Evolutionary relationships and population genetics of the Afrotropical leaf-nosed bats (Chiroptera, Hipposideridae)

Abstract

The Old World leaf-nosed bats (Hipposideridae) are aerial and gleaning insectivores that occur throughout the Paleotropics. Both their taxonomic and phylogenetic histories are confused. Until recently, the family included genera now allocated to the Rhinonycteridae and was recognized as a subfamily of Rhinolophidae. Evidence that Hipposideridae diverged from both Rhinolophidae and Rhinonycteridae in the Eocene confirmed their family rank, but their intrafamilial relationships remain poorly resolved. We examined genetic variation in the Afrotropical hipposiderids Doryrhina, Hipposideros, and Macronycteris using relatively dense taxon-sampling throughout East Africa and neighboring regions. Variation in both mitochondrial (cyt-b) and four nuclear intron sequences (ACOX2, COPS, ROGDI, STAT5) were analyzed using both maximum likelihood and Bayesian inference methods. We used intron sequences and the lineage delimitation method BPP—a multilocus, multi-species coalescent approach—on supported mitochondrial clades to identify those acting as independent evolutionary lineages. The program StarBEAST was used on the intron sequences to produce a species tree of the sampled Afrotropical hipposiderids. All genetic analyses strongly support generic monophyly, with Doryrhina and Macronycteris as Afrotropical sister genera distinct from a Paleotropical Hipposideros; mitochondrial analyses interpose the genera Aselliscus, Coelops, and Asellia between these clades. Mitochondrial analyses also suggest at least two separate colonizations of Africa by Asian groups of Hipposideros, but the actual number and direction of faunal interchanges will hinge on placement of the unsampled African-Arabian species H. megalotis. Mitochondrial sequences further identify a large number of geographically structured clades within species of all three genera. However, in sharp contrast to this pattern, the four nuclear introns fail to distinguish many of these groups and their geographic structuring disappears. Various distinctive mitochondrial clades are consolidated in the intron-based gene trees and delimitation analyses, calling into question their evolutionary independence or else indicating their very recent divergence. At the same time, there is now compelling genetic evidence in both mitochondrial and nuclear sequences for several additional unnamed species among the Afrotropical Hipposideros. Conflicting appraisals of differentiation among the Afrotropical hipposiderids based on mitochondrial and nuclear loci must be adjudicated by large-scale integrative analyses of echolocation calls, quantitative morphology, and geometric morphometrics. Integrative analyses will also help to resolve the challenging taxonomic issues posed by the diversification of the many lineages associated with H. caffer and H. ruber.

Introduction

The Old World leaf-nosed bats, family Hipposideridae, currently include seven genera and 90 species of insectivorous bats distributed over much of the Paleotropics (Monadjem 2019; Simmons and Cirranello 2019). Both the taxonomic and phylogenetic histories of this family are confused. Throughout much of its history (e.g., Koopman 1989), Hipposideridae was considered either a subfamily of the Rhinolophidae (the horseshoe bats) or as its sister family within the Rhinolophoidea. Recently, however, the “trident bats” (Cloeotis, Paratriaenops, Rhinonicteris, and Triaenops) were shown to comprise a family-ranked group, the Rhinonycteridae, which is separate from and sister to the Hipposideridae (Foley et al. 2015; Armstrong et al. 2016). Even the genus Hipposideros Gray, 1831, as it was traditionally understood, appears paraphyletic with respect to the allied genera Asellia, Aselliscus, Coelops, and Anthops (Foley et al. 2015; Amador et al. 2018). Re-validation of Macronycteris Gray, 1866 and Doryrhina Peters, 1871 for groups of Afrotropical endemic species more closely related to each other than to African and Asian members of Hipposideros sensu stricto resolved a number of those issues (Foley et al. 2017).

The species richness of Doryrhina, Macronycteris, and Hipposideros differs widely. Most authors recognize two species of Doryrhina (D. cyclops and D. camerunensis), five species of Macronycteris (M. commersoni, M. cryptovalorona, M. gigas, M. thomensis, and M. vittata), and 83 species of Hipposideros, 10 of which occur in Africa (Monadjem 2019; Simmons and Cirranello 2019). These are H. beatus, H. caffer, H. curtus, H. fuliginosus, H. lamottei, H. ruber, and H. tephrus in the bicolor group of Hipposideros; H. jonesi and H. marisae in the speoris group, and H. megalotis in the megalotis group (Hill 1963; Murray et al. 2012; Monadjem 2019). In addition, three extinct species of hipposiderid are known from the region: †Macronycteris besaoka (Madagascar), †Hipposideros amenhotepos (Egypt), and †H. kaumbului (Ethiopia). Type localities for valid species, subspecies, and synonyms for these three genera in Africa and Madagascar appear in Figure 1; after the removal of Doryrhina and Macronycteris taxa, group assignments for the species remaining in Hipposideros appear in Table 1.

Figure 1.

Type localities for Afrotropical hipposiderids: Doryrhina, blue symbols; Hipposideros, white symbols; Macronycteris, black symbols. Stars denote valid species, whereas circles indicate taxa considered as subspecies or synonyms. Localities are projected onto the biome map of Olson et al. (2001). Taxa depicted are: Hipposideros abae J. A. Allen,1917; †Hipposideros (Pseudorhinolophus) amenhotepos Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders & Gingerich, 2015; Phyllorhina angolensis Seabra, 1898; Hipposideros caffer var. aurantiaca De Beaux, 1924; Hipposideros beatus K. Andersen, 1906; †Hipposideros besaoka Samonds, 2007; Phyllorrhina bicornis Heuglin, 1861; Hipposideros braima Monard, 1939; Hipposideros caffer Sundevall, 1846; Phyllorhina caffra Peters, 1852; Hipposideros camerunensis Eisentraut, 1956; Hipposideros caffer centralis K. Andersen, 1906; Rhinolophus Commersonii É. Geoffroy, 1813; Hipposideros cryptovalorona Goodman, Schoeman, Rakotoarivelo & Willows-Munro, 2016; Hipposideros curtus G. M. Allen, 1921; Phyllorrhina cyclops Temminck, 1853; Phyllorrhina fuliginosa Temminck, 1853; Hipposideros gigas gambiensis K. Andersen, 1906; Rhinolophus gigas Wagner, 1845; Phyllorrhina gracilis Peters, 1852; Hipposideros caffer guineensis K. Andersen, 1906; Hipposideros jonesi Hayman, 1947; †Hipposideros kaumbului Wesselman, 1984; Hipposideros lamottei Brosset, 1985; Hipposideros langi J. A. Allen, 1917; Hipposideros marisae Aellen, 1954; Phyllorhina Commersoni, var. marungensis Noack, 1887; Hipposideros beatus maximus Verschuren, 1957; Phyllorrhina megalotis Heuglin, 1861; Rhinolophus micaceus de Winton, 1897; HipposiderosCommersoni mostellum Thomas, 1904; Hipposideros nanus J. A. Allen, 1917; Hipposideros gigas niangarae J. A. Allen, 1917; Hipposideros caffer niapu J. A. Allen, 1917; Phyllorrhina rubra Noack, 1893; Hipposideros sandersoni Sanderson, 1937; Hipposideros tephrus Cabrera, 1906; Phyllorhina Commersoni, var. thomensis Bocage, 1891; Hipposideros gigas viegasi Monard, 1939; Phyllorhina vittata Peters, 1852.

