Rhinolophus smithersi, 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 : 19

Rhinolophus smithersi new species urn:lsid:zoobank.org:act:E78B0F44-7534-4991-975AF176CB668DDE

Smithers’s Horseshoe Bat

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

Holotype. NMZB 33647 , alcohol and cleaned skull. Collector Number FWC 4764. Female collected 26 October 2000 by F. P. D. Cotterill and P. J. Taylor. Skull and mandible in good condition.

Type locality. Ngolangola Gorge at confluence with Lutope River , Sebungwe District, Gokwe Communal Land, NW Zimbabwe; 18 ° 17 ' 05 " S; 28 ° 05 ' 00 " E; elevation 1000 m asl.

Diagnosis. Peak echolocation frequency at 44–46 kHz which is considerably higher than all other members of this speciescomplex (33–42 kHz). Very small-cranium (mean CCL 23.4 mm; 23.0– 24.3 mm; n = 6; 23.2 mm in holotype), the holotype having a disproportionately wide noseleaf (14 mm; 53% of skull length) (although this is not as pronounced in the series from Pafuri, NE South Africa; mean noseleaf width 10.7 mm; 39% of skull length), it is immediately distinguishable from other members of the R. hildebrandtii complex on size and peak frequency ( Figs. 3 ; 4 ; Table 5). Genotypes of R. smithersi are members of Clades 1d and 1e ( Fig. 2 , Fig. S1 ).

Paratype. NMZB 33652 , male alcohol and cleaned skull, Collector Number FWC 4769. Collected 26th October 2000 by F.P.D. Cotterill and P.J. Taylor from Lutope-Ngolangola Gorge, Zimbabwe .

Description. Apart from its distrinctly smaller size, conforming generally to the keys and published description for R. hildebrandtii s.l. [ 12], i.e. a relatively large-sized (forearm length 60.7 mm in holotype) bat with medium or large ears, wide horseshoe (10–14 mm) covering the muzzle, sella constricted at its proximal third, almost parallel-sided above and having long hairs, lancet relatively long and straight-sized, lower lip with single median groove, upper parts greyish-brown, with individual hairs long and unicoloured, underparts same colour or very slightly paler, skull large and heavily built. Lower lip with single mental groove. The sagittal crest is very well developed in relation to the nasal inflation and the rostral depression is weakly developed. The anterior premolar is small-sized and external to the toothrow ( Table 5).

Distribution. Known from Lutope-Ngolangola Gorge south of the Zambezi Escarpment in NW Zimbabwe and also from Pafuri in the Limpopo Valley in the foothills of the Soutpansberg Mountains of northern Limpopo Province, South Africa but likely more widespread across savanna woodlands of the Limpopo and Zambezi valleys, and their escarpments (the Gwembe horst, and the Soutpansberg and Waterberg Mountains, respectively). Accurate delimitation of this species’ range is subject to further collecting and reappraisal of existing museum material. This applies to papers by M.B. Fenton and co-workers based on intensive studies of the bat fauna, which included populations of this species complex in the Sengwa Wildlife Research Area, only, 5 km north of the type locality of R. smithersi [120,121,122,123,124,125,126]. These all reported the largest rhinolophid studied as R. hildebrandtii , but these data would refer to both R. mossambicus and R. smithersi .

Ecology. Poorly known; this species occurs sympatrically with R. mossambicus at one locality in miombo savanna on Karoo Sandstone, dominated by trees of Brachystegia glaucescens , and including large specimens of baobabs, Adansonia digitata ; albeit much of the landscape has been converted to cotton fields. More diverse riparian woodland fringes the Lutope and Ngolanola rivers as well as along the Limpopo River at Pafuri. Daylight roosts were not located but these bats could use caves in the sandstone cliffs and/or hollows in baobabs.

Etymology. We selected the specific epithet in recognition of the late Reay Henry Noble Smithers (1907–1987), former Director of the National Museums of Zimbabwe, prodigious collector and researcher of mammals including bats, and author of important regional texts on the mammalogy of Botswana, Zimbabwe and Mozambique, notably his definitive monograph – The Mammals of the Southern African Subregion subsequently updated and revised [ 127]. The species name combines the surname Smithers and genitive singular case-ending ‘‘i’’ indicative of masculine 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 3. Consensus tree for the cytochrome b dataset for representative genotyped specimens of the Rhinolophus hildebrandtii complex. The topology represents the consensus topology from a 20 million MCMC run implemented in BEAST. Estimates of divergence times (million years ago; Mya) are indicated adjacent to nodes or above branches and grey bars indicate 95% HPD values. The split between the Hipposideridae and Rhinolophidae was used as the calibration point. Taxa names include museum/field numbers which correspond to Appendix S1 or GenBank accession numbers and abbreviations are: RcfH - R. cf. hildebrandtiiı RD - R. darlingiı RE - R. eloquensı RF - R. fumigatusı RH - R. hildebrandtii s.l.ı RL - R. landeri and RR - R. ruwenzorii. Localitiesı where availableı are providedı abbreviations include SA - South Africaı MZ - Mozambiqueı and ZW - Zimbabweı and the numbers in parentheses correspond with place names in Table S1 and Fig. 2 for Clade 1 and 2 individuals. doi:10.1371/journal.pone.0041744.g003

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