holotype; mean 32.8±0.24 kHz, n = 7 in type series) which immediately distinguishes this species from all others. Noseleaf extremely wide, 13.5–16.3 mm (15.5 mm in holotype), exceeding the previously described range (12–15) for the species [ 12]. Body size and cranial size very large ( Table 5; FL 66–68 mm; holotype 65.9 mm; GSL 29–30 mm; CCL 24–26 mm; holotype 29.9 mm and 25.6 mm respectively). In body and cranial size this species overlaps with mabuensis ; however the latter species has a distinct echolocation frequency (ca 38 kHz) and a distinct baculum (‘‘ Type 2’’). Anterior upper premolar conspicuous and located within tooth row or entirely absent. In the holotype and in three additional specimens, the anterior premolar is prominent and located within the toothrow. In another three specimens, the anterior premolar is absent (and the canine and posterior premolar in contact). In another specimen the prominent anterior premolar is situated in the toothrow on the right side but absent on the left side, leaving a small gap between canine and posterior premolar. In no cases were specimens found to have small anterior premolars situated external to the toothrow, thus distinguishing this species from hildebrandtii s.s. [ 12]. The baculum (total length 3.5 mm in holotype) has a unique shape (‘‘ Type 1’’; Fig. 9 ) characterised by narrow, laterally compressed and ventrally deflected shaft, emarginated distal portion and spatulate tip, which is not found in other species of the hildebrandtii complex in which the baculum has been described ( mossambicus and mabuensis ). Genotypes of R.cohenae are members of Clade 1a ( Fig. 2 ).

Paratypes. DM 7886 (adult male, alcohol skin and skull, collected by L. Cohen on 27 September 2004 from type locality), DM11620 , DM 11919 , DM 11918 (adult males, alcohol skin and skull, collected by S. Stoffberg and L. Cohen on 27 September 2009) .

Description. Dorsal and ventral colour similar to R. hildebrandtii s.s., but larger in noseleaf, skull and external dimensions as described above. The noseleaf shows the typical low, rounded shape of the connecting process in lateral view, in the holotype ( Fig. 8b ) and in other specimens examined (e.g. Fig. 8a ). Lower lip with single mental groove. The robust skull is represented in the mean cranial dimensions ( Table 5; Fig. 5 ). In lateral view the skull has a prominent rostral chamber that extends to the same height as the sagittal crest; albeit the cranium is distinctly elongated and flattened in lateral profile ( Fig. 6 ). In dorsal view the V-shaped inter-orbital basin formed by the supraorbital ridges is very prominent, with the anterior root of the sagittal crest displaced posteriorly ( Fig. 6 ).

Distribution. Known only from three closely-spaced localities in Mpumalanga Province of South Africa in the vicinity of Nelspruit.

Ecology. The type locality of Barberton is located in the Savanna Biome [ 114] at 690 m, close to the border of the Grassland Biome. Two nearby additional localities (Sudwala and Mayo Mines) are located in the Grassland Biome (Mesic Highveld Grassland) at altitudes between 900 and 1100 m.

Etymology. We selected the specific epithet to recognize Lientjie Cohen who collected the type specimen. She has contributed significantly to the conservation of bats in South Africa, particularly in Mpumalanga Province. The species name combines the surname Cohen and genitive singular case-ending ‘‘ae’’ indicative of feminine gender.

Specimens examined. See Table S1.

Figures

Figure 1. Portraits of (a) Rhinolophus smithersi species novo, and (b) Rhinolophus mossambicus species novo, two of four new cryptic species described herein within the R. hildebrandtii complex. doi:10.1371/journal.pone.0041744.g001

Figure 2. Map of southern, central and eastern Africa indicating localities of individuals of R. hildebrandtii species-complex included in this study. Grey-shaded area represents elevations in excess of 600 m a.s.l. Closed squares indicate museum specimens from which craniometric data were obtained. Open symbols indicate specimens genotyped in this study. The distribution of the three major clades is based on cytochrome b (see Figure 2): open circles = Clade 1; open squares = Clade 2; open diamonds = Clade 3. Closed squares enclosed in open symbols indicate localities where both molecular and morphological data were available for selected specimens. Numbers refer to respective localities listed in Table S1. ‘‘T’’ indicates the type localities of R. eloquens in Uganda and R. hildebrandtii in Kenyaı respectively. doi:10.1371/journal.pone.0041744.g002