Table 1.

Species groups of Hipposideros (modified from Murray et al. 2012 to include newly recognized forms and to remove species now recognized in Doryrhina and Macronycteris).

Armiger group	calcaratus subgroup	H. macrobullatus	H. lankadiva
H. alongensis	H. calcaratus c	H. maggietaylorae	H. lekaguli
H. armiger	H. cervinus c	H. nequam	H. pelingensis
H. griffini a	H. coxi c	H. obscurus	Larvatus group
H. pendelburyi a	H. galeritus c	H. orbiculus	H. grandis
H. turpis	ruber subgroup	H. papua	“H. khasiana” a,g
Bicolor group	H. abae d	H. pygmaeus	H. larvatus
ater subgroup	H. beatus e	Boeadii group	H. madurae
H. ater b	H. caffer e	H. boeadii	H. sorenseni
H. atrox a	H. fuliginosus e	Cyclops group f	H. sumbae
H. bicolor b	H. lamottei e	H. corynophyllus	Megalotis group
H. breviceps b	H. ruber e	H. edwardshilli	H. megalotis
H. cineraceus b	H. tephrus a,e	H. muscinus	Pratti group
H. coronatus b	subgroup uncertain	H. semoni	H. lylei
H. dyacorum b	H. cruminiferus	H. stenotis	H. pratti
H. einnaythu a,b	H. curtus	H. wollastoni	H. scutinares
H. halophyllus b	H. doriae	Diadema group	Speoris group
H. khaokhouayensis b	H. durgadasi	H. demissus	H. jonesi h
H. nicobarulae a,b	H. fulvus	H. diadema	H. marisae g
H. pomona b	H. gentilis a	H. dinops	H. speoris
H. ridleyi b	H. hypophyllus	H. inexpectatus
H. rotalis b	H. kunzi a	H. inornatus

(Endnotes) a Added to species list subsequent to Murray et al. (2012) b Recognized in the Ater species group by Monadjem (2019) c Recognized in the Calcaratus species group by Monadjem (2019) d Formerly listed in the Speoris group but transferred to the Ruber group by Monadjem (2019) e Recognized in the Ruber species group by Monadjem (2019) f H. cyclops and H. camerunensis are now recognized as members of Doryrhina; listed species were treated as Doryrhina inMonadjem (2019) on the basis of similar morphology but were recognized as the Muscinus group by Tate (1941); they might represent an unnamed genus or subgenus. g Invalid name accorded to what is likely a real biological entity (cf. Monadjem 2019) h Formerly in the Bicolor species group but transferred to the Speoris group by Monadjem (2019).

As suggested by their checkered taxonomic history, phylogenetic understanding of the Hipposideridae has slowly come into focus. Doryrhina and Macronycteris are two of a dozen generic-group names that were synonymized with Hipposideros for all of the 20^th century (Miller 1907; Allen 1939; Koopman 1994). Instead of subgenera, taxonomists used the species groups delineated by Andersen (1918) and refined by Tate (1941) and Hill (1963) in their generic revisions based on morphology. Assessment of rhinolophoid relationships using an intron supermatrix (Eick et al. 2005) confirmed the early divergence of hipposiderids and rhinolophids (estimated at 41 Ma), thereby substantiating their rank as a separate families. Despite earlier suppositions that the area of origin for Hipposideridae was in Asia (Koopman 1970; Bogdanowicz and Owen 1998) or Australia (Hand and Kirsch 1998), Eick et al. (2005) clearly demonstrated the ancestry of the family (and superfamily) was in Africa. A recent supermatrix analysis with the most comprehensive taxonomic sampling (42 species; Amador et al. 2018) confirmed the early divergence of hipposiderids and rhinolophids at 41.3 Ma, but this analysis questioned the validity of both Doryrhina and Macronycteris. Amador et al. attributed the paraphyly of Hipposideros sensu lato documented by Foley et al. (2015) to their limited taxonomic sampling. Amador et al. (2018) also challenged the integrity of the commersoni, cyclops, speoris, and bicolor species groups, arguing that all African species save for H. jonesi belonged in a single, exclusively African species group.

Although new species of hipposiderids are regularly discovered and described in Asia (Robinson et al. 2003; Guillen-Servent and Francis 2006; Bates et al. 2007; Douangboubpha et al. 2011; Thong et al. 2012; Murray et al. 2018), the pace of discovery has been much slower in Africa. Only one extant species has been described since the recognition of Hipposideros lamottei (Brosset 1985 [“1984”]), and that one was from Madagascar (Goodman et al. 2016). Surveys of mitochondrial sequences from African hipposiderids have strongly suggested that supposedly widespread species such as Hipposideros caffer and H. ruber actually represent complexes of cryptic species (Vallo et al. 2008, 2011; Monadjem et al. 2013). Phylogenetic analyses (e.g., Vallo et al. 2008) show that these named species complexes are not monophyletic, resolving clades comprised of bats identified as both H. caffer and H. ruber. These studies have characterized the clades in both morphological and genetic terms, even establishing them in sympatry (see also Vallo et al. 2011). However, the uncertain relationship of the identified clades to the many names already proposed for Afrotropical hipposiderids, many based on incomplete or formalin-preserved specimens, has precluded formally naming them. Incomplete geographic sampling and the lack of evidence from nuclear genes for these populations has also clouded interpretations of this mitochondrial diversity.

Our field surveys in Eastern Africa and adjoining regions offer a new basis for considering the taxonomy and phylogenetics of Afrotropical hipposiderids. We sought to answer these questions: (1) Is there compelling evidence to support the recognition of Doryrhina and Macronycteris as distinct Afrotropical genera alongside the Paleotropical Hipposideros? (2) Which species belong to these groups? (3) Are the traditional species groups of African hipposiderids monophyletic? Using both mitochondrial and nuclear intron sequences, we also evaluate the question of cryptic species among African hipposiderids and the possibility of mitochondrial-nuclear discordance.

Material and methods

Selection of taxa and sampling

Our genetic dataset is based on 453 hipposiderid individuals, the vast majority being represented by museum vouchers. We generated original genetic data from 319 individuals collected at 102 georeferenced localities, and complemented them with 134 mitochondrial sequences from 90 localities downloaded from GenBank (we obtained new sequence data for five individuals with prior GenBank records; see Suppl. material 1: Figure S1 and Appendix I). All individuals were sequenced for Cytochrome-b (cyt-b) in order to maximize assessment of genetic diversity; however, redundant haplotypes were removed for subsequent phylogenetic analyses (see Appendix I for complete list of individuals sequenced). The bats newly sequenced for this study were obtained over several decades in the course of small mammal surveys across sub-Saharan Africa and Madagascar, with relatively dense sampling in East Africa. Initial assignment of East African individuals to species was determined using meristic, mensural, and qualitative characters published in the bat keys of Thorn et al. (2009) and Patterson and Webala (2012). Collection methods followed mammal guidelines for the use of wild mammals in research and education (Sikes and the Animal Care and Use Committee of the American Society of Mammalogists 2016) and the most recent collections were approved under Field Museum of Natural History’s IACUC #2012-003. Only GenBank records for cyt-b were available for records of the Arabian-North African hipposiderid Asellia, which was included for context in the phylogenetic analyses. Lacking information from nuclear introns, we draw no firm conclusions from their placement and do not discuss Asellia in this paper (see Benda et al. 2011; Bray and Benda 2016).

