Rhinolophus hildebrandtii Petersı 1878

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 : 17-19

Rhinolophus hildebrandtii Petersı 1878

Hildebrandt’s Horseshoe Bat

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

Holotype. ZMB 5378 (male collected by S. Hildebrandt in July 1877 from Ndi , Taita, Kenya). Cotype ZMB 5379 (male collected by S. Hildebrandt in July 1877 from Ndi , Taita, Kenya).

Type locality. Ndi , Taita, Kenya, 2 ° 46 ' 43 " S; 38 ° 46 ' 18 " E; elevation 390 m asl.

Diagnosis. Individuals from two localities in Kenya, including the type locality of Taita District, had a peak frequency of 42 kHz [ 15]. Intermediate in body and cranial size, with the type and co-type overlapping in cranial dimensions with members of Clade 2 from which they are however easily distinguished genetically based on sequences from three mitochondrial and one nuclear gene (cytochrome b uncorrected genetic divergence 7.7–9.0%). Although there is minor overlap in individual cranial and forearm variables ( Table 5), they can be distinguished completely by PCA of cranial dimensions from other members of Clade1 which are distinctly smaller (Lineages 1d, 1e) or larger (Lineages 1a, 1b) ( Fig. 5b ). The anterior upper premolar, when present is moderate-sized but situated external to the toothrow (with the canine and P 4 in contact) [ 12]. In this respect, they are distinguished from cohenae which have relatively conspicuous anterior premolars within the toothrow and smithersi in which the anterior premolar is small to minute and external to the toothrow. Genotypes of R. hildebrandtii are members of Clade 1c ( Fig. 2 ).

Freq.= peak (time expansion or full-spectrum detector) or maximum (ANABAT detector) frequency; NLW = nasal width; FL = forearm length; GSL = greatest skull length; CCL = condylo-canine skull length. *Echolocation frequency of holotype not measured but assumed on basis of two released individuals of the same species at Mt Mabu recorded during the same month (October 2008). **We assume the holotype and paratype specimens from the Taita District may have had a CF frequency of ca 42 kHz based on recent recordings from the type locality. ***The holotype was not recorded but we assume a CF frequency of 35–38 kHz based on recordings of four bats taken from nearby localities in northern Mozambique. doi:10.1371/journal.pone.0041744.t005

Paratype. BM 79.1.21.1 , Taita (un-sexed individual collected in 1877 by S. Hildebrandt) .

Description. Follows the general description for the species [ 12]; however, hildebrandtii s.s. as defined here (restricted to East Africa) is intermediate-sized in most external and cranial dimensions and varies from the range of values reported previously for R. hildebrandtii s.l. [ 12], for example noseleaf width in nine East African specimens examined varied from 10–14 mm, rather than 12–15 mm given by previous description [ 12].

Distribution. As here defined, and based only on limited material examined from Kenya and Tanzania in the Natural History Museum, London and Berlin Museum in this study (n = 11; mensural data reported in Table 5), we restrict the distribution of hildebrandtii s.s. to East Africa, whilst recognising that northern populations in Democratic Republic of Congo, Ethiopia and southern Sudan (now South Sudan) are possibly referable to this species (as discussed by Koopman 1975, [ 13]. Moreover, given that two individuals with peak frequency of 37 kHz from Lutope-Ngolangola in NW Zimbabwe grouped genetically with individuals from coastal Mozambique assigned to Clade 2, it seems probable that hildebrandtii as here defined (coinciding with the divergent Lineage 1c) does not occur in southern Africa. Nevertheless, the taxonomic affinities of material from Zambia and Malawi remain unclear as craniometric analysis was unable to differentiate molecular clades and lineages at this scale ( Fig. S2 ). Future studies need to incorporate molecular and bacular data from central, equatorial and northeastern Africa (particularly Zambia, Malawi, DRC and Ethiopia) to test the species limits of hildebrandtii . It is possible that yet more cryptic species remain to be discovered.

Ecology. This species was recorded from montane and submontane forests in the Usambara Mountains of the Eastern Arc Range in Tanzania; means and ranges given for external and cranial measurements for four individuals from this population conform very closely to our data [ 119].

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