Figure 4. Morphometric variation in a series representing the R. hildebrandtii complex from Lutope-Ngolangola, Zimbabwe: a) biplot of forearm length versus noseleaf width and b) PCA of five craniometric variables (M3M3, CM3, IOC, NW, NH) in 26 individuals of known (37 or 46 kHz) and unknown (?) frequency. Females indicated by open circlesı males by closed circles or crosses or asterisk. Voucher specimens for molecular sequencing study indicated by asterisk (Clade 1e: = smithersii sp. nov.; see Taxonomic Conclusions) and crosses (Clade 2: = mossambicus sp. nov.; see Taxonomic Conclusions). Hereafterı all individuals with a frequency of 37 kHz were assumed to belong to Clade 2 (mossambicus sp. nov.) and the 46 kHz individual was assumed to belong to Clade 1e (smithersi sp. nov.). doi:10.1371/journal.pone.0041744.g004

Figure 5. Canonical variates analysis (CVA) (a) of 10 cranial variables in five groups of the Rhinolophus hildebrandtii complex defined by molecular analysis; and PCA (b) of five cranial variables for sample in (a) with type series of hildebrandtii (‘‘H¹’’) and eloquens (‘‘E¹’’) added. Open circles = Clade 1a (= cohenae sp. nov.); closed circles = Clade 1b (= mabuensis sp. nov.); shaded circles = Clade 1d (= smithersi sp. nov.; Pafuri); asterisk enclosed in circle = Clade 1e (= smithersi sp. nov.; Zimbabwe); open squares = Clade 2 (mossambicus sp. nov.; Mozambique); shaded squares = Clade 2 (mossambicus sp. nov.; Lutopeı Zimbabwe); open diamonds = R. eloquens type series (Clade 3); crosses in circles = R. hildebrandtii type and co-type (Clade 1c). doi:10.1371/journal.pone.0041744.g005

Figure 6. Relative warps analysis (RWA) of 13 dorsal cranial landmarks from 22 individuals of R. hildebrandtii s.l. belonging to two molecular clades and two lineages of Clade 1 (see Fig. 3). Revised taxon names are provided in parentheses (see Taxonomic Conclusions). Skulls which were included in this analysis are indicated in Table S1. Symbols as in Fig. 5. Thin plate splines (grids) show landmark distortions represented by extremes of variation on RW1 (left = negative; right = positive) and RW2 (bottom = negative; top = positive) axes. The two skull photographs at the bottom are of actual specimens representing the negative (left: TM 41997ı smithersi from Pafuri) and positive (right: DM 11560ı cohenae from Mayoı Mpumalanga Province) extremes of variation on RW1. Landmark positions (filled circles) are shown in the photograph in the centre. doi:10.1371/journal.pone.0041744.g006

Figure 8. Photographs showing lateral views of noseleafs of selected individuals (including holotypes of new species) of the Rhinolophus hildebrandtii complex representing molecular Lineage 1a (= cohenae sp. nov.; a–b), Lineage 1b (= mabuensis sp. nov.; c) and Clade 2 (= mossambicus sp. nov.; d–f). a = DM 7886 (cohenae sp. nov.;Barberton Tunnel, Mpumalanga Province, South Africa); b = DM 8626 (cohenae sp. nov.; Barberton Tunnel, Mpumalanga Province, South Africa; Holotype); c = DM 10842 (mabuensis sp. nov.; Mt Mabu, Mozambique; Holotype); d = DM 8578 (mossambicus sp. nov.; Niassa Game Reserve, Mozambique; Holotype); e = DM 8579 (mossambicus sp. nov.; Chinizuia, Mozambique); f = DM 8577 (mossambicus sp. nov.; Namapa, Mozambique). doi:10.1371/journal.pone.0041744.g008

Figure 9. Dorsal (D), ventral (V) and lateral (L) view of bacula (tips on right) from four individuals (a–d) from Mpumalanga (Clade 1a = cohenae sp. nov.), two (e–f) from lowland sites in Mozambique (Clade 2 = mossambicus sp. nov.) and one (g) from Mt Mabu in Mozambique (Clade 1b = mabuensis sp. nov.). a = DM 11558 (Sudwala); b = DM 11620 (Barberton Tunnel; Topotype of cohenae); c = DM 11560 (Mayo); d = DM 11618 (Barberton Tunnel); e = DM 8580 (Gorongosa); f = DM 8578 (Niassa GR; Holotype of mossambicus); g = DM 10842 (Mt Mabu; Holotype of mabuensis). Bacula of Clade 1a (cohenae sp. nov.) have spatulate tip (rounded in Clades 2 (mossambicus sp. nov.) and 1b (mabuensis sp. nov.))ı typically emarginated basal portion (less so in Clades 2 and 1b) and shaft laterally compressed (cylindrical in Clades 2 and 1b) and sloping downwards in lateral view (horizontal in Clades 2 and 1b). doi:10.1371/journal.pone.0041744.g009