Appendix I contains the institutions and voucher numbers, GenBank accession numbers, and locality information for our samples. The fact that museum voucher specimens were used wherever possible for the genetic analyses permits the genetic analysis to serve as a foundation for integrative taxonomic analyses of dental, cranial, and skeletal variation, using the same specimens. To avoid adding to current taxonomic confusion, we take a conservative approach in assigning names to clades in our analyses. Where a clade’s taxonomic identity was ambiguous or unknown, we referred to it simply as a numbered clade. Integrative taxonomic diagnoses of the various clades supported by our analyses will be necessary to determine which, if any, existing names may apply to them. However, to relate our results to those of earlier studies of African Hipposideros (Vallo et al. 2008, 2011; Monadjem et al. 2013), we cross-referenced specimens used in two or more analyses to equate the various non-binomial names that have been applied to these cryptic lineages.

DNA extraction, amplification, and sequencing

Genomic DNA from preserved tissue samples was extracted using the Wizard SV 96 Genomic DNA Purification System (Promega Corporation, WI, USA). Fresh specimens were sequenced for mitochondrial cytochrome-b (cyt-b), using the primer pair LGL 765F and LGL 766R (Bickham et al. 1995; Bickham et al. 2004), and four unlinked autosomal nuclear introns: ACOX2 intron 3, COPS7A intron 4, ROGDI intron 7 (Salicini et al. 2011), and STAT5B (Matthee et al. 2001) for hipposiderid specimens and the sister group Triaenops afer (Rhinonycteridae; see Table 1 for primer information). PCR amplification, thermocycler conditions, and sequencing were identical to Patterson et al. (2018) and Demos et al. (2018). Sequences were assembled and edited using GENEIOUS PRO v.11.1.5 (Biomatters Ltd). Sequence alignments were made using MUSCLE (Edgar 2004) with default settings in GENEIOUS. Protein coding data from cyt-b were translated to amino acids to determine codon positions and confirm the absence of premature stop codons, deletions, and insertions. Several gaps were incorporated in the nuclear intron alignments, but their positions were unambiguous.

Sequence alignments used in this study have been deposited on the FIGSHARE data repository (https://doi.org/10.6084/m9.figshare.11936250). All newly generated sequences were deposited in GenBank with accession numbers MT149315–MT149893 (see also Appendix I).

Phylogenetic analyses

jMODELTEST2 (Darriba et al. 2012) on CIPRES Science Gateway v. 3.3 (Miller et al. 2010) was used to determine the sequence substitution models that best fit the data using the Bayesian Information Criterion (BIC) for cyt-b and the four nuclear introns. PARTITIONFINDER2 (Lanfear et al. 2016) on CIPRES was used to determine the sequence substitution models for the concatenated alignment of four nuclear introns using the Bayesian Information Criterion (BIC) with the ‘greedy’ search algorithm. Uncorrected sequence divergences (p-distances) between and within species/clades were calculated for cyt-b using MEGA X v. 10.0.5 (Kumar et al. 2018). Maximum-likelihood (ML) analyses were performed using the program IQ-TREE v. 1.6.10 (Chernomor et al. 2016; Nguyen et al. 2015) on the CIPRES portal for separate gene trees (cyt-b, ACOX2, COPS7A, ROGDI, and STAT5B) and a concatenated alignment, partitioned by gene, using the four nuclear introns. As in Hillis and Bull (1993), nodes supported by bootstrap values (BP) ≥ 70% were considered strongly supported. Gene tree analyses under a Bayesian Inference (BI) framework were inferred in MRBAYES v. 3.2.7 (Ronquist et al. 2012) on the CIPRES portal for the same set of genes as the ML analyses. Two independent runs were conducted in MrBayes, and nucleotide substitution models were unlinked across partitions for each nuclear locus in the concatenated alignment. Four Markov chains were run for 1 × 10⁸ generations for individual gene trees, and 2 × 10⁷ generations for the concatenated analysis, using default heating values and sampled every 1000^th generation. A conservative 25% burn-in was applied and stationarity of the MRBAYES results was assessed in Tracer v. 1.7 (Rambaut et al. 2018). Majority-rule consensus trees were constructed for each Bayesian analysis. Following Erixon et al. (2003), nodes supported by posterior probabilities (PP) ≥0.95 were considered strongly supported.

Haplotype networks for cyt-b were inferred using the median-joining network algorithm in PopArt v. 1.7 (Leigh and Bryant 2015). Separate analyses were carried out for the following clades, each consisting of four subclades: (1) Doryhina (D. camerunensis, D. cf. camerunensis, D. cyclops1, and D. cyclops2); (2) Macronycteris (M. commersoni, M. cryptovalorona, M. gigas, and M. vittata); (3) Hipposideros caffer1–4; (4) Hipposideros caffer5–8; and (5) Hipposideros ruber1–4.

Hipposiderid taxa included in the species tree analyses were assigned to either species or numbered clades based on clade support in the ML and BI gene-tree analyses of the cyt-b dataset. This in turn identified populations to be used as ‘candidate species’ in a coalescent-based species-tree approach implemented in StarBEAST2 (Ogilvie et al. 2017), an extension of BEAST v. 2.5.1 (Drummond et al. 2012; Bouckaert et al. 2014). Species tree analysis was conducted using the four nuclear intron alignments. Substitution, clock, and tree models were unlinked across all loci. A lognormal relaxed-clock model was applied to each locus under a Yule tree prior and a linear with constant root population size model. Four independent replicates were run with random starting seeds, and chain lengths of 1 × 10⁸ generations and parameters were sampled every 5,000 steps. For the StarBEAST2 analyses, evidence of convergence and stationarity of posterior distributions of model parameters was assessed based on ESS values >200 and examination of trace files in Tracer v. 1.7. The burn-in was set at 10% and separate runs were assembled using LOGCOMBINER v. 2.5.1 and TREEANNOTATOR v. 2.5.1 (Rambaut et al. 2018).

Coalescent lineage delimitation

Based on the well supported clades obtained in the cyt-b gene tree analyses and available intron samples, a lineage delimitation scenario with 18 candidate species was tested. We inferred the evolutionary isolation of their gene pools using the phased nuclear DNA dataset (ACOX2, COPS7A, ROGDI, and STAT5A; 104 individuals) for joint independent lineage delimitation and species-tree estimation evaluated under the multi-species coalescent model using the program BPP v. 3.3 (Yang and Rannala 2014; Rannala and Yang 2017). This analysis was carried out to guide future investigations of the species status of evolutionarily isolated lineages inferred here. Supported lineages will be examined using an integrative species taxonomic approach, including morphological, morphometric, and acoustic characters, as well as ectoparasite associations and distributional data. Species/clade memberships for BPP were identical to individuals assigned to lineages in the species tree analyses. The validity of our assignment of individuals to populations was tested using the guide-tree-free algorithm (A11) in BPP. Because the probability of delimitation by BPP is sensitive to selected parameters (Leaché and Fujita 2010; Yang 2015), we evaluated two independent runs for each of four different combinations of divergence depth and effective population sizes priors (τ and θ, respectively; Table 2). Two independent MCMC chains were run for 5 × 10⁴ generations. The burn-in was 20% and samples drawn every 50^th generation. In total, eight BPP runs were carried out using four phased nuclear intron alignments. Lineages were considered to be statistically well supported when the delimitation posterior probabilities generated were ≥0.95 under all four combinations of priors.

