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Link to original content: https://pubmed.ncbi.nlm.nih.gov/22984399
Four new bat species (Rhinolophus hildebrandtii complex) reflect Plio-Pleistocene divergence of dwarfs and giants across an Afromontane archipelago - PubMed Skip to main page content
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. 2012;7(9):e41744.
doi: 10.1371/journal.pone.0041744. Epub 2012 Sep 12.

Four new bat species (Rhinolophus hildebrandtii complex) reflect Plio-Pleistocene divergence of dwarfs and giants across an Afromontane archipelago

Peter J Taylor  1 , Samantha Stoffberg, Ara Monadjem, Martinus Corrie Schoeman, Julian Bayliss, Fenton P D Cotterill
Affiliations

Four new bat species (Rhinolophus hildebrandtii complex) reflect Plio-Pleistocene divergence of dwarfs and giants across an Afromontane archipelago

Peter J Taylor et al. PLoS One. 2012.

Abstract

Gigantism and dwarfism evolve in vertebrates restricted to islands. We describe four new species in the Rhinolophus hildebrandtii species-complex of horseshoe bats, whose evolution has entailed adaptive shifts in body size. We postulate that vicissitudes of palaeoenvironments resulted in gigantism and dwarfism in habitat islands fragmented across eastern and southern Africa. Mitochondrial and nuclear DNA sequences recovered two clades of R. hildebrandtii senso lato which are paraphyletic with respect to a third lineage (R. eloquens). Lineages differ by 7.7 to 9.0% in cytochrome b sequences. Clade 1 includes R. hildebrandtii sensu stricto from the east African highlands and three additional vicariants that speciated across an Afromontane archipelago through the Plio-Pleistocene, extending from the Kenyan Highlands through the Eastern Arc, northern Mozambique and the Zambezi Escarpment to the eastern Great Escarpment of South Africa. Clade 2 comprises one species confined to lowland savanna habitats (Mozambique and Zimbabwe). A third clade comprises R. eloquens from East Africa. Speciation within Clade 1 is associated with fixed differences in echolocation call frequency, and cranial shape and size in populations isolated since the late Pliocene (ca 3.74 Mya). Relative to the intermediate-sized savanna population (Clade 2), these island-populations within Clade 1 are characterised by either gigantism (South African eastern Great Escarpment and Mts Mabu and Inago in Mozambique) or dwarfism (Lutope-Ngolangola Gorge, Zimbabwe and Soutpansberg Mountains, South Africa). Sympatry between divergent clades (Clade 1 and Clade 2) at Lutope-Ngolangola Gorge (NW Zimbabwe) is attributed to recent range expansions. We propose an "Allometric Speciation Hypothesis", which attributes the evolution of this species complex of bats to divergence in constant frequency (CF) sonar calls. The origin of species-specific peak frequencies (overall range = 32 to 46 kHz) represents the allometric effect of adaptive divergence in skull size, represented in the evolution of gigantism and dwarfism in habitat islands.

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Conflict of interest statement

Competing Interests: The authors have declared that no competing interests exist.

Figures

Figure 1
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.
Figure 2
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 Kenya, respectively.
Figure 3
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. hildebrandtii, RD - R. darlingi, RE - R. eloquens, RF - R. fumigatus, RH - R. hildebrandtii s.l., RL - R. landeri and RR - R. ruwenzorii. Localities, where available, are provided, abbreviations include SA - South Africa, MZ - Mozambique, and ZW - Zimbabwe, and the numbers in parentheses correspond with place names in Table S1 and Fig. 2 for Clade 1 and 2 individuals.
Figure 4
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 circles, 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). Hereafter, 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.).
Figure 5
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 (“H*”) and eloquens (“E*”) 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.; Lutope, Zimbabwe); open diamonds = R. eloquens type series (Clade 3); crosses in circles = R. hildebrandtii type and co-type (Clade 1c).
Figure 6
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 41997, smithersi from Pafuri) and positive (right: DM 11560, cohenae from Mayo, Mpumalanga Province) extremes of variation on RW1. Landmark positions (filled circles) are shown in the photograph in the centre.
Figure 7
Figure 7. Relative warps analysis (RWA) of 12 lateral cranial landmarks from 23 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 is 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: DM 8577, mossambicus from Namapa, Mozambique) and positive (right: DM 11560, cohenae from Mayo, Mpumalanga Province) extremes of variation on RW1. Landmark positions (filled circles) are shown in the photograph in the centre.
Figure 8
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).
Figure 9
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.)), 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).
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This research was supported by logistic and financial support from the Darwin Initiative (Darwin Initiative Award 15/036: Monitoring and Managing Biodiversity Loss in South-East Africa's Montane Ecosystems), the Mulanje Mountain Conservation Trust, the African Butterfly Research Institute and the Instituto de Investigação Agrária de Moçambique to JB. PJT and SS received grants from the National Research Foundation of South Africa. SS was further supported by a Claude Leon Foundation Postdoctoral Fellowships. FPDC thanks the Bay Foundations for support over the period core data were collected. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

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