Table 2.

Primer information and chosen substitution models for regions amplified in this study. Substitution models before “/” are the best-supported models inferred by jMODELTEST2 and models after “/” indicate those inferred by PARTITIONFINDER2 for the concatenated intron alignment.

Primer name	Sequence	Primer publication	Substitution model
ACOX2-3-F	5’-CCTSGGCTCDGAGGAGCAGAT-3’	Salicini et al. 2011	K80+G / K81+G
ACOX2-3-R	5’-GGGCTGTGHAYCACAAACTCCT-3’
COPS7A-4-F	5’-TACAGCATYGGRCGRGACATCCA-3’	Salicini et al. 2011	HKY / K80
COPS7A-4-R	5’-TCACYTGCTCCTCRATGCCKGACA-3’
ROGDI-7-F	5’-CTGATGGAYGCYGTGATGCTGCA-3’	Salicini et al. 2011	K80+G / K81+G
ROGDI-7-R	5’-CACGGTGAGGCASAGCTTGTTGA-3’
STAT5B-16-F	5’--CTGCTCATCAACAAGCCCGA-3’	Matthee et al. 2001	GTR+G / K81+G
STAT5B-16-R	5’-GGCTTCAGGTTCCACAGGTTGC-3’
cyt-b-LGL-765-F	5’-GGCTTCAGGTTCCACAGGTTGC-3’	Trujillo et al. 2009	GTR+I+G
cyt-b -LGL-766-R	5’-GTTTAATTAGAATYTYAGCTTTGGG-3’

Table 3.

Prior Schemes (PS) used in BPP analyses. Prior distributions on τ represent two relative divergence depths (deep and shallow) and on θ represent two relative effective population sizes (large and small) scaling mutation rates.

PS	Effective pop. size	Divergence depth	Gamma distribution for prior
1	Large	Deep	θ = Γ [1, 10] and τ = Γ [1, 10]
2	Large	Shallow	θ = Γ [1, 10] and τ = Γ [2, 2000]
3	Small	Shallow	θ = Γ [2, 2000] and τ = Γ [2, 2000]
4	Small	Deep	θ = Γ [2, 2000] and τ = Γ [1, 10]

Results

In terms of cyt-b sequence divergence, clades within Doryrhina are separated by 3.0–5.7% genetic distances, whereas less than 3% separates the four recognized species of Macronycteris. Between Afrotropical Hipposideros, the greatest distances separate H. jonesi from other lineages (13.4–16.1%). The various numbered clades allied to Hipposideros caffer differ from one another in cyt-b sequences by 2.5–10.3% and clades allied to H. ruber differ by 3.0–8.2% (Table 4).

Table 4.

Uncorrected cyt-b p-distances between (off diagonal) and within (on diagonal) Afrotropical hipposiderid clades, showing the number of base differences per site averaged over all sequence pairs between groups. The analysis involved 386 nucleotide sequences and all ambiguous positions were removed; na (not available) reflects a sample size of one individual.

		[1]	[2]	[3]	[4]	[5]	[6]	[7]	[8]	[9]	[10]	[11]	[12]	[13]	[14]	[15]
[1]	Doryrhina camerunensis	0.003
[2]	Doryrhina cf. camerunensis	0.055	na
[3]	Doryrhina cyclops 1	0.057	0.048	0.008
[4]	Doryrhina cyclops 2	0.055	0.041	0.030	0.006
[5]	Hipposideros abae	0.176	0.173	0.168	0.166	0.033
[6]	Hipposideros beatus 1	0.152	0.156	0.160	0.152	0.116	0.007
[7]	Hipposideros beatus 2	0.146	0.149	0.151	0.147	0.117	0.044	0.006
[8]	Hipposideros caffer 1	0.157	0.154	0.150	0.148	0.108	0.103	0.106	0.006
[9]	Hipposideros caffer 2	0.153	0.150	0.150	0.147	0.106	0.096	0.108	0.045	0.01
[10]	Hipposideros caffer 3	0.152	0.148	0.150	0.149	0.108	0.110	0.111	0.046	0.052	0.011
[11]	Hipposideros caffer 4	0.162	0.159	0.160	0.154	0.106	0.105	0.114	0.077	0.078	0.079	0.018
[12]	Hipposideros caffer 5	0.150	0.155	0.159	0.148	0.113	0.091	0.096	0.095	0.101	0.103	0.098	0.005
[13]	Hipposideros caffer 6	0.155	0.156	0.164	0.153	0.112	0.093	0.102	0.090	0.097	0.099	0.094	0.028	0.011
[14]	Hipposideros caffer 7	0.151	0.154	0.161	0.148	0.108	0.084	0.094	0.094	0.094	0.098	0.096	0.025	0.032	0.011
[15]	Hipposideros caffer 8	0.154	0.155	0.160	0.152	0.111	0.090	0.092	0.092	0.095	0.102	0.093	0.033	0.039	0.029	0.021
[16]	Hipposideros fuliginosus	0.155	0.149	0.154	0.142	0.101	0.095	0.096	0.078	0.084	0.087	0.094	0.088	0.086	0.080	0.085
[17]	Hipposideros jonesi	0.153	0.143	0.154	0.147	0.160	0.145	0.138	0.135	0.140	0.140	0.139	0.134	0.134	0.137	0.139
[18]	Hipposideros lamottei	0.171	0.173	0.174	0.163	0.106	0.097	0.118	0.094	0.086	0.095	0.097	0.057	0.059	0.052	0.060
[19]	Hipposideros cf. lamottei	0.158	0.158	0.156	0.149	0.107	0.103	0.107	0.091	0.099	0.095	0.097	0.053	0.058	0.052	0.054
[20]	Hipposideros marisae	0.177	0.180	0.178	0.172	0.159	0.153	0.153	0.144	0.148	0.140	0.152	0.148	0.144	0.154	0.159
[21]	Hipposideros ruber 1	0.155	0.151	0.152	0.142	0.104	0.099	0.101	0.081	0.084	0.085	0.086	0.090	0.081	0.082	0.082
[22]	Hipposideros ruber 2	0.155	0.155	0.156	0.142	0.103	0.097	0.102	0.081	0.082	0.089	0.089	0.086	0.078	0.083	0.081
[23]	Hipposideros ruber 3	0.159	0.149	0.155	0.147	0.105	0.094	0.101	0.084	0.082	0.096	0.088	0.090	0.083	0.082	0.082
[24]	Hipposideros ruber 4	0.153	0.147	0.151	0.141	0.100	0.094	0.097	0.073	0.073	0.085	0.082	0.083	0.077	0.078	0.078
[25]	Hipposideros cf. ruber	0.164	0.161	0.161	0.153	0.099	0.094	0.100	0.086	0.084	0.095	0.093	0.081	0.080	0.080	0.083
[26]	Macronycteris commersoni	0.157	0.156	0.155	0.145	0.164	0.159	0.154	0.164	0.169	0.168	0.171	0.166	0.170	0.167	0.162
[27]	Macronycteris cryptovalorona	0.147	0.143	0.147	0.142	0.157	0.152	0.146	0.156	0.162	0.161	0.165	0.159	0.164	0.158	0.157
[28]	Macronycteris gigas	0.151	0.149	0.154	0.144	0.164	0.163	0.158	0.162	0.168	0.165	0.170	0.165	0.171	0.164	0.165
[29]	Macronycteris vittata	0.147	0.148	0.149	0.140	0.162	0.158	0.147	0.160	0.161	0.166	0.165	0.164	0.169	0.164	0.163

Table 4.

Continued.

		[16]	[17]	[18]	[19]	[20]	[21]	[22]	[23]	[24]	[25]	[26]	[27]	[28]	[29]
[16]	Hipposideros fuliginosus	0.003
[17]	Hipposideros jonesi	0.137	0.008
[18]	Hipposideros lamottei	0.100	0.161	0.038
[19]	Hipposideros cf. lamottei	0.091	0.142	0.054	0.006
[20]	Hipposideros marisae	0.145	0.092	0.156	0.158	na
[21]	Hipposideros ruber 1	0.082	0.144	0.093	0.087	0.152	0.013
[22]	Hipposideros ruber 2	0.084	0.147	0.093	0.091	0.157	0.030	0.007
[23]	Hipposideros ruber 3	0.082	0.142	0.085	0.088	0.158	0.052	0.057	0.022
[24]	Hipposideros ruber 4	0.078	0.140	0.087	0.088	0.154	0.053	0.051	0.057	na
[25]	Hipposideros cf. ruber	0.084	0.142	0.094	0.089	0.157	0.081	0.081	0.082	0.072	0.033
[26]	Macronycteris commersoni	0.155	0.186	0.183	0.167	0.193	0.163	0.162	0.161	0.168	0.174	0.012
[27]	Macronycteris cryptovalorona	0.149	0.176	0.170	0.163	0.185	0.161	0.161	0.158	0.160	0.162	0.028	0.003
[28]	Macronycteris gigas	0.153	0.179	0.178	0.169	0.190	0.165	0.165	0.161	0.165	0.169	0.026	0.029	0.012
[29]	Macronycteris vittata	0.150	0.181	0.180	0.168	0.192	0.158	0.159	0.158	0.159	0.164	0.026	0.029	0.027	0.006

Maximum likelihood and Bayesian phylogenies from a 452-individual alignment of cyt-b are shown in Suppl. material 2: Figures S2, Suppl. material 3: Figures S3. Identical haplotypes were pruned from this tree to produce the 303 unique-haplotype alignment shown in Figure 2. The 303 haplotype alignment used in the ML and BI gene tree analyses ranged from 413 to 1140 base pairs (bp) in length (89.9% complete matrix). Only the Bayesian topology is shown, but both posterior probabilities and bootstrap values are depicted at common, well supported nodes. Multiple, geographically cohesive clades are evident for the three widely distributed Afrotropical Hippposideros, H. beatus, H. caffer, and H. ruber.

Figure 2.

Parts A and B. Phylogeny of Hipposideridae based on Bayesian analysis of 303 cyt-b sequences. Colored lines denote well supported clades and symbols denote nodal support: red circles, BS ≥ 70%, PP ≥ 0.95; black circles BS ≥ 70%, PP ≤ 0.95; open circles BS ≤ 70%, PP ≥ 0.95.

Figure 2.

Continued.

Substitution networks for cyt-b haplotypes for Doryrhina, Macronycteris, and Hipposideros are shown in Figures 3, 4, showing the genetic and geographic relationships of the clades identified in Figure 2.

Figure 3.

Substitution network plots for Afrotropical hipposiderids ADoryrhinaBMacronycteris.

Figure 4.

Substitution network plots for Afrotropical hipposiderids AHipposideros caffer clades 1–4 BHipposideros caffer clades 5–8 CH. ruber clades.

Maximum likelihood and Bayesian phylogenies from a 103-individual alignment of four concatenated introns for Doryrhina, Macronycteris, and Hipposideros are shown in Figure 5. Many of the numbered clades in Figures 2–4 are jumbled in Figure 5; they are not recovered as monophyletic units and the geographic structure evident in mtDNA analyses disappears.

Figure 5.

Phylogeny of Hipposideridae based on Bayesian analysis of 103 concatenated nuclear intron sequences. Numbers denote posterior probabilities (BI) and bootstrap percentages (ML); red circles at more terminal nodes indicate BS ≥ 70%, PP ≥ 0.95.

A species tree generated using StarBEAST from the four introns appears in Figure 6. It depicts well-supported relationships among the various clades allied with H. caffer, H. ruber, and H. beatus. Remarkably, and in contrast with the concatenated analyses, it shows support for the Asian dyad H. diadema and H. larvatus as sister to these ruber subgroup members, with the Asian ater subgroup outside this pairing. There is little support for the deeper phylogenetic nodes.

Figure 6.

Species tree Hipposideridae based on StarBEAST analysis of four introns. Posterior probabilities appear at all nodes.

Discussion

Overall genetic variability

The three Afrotropical hipposiderid genera differ substantially in terms of their internal genetic differentiation. Clades of Hipposideros are separated by cyt-b p-distances averaging 9.7% (2.5–16.1%), whereas Doryrhina clades average p-distances of 4.8% (3.0–5.7%) and Macronycteris clades 2.7% (2.6–2.9%). Distance values for these genera tend to fall at the lower end of values obtained with similar sampling intensity for species-ranked clades in other Afrotropical bat genera: 2.5% for Otomops (Patterson et al. 2018), 9.3% for Miniopterus (Demos et al. 2020), 10% for Scotophilus and Rhinolophus (Demos et al. 2018, 2019a), 13.5% for Myotis (Patterson et al. 2019), and 17% for Nycteris (Demos et al. 2019b). Fewer cyt-b substitutions on average for these hipposiderids does not limit support for individual clades, and because distances do not approach those characteristic of substitutional saturation, the cyt-b tree recovers much of the deeper phylogenetic structure evident with nuclear intron sequences (compare Figs 2, 5).

Phylogenetics

Both cyt-b and intron analyses securely recovered Doryrhina, Macronycteris, and Hipposideros as monophyletic. Doryrhina + Macronycteris are sister to the remaining hipposiderids. However, only the cyt-b analysis included the hipposiderid genera Aselliscus, Coelops, and Asellia alongside Hipposideros. That analysis recovered all four genera as monophyletic with strong support. Aselliscus and Coelops were recovered as sister to Hipposideros, with Asellia joining later, but these relationships lacked confident support.

Using a supermatrix approach on exemplars of 46 species of hipposiderids, Amador et al. (2018) found Hipposideros sensu stricto to be paraphyletic. They recovered a mostly Asian group of Hipposideros as sister to two subclades, Coelops + Aselliscus and Asellia + African hipposiderids excluding H. jonesi, which was recovered with the Asian taxa. Paraphyly in this molecular analysis echoed earlier indications of Hipposideros paraphyly from morphology (Bogdanowicz and Owen 1998; Hand and Kirsch 1998, 2003). In another supermatrix analysis of exemplars belonging to 49 hipposiderid species, Shi and Rabosky (2015) failed to recover Macronycteris as monophyletic; M. commersoni was sister to all remaining hipposiderids, but strangely it did not group with M. gigas. When the anomalous position of M. commersoni in their tree is ignored, their topology is highly similar to that of Figure 2, except that Asellia (Aselliscus, Coelops) become the sister of Hipposideros (Macronycteris, Doryrhina), rather than sister of just Hipposideros. Using both mitochondrial and nuclear loci, Lavery et al. (2014) found that 17 species of Asian, Oceanian and Australasian Hipposideros were monophyletic with respect to the genera Aselliscus, Coelops, and Anthops. Clearly, missing data and missing taxa compromise all of these phylogenetic appraisals, so that the question of hipposiderid and Hipposideros monophyly remains open. However, subject to its sampling limitations, there is clear support in our analyses of monophyly for Doryrhina, Macronycteris, and Hipposideros as we apply these names.

Despite employing different mitochondrial and nuclear loci and using different sets of taxa, the phylogeny recovered by Lavery et al. (2014) is largely congruent with that in Figure 5. Their earliest diverging species group of Hipposideros is the calcaratus group, not represented in our tree unless H. obscurus is a member (Table 1). Their next diverging unit is the diadema group, which is also positioned near the base of our tree. Their other two groups are paired: the galeritus group (which includes H. cervinus, indicating that this species is misclassified as calcaratus member) joined with the bicolor/ater group. In our intron analysis (Fig. 5), members of the larvatus and diadema groups join H. obscurus as sister to all remaining Hipposideros groups. The remainder form a trichotomy: H. coronatus, typically considered in the bicolor group; H. pygmaeus and H. cervinus, which are listed in different groups but were both considered members of the galeritus unit by Tate (1941); and the erstwhile bicolor group (sensu Hill 1963), which was subdivided into the ater subgroup (for Asian, Oceanian, and Australasian species) and the ruber subgroup (for Atrotropical ones) by Monadjem (2019).

The ater subgroup members included in our mitochondrial analysis (Fig. 2) form a well-supported clade consisting of H. bicolor, H. cineraceus, H. pomona, H. doriae, H. ater, H. khaokhouayensis, H. rotalis, H. halophyllusH. dyacorum, H. ridley, and H. durgadasi. This group is sister to all analyzed members of the ruber subgroup: the various clades allied with Hipposideros beatus, H. caffer, and H. ruber, as well as individuals of the Afrotropical species H. lamottei and H. fuliginosus. H. abae, which was previously considered in the speoris group (Simmons 2005; Murray et al. 2012), is clearly a member of the ruber group. Outside this pairing are the Asian species H. cervinus, H. coronatus, H. coxi, H. obscurus, and H. pygmaeus. Two Afrotropical species also lie outside the ruber + ater clade: H. jonesi and H. marisae, both thought to belong to the speoris group, appear as sisters in Figure 2A.

Parsimony, topological position, and the strong support of branching relationships in the mitochondrial and intron trees (Fig. 5; also Lavery et al. 2014) make it clear that the Afrotropical ruber group represents a comparatively recent colonization event from Asian ancestors–the ruber group is sister to the ater group and this pair has Asian sisters. However, although the basal dichotomy within Hipposideros includes an all Asian clade, lack of support for its sister(s) clouds the phylogenetic position of the H. jonesi-H. marisae clade–possibly sister to all sampled Hipposideros but more likely sandwiched between Asian clades. In any case, Figure 2 suggests that the H. jonesi-H. marisae clade resulted from an earlier African-Asian colonization event.

The lack of agreement in the phylogenetic position of H. diadema and H. larvatus between the concatenated intron tree (Fig. 5) and the species tree (Fig. 6) deserves comment, as both analyses were based on the same genetic dataset. The position of H. diadema-H. larvatus as sister to the ruber group (Fig. 6) runs counter to both our other genetic analyses (Figs 2, 5) and morphological assessments (Hill 1963; Murray et al. 2012; Table 1). This discrepancy is likely due to the generally weaker support for deep nodes within the tree; in the absence of saturation, this is often taken as evidence of rapid evolutionary radiations (e.g., Almeida et al. 2011). Lanier and Knowles (2014) used simulated data on deep phylogenies to show that species-tree methods do account for coalescent variance at deep nodes but that mutational variance among lineages poses the primary challenge for accurate reconstruction. In either case, vastly expanded genetic sampling via NGS techniques offers the most plausible avenues to clearer resolution.

However, the highly distinctive species H. megalotis belongs to its own species group (Table 1) and has not been included in any genetic analysis. Distributed in the Horn of Africa and the Arabian Peninsula, H. megalotis is the only hipposiderid with a fold of skin joining the base of the ear pinnae. Its uniquely specialized auditory system and derived dentition (e.g., loss of anterior premolars and enlargement of outer lower incisors), led Hill (1963) to regard it as a species that diverged early from the other groups of African Hipposideros. Including this species in future analyses would shed light on the group’s biogeography. Were there three colonizations of Africa by Asian groups of Hipposideros or could H. megalotis be sister to all Asian lineages of this genus? This information would greatly clarify ancestral geographic range inference.

Species limits

The lineage delimitation analyses indicate that a number of hipposiderid lineages are either unnamed or unidentified, and also that a number of recognized species may not be genetically and evolutionarily independent.

Previous studies had indicated that both Hipposideros caffer (Vallo et al. 2008) and H. ruber (Vallo et al. 2011) appear to be complexes of cryptic species. The two are traditionally distinguished on the basis of size and pelage color, H. ruber being the larger and more brightly colored form, but this distinction is clouded by geographic variation in size and the presence of both reddish and gray-brown phases in both species. Our mitochondrial analyses identified four H. ruber lineages and eight H. caffer lineages in two distinct groupings among the sampled populations (Fig. 2). Four of the caffer lineages and three of the ruber clades were identified as putative species by the BPP analyses (Table 5). The large number of clades in East Africa is remarkable: Kenya and Tanzania each support four of the eight clades allied with H. caffer, and all but one of the eight clades known from throughout the continent occur in one or the other East African country. This undoubtedly reflects the region’s great landscape diversity, where West and Central African rainforests reach their eastern limit, southern savannas reach their northern limits, the Sahel reaches its southern limits, and all are riven by the African Rift Valley. It also is a product of our sampling intensity (see Suppl. material 1: Fig. S1).

Table 5.

Lineage delimitation results from BPP based on the four intron dataset for mtDNA-supported clades of Afrotropical Hipposideridae. PS1-PS4 refer to four different prior schemes based on population size and age of divergence priors (see Table 3 for parameter details). Bold font indicates that the putative species was delimited under all parameter settings.

Putative Species	PS1	PS2	PS3	PS4
Doryrhina camerunensis	0.30	0.76	0.95	0.51
D. cf. camerunensis	0.32	0.73	0.97	0.79
D. cyclops 2	0.23	0.68	0.95	0.51
Hipposideros beatus 2	1	1	1	1
H. caffer 1	0.99	0.99	0.99	0.99
H. caffer 2	0.99	1	1	1
H. caffer 3	0.99	0.99	0.99	0.99
H. caffer 5	0.14	0.18	0.11	0.08
H. caffer 6	0.56	0.61	0.85	0.82
H. caffer 7	0.14	0.18	0.11	0.08
H. caffer 8	0.99	0.99	0.99	0.99
H. ruber 1	0.99	0.99	0.99	0.99
H. ruber 2	1	0.99	1	0.99
H. ruber 4	0.99	0.99	0.99	0.99
Macronycteris commersoni	0.24	0.72	0.94	0.52
M. cryptovalorona	0.35	0.81	0.97	0.76
M. gigas	0.09	0.43	0.91	0.38
M. vittata	0.11	0.44	0.90	0.34

Because some cyt-b sequences were used in multiple studies of this group, it is possible to relate our clade labels to those used by earlier studies (Table 6). Based on attributions made on morphological grounds by Vallo et al. (2008) and Monadjem et al. (2013), some well-supported but unnamed clades in our analysis can be identified. For instance, caffer1 has a distributional range and includes specimens previously identified as Hipposideros tephrus (Appendix I), while specimens of caffer4 come from near the type locality of H. caffer Sundevall, 1846, and may well represent that species. However, no samples confidently identified as H. ruber from the vicinity of its type locality have been sequenced, leaving the application of that name to clades in any of these trees purely conjectural. Applying formal names only after integrative taxonomic assessment is a responsible course as multispecies coalescent models like BPP can lead to over-splitting of species, especially when applied to geographically variable species complexes with parapatric distributions (Chambers and Hillis 2020).

Table 6.

Clade names and associated binomials (if used) for three analyses of cryptic lineages within the ruber species group of Hipposideros. No genetic analysis of this group has included type material; consequently, the application of binomials hinges on the robustness of ancillary morphological analyses, which were not conducted in our study. Boldfaced names denote clades supported by all four prior schemes in our BPP delimitation analyses.

Vallo et al. (2008)		Monadjem et al. (2013)		This paper
A1	H. caffer			caffer 4
		A1a	H. caffer	caffer 4
		A1b	H. caffer	caffer 4
A2	H. caffer		H. caffer tephrus	caffer 1
B	H. ruber			caffer 5
		B1	H. ruber	cf. lamottei
		B2	H. ruber	caffer 7
C1	H. ruber			ruber 1, ruber 2
		C1a	H. cf. ruber	ruber 1
		C1b	H. cf. ruber	ruber 1
C2	H. ruber	C2	H. cf. ruber	ruber 3
				ruber 4
D	H. ruber		H. cf. ruber	cf. ruber
		D1	H. cf. ruber	cf. ruber
		D2	H. cf. ruber	cf. ruber
–		E1	H. cf. ruber	cf. ruber
–		E2	H. cf. ruber	cf. ruber
–		–		caffer 2
–		–		caffer 3
–		–		caffer 6
–		–		caffer 8
abae	H. abae	abae	H. abae	abae
beatus	H. beatus	beatus	H. beatus	beatus1, beatus 2
fuliginosus	H. fuliginosus	fuliginosus	H. fuliginosus	fuliginosus
lamottei	H. lamottei	lamottei	H. lamottei	lamottei

Doryrhina is a poorly known genus characterized morphologically by the peculiar club-shaped processes on the central and posterior nose leaves. This trait is shared by the two recognized African species, D. cyclops and D. camerunensis, which differ chiefly in size (the latter is larger, with forearm lengths >75 mm). Although D. cyclops is considered to be monotypic, mitochondrial sequences clearly separate West African populations in Liberia and Senegal (cyclops1) from Central African populations in Gabon and Central African Republic (cyclops2), and these are substantially separated from D. camerunensis and a specimen referred to that species from Tanzania (Figs 2, 3). However, both the intron analysis (Fig. 5) and the species tree (Fig. 6) show little or no geographic structure. The BPP analyses confirm that none of the mitochondrial clades is behaving as an independent evolutionary lineage (Table 5). Geographic structure in mtDNA but continent-wide admixture in the nuclear genome could result from either male-biased dispersal with female philopatry or highly structured seasonal migrations, which are known in other hipposiderids. In any case, the genetic patterns of Doryrhina are hard to reconcile with its space-use behavior; individuals appear to have very small home ranges, on the order of a few hectares (Monadjem 2019). An integrative taxonomic review of the genus Doryrhina is needed to determine the validity of D. cyclops and D. camerunensis. It would also shed light on whether six Australo-Papuan species tentatively allocated to that genus (cf. Monadjem 2019) belong there or elsewhere. Tate (1941) had earlier allocated those species to the Australasian muscinus group, convergent on but separate from his Afrotropical cyclops group, but Hill (1963) later united these groups.

Our analysis included four of the five recognized species of Macronycteris, lacking only M. thomensis, which is endemic to São Tomé Island in the Gulf of Guinea. Two species, M. gigas and M. vittata, occur on the African mainland and two others, M. commersoni and M. cryptovalorona, occur on Madagascar. Macronycteris cryptovalorona was named only in 2016, on the basis of its strong genetic divergence from M. commersoni; it appears in Figure 2 as sister to all three remaining species of Macronycteris. Despite a search for diagnostic characters, Goodman et al. (2016) could not distinguish it morphologically from M. commersoni. Both species are known to occur in the same caves in south central and southwestern Madagascar (Goodman et al. 2016; Rakotoarivelo et al. 2019). On the other hand, M. vittata and M. gigas are distinguished typically on the basis of size and pelage color (cf. Monadjem 2019). They are also known to occur together in the same cave (Shimoni Cave in Kwale, Kenya; Webala et al. 2019), where they utilize echolocation calls with different peak frequencies: vittata at 64–70 kHz and gigas at 53.4–54.8 kHz. Both in Africa and on Madagascar, these pairs of taxa appear to act as distinct species, but the monophyly evident in the cyt-b sequences (Figs 2, 4) disappears in the nuclear intron analyses. BPP analyses fail to resolve any of the Macronycteris species, and none appear as monophyletic in the concatenated intron analyses.

Our results clearly underscore the importance of using multilocus datasets to evaluate phylogenetic and phylogeographic relationships at the genus and species level in mammals. Use of a single genetic system may lead to widely divergent conclusions regarding species identity and distribution. Toews and Brelsford (2012) reviewed cases of mito-nuclear discordance in animals generally. Fully 18% of the cases they reviewed had discordant patterns of mitochondrial and nuclear DNA. In most cases, such patterns are attributable to adaptive introgression of mtDNA, demographic disparities, and sex-biased asymmetries; in some cases they found evidence for hybrid zone movement or human agency. Discordant patterns of variation between mitochondrial and nuclear DNA have been reported in at least six other families of bats (Nesi et al. 2011; Furman et al. 2014; Naidoo et al. 2016; Hassanin et al. 2018; Demos et al. 2019a; Gürün et al. 2019). Gürün et al. (2019) implicated the role of sex-biased dispersal in causing such discordance, male dispersal spreading nuclear variation farther and faster than the movement of mitochondria. This may be a more general pattern in bats (see also Demos et al. 2019b). To understand the processes responsible for these discordant patterns of genome evolution, extensive genomic sampling and far fuller knowledge of natural history will be required.

Appendix I

Genetic sampling of Hipposideridae. Wherever possible, the voucher numbers associated with the genetic samples are specified. Accession numbers identify sequences downloaded from GenBank or accessioned to Genbank for this study. The designation 'redundant' indicates a cyt-b sequence that was omitted from the 303 individual alignment because of its identity to another. Institutional acronyms are as follows: AMNH – American Museum of Natural History, New York; CM – Carnegie Museum, Pittsburgh; CVVD – ?; DM – Durban Natural Science Museum, Durban; EBD – Estación Biológica de Doñana, Sevilla; FMNH – Field Museum of Natural History, Chicago; IEBR-T – Thong collection at Institute of Ecology and Biological Resources, Hanoi; IVB – Institute of Vertebrate Biology, Brno; KU – Biodiversity Institute and Natural History Museum, University of Kansas, Lawrence; Czech Academy of Sciences, Prague; LSUMZ – Lousiana State University Museum of Natural Science-Mammal Tissues, Baton Rouge; NHMOU – Natural History Museum of Osmania University, Hyderabad; NMK – National Museums of Kenya, Nairobi; NMP – National Museum, Prague; PSUZC – Princess Maha Chakri Sirindhorn Natural History Museum, Songkhla; ROM – Royal Ontario Museum, Toronto; SMF – Senckenberg Museum, Frankfurt; TM – Transvaal Museum, Pretoria; TTU – Texas Tech University Museum, Lubbock; UADBA – Université d'Antananarivo, Département de Biologie Animale, Antananarivo; UNIMAS – University of Malaysia Sarawak Natural History Museum, Kuching.

Voucher	cyt-b	ACOX2	COPS	ROGDI	STAT5	ScientificName	Country	Latitude	Longitude
	KU316954					Asellia arabica	Oman	17.100	54.080
	KU316958					Asellia italosomalica	Yemen	12.670	54.120
	JF438999					Asellia tridens	Libya	24.933	10.167
IEBR-T	KU161572					Aselliscus dongbacana	Vietnam	22.360	105.395
	LC426460					Aselliscus stoliczkanus	China
	DQ888675					Aselliscus tricuspidatus	Vanuatu	-15.307	166.926
DM 8021	FJ457616					Cloeotis percivali	Swaziland	-25.817	31.283
DM 8026	FJ457615					Cloeotis percivali	Swaziland	-25.817	31.283
	DQ888674					Coelops frithi	Taiwan	21.948	120.780
FMNH 148981	redundant					Doryrhina camerunensis	Burundi	-2.100	29.383
FMNH 148982	MT149719	MT149615	MT149513	MT149418	MT149317	Doryrhina camerunensis	Burundi	-2.850	29.400
NMK 187403	redundant	MT149616	MT149514	MT149419	MT149318	Doryrhina camerunensis	Kenya	0.344	34.857
NMK 187418	MT149720					Doryrhina camerunensis	Kenya	0.344	34.857
FMNH 165159	MT149721					Doryrhina camerunensis	Uganda	1.683	31.533
FMNH 165160	MT149722					Doryrhina camerunensis	Uganda	1.683	31.533
FMNH 223198	MT149723					Doryrhina camerunensis	Uganda	0.445	32.889
FMNH 223551	redundant	MT149617	MT149515	MT149420	MT149319	Doryrhina camerunensis	Uganda	0.445	32.889
FMNH 224066	MT149724					Doryrhina camerunensis	Uganda	0.501	30.426
FMNH 224068	MT149725					Doryrhina camerunensis	Uganda	0.501	30.426
FMNH 153929	MT149726	MT149618	MT149516	MT149421	MT149320	Doryrhina cf. camerunensis	Tanzania	-4.942	38.733
DM 12626	KF551833					Doryrhina cyclops1	Liberia	7.553	-8.492
IVB S261	EU934465					Doryrhina cyclops1	Senegal	12.883	-12.717
IVB S747	EU934466					Doryrhina cyclops1	Senegal	13.333	-13.217
FMNH 227409	MT149727	MT149619	MT149517	MT149422	MT149321	Doryrhina cyclops2	Central African Republic	13.033	16.410
FMNH 227410	MT149728	MT149620	MT149518	MT149423	MT149322	Doryrhina cyclops2	Central African Republic	3.033	16.410
FMNH 167772	MT149729	MT149621	MT149519	MT149424	MT149323	Doryrhina cyclops2	Gabon	-2.283	10.497
FMNH 167773	MT149730	MT149622	MT149520	MT149425	MT149324	Doryrhina cyclops2	Gabon	-2.283	10.497
NMP 91850	EU934446					Hipposideros abae	Benin	7.783	2.267
NMP 91851	EU934447					Hipposideros abae	Benin	7.783	2.267
IVB S822	EU934448					Hipposideros abae	Senegal	12.350	-12.317
IEBR-T 90806.7	JN247006					Hipposideros alongensis	Vietnam
YN07C123	JX849159					Hipposideros armiger	China	23.600	102.002
UNIMAS 729	EF108140 redundant					Hipposideros ater	Malaysia	1.407	110.169
UNIMAS 1577	EF108139 redundant					Hipposideros ater	Malaysia	3.801	113.785
KU 164242	MT149731	MT149623	MT149521	MT149426	MT149325	Hipposideros ater	Philippines	13.796	120.159
KU 164243	MT149732	MT149624	MT149522	MT149427	MT149326	Hipposideros ater	Philippines	13.796	120.159
KU 164712	MT149733	MT149625	MT149523	MT149428	MT149327	Hipposideros ater	Philippines	19.085	121.241
ROM 100579	FJ347975					Hipposideros beatus1	Ivory Coast	6.930	-7.217
DM 13241	KF551829					Hipposideros beatus1	Liberia	7.553	-8.492
DM 13242	KF551830					Hipposideros beatus1	Liberia	7.553	-8.492
FMNH 227406	MT149734	MT149613	MT149524	MT149429	MT149328	Hipposideros beatus2	Central African Republic	3.033	16.410
FMNH 149406	FJ347976					Hipposideros beatus2	Democratic Republic of Congo	-1.417	28.583
FMNH 215440	MT149735	MT149626	MT149525	MT149430	MT149329	Hipposideros beatus2	Kenya	0.352	34.865
NMK 184861	MT149736	MT149627	MT149526	MT149431	MT149330	Hipposideros beatus2	Kenya	0.356	34.861
NMK 184864	redundant					Hipposideros beatus2	Kenya	0.360	34.861
NMK 184870	MT149737					Hipposideros beatus2	Kenya	0.352	34.865
FMNH 192931	MT149738					Hipposideros beatus2	Tanzania	-1.094	31.515
FMNH 192932	MT149739					Hipposideros beatus2	Tanzania	-1.094	31.515
FMNH 192933	redundant	MT149628	MT149527	MT149432	MT149331	Hipposideros beatus2	Tanzania	-1.094	31.515
FMNH 164972	MT149740					Hipposideros beatus2	Uganda	1.733	31.467
FMNH 165157	redundant	MT149629	MT149528	MT149433	MT149332	Hipposideros beatus2	Uganda	1.683	31.533
LSUMZ MT-4482	MT149741	MT149630	MT149529	MT149434	MT149333	Hipposideros bicolor	Malaysia	1.970	103.500
LSUMZ MT-4489	MT149742	MT149631	MT149530		MT149334	Hipposideros bicolor	Malaysia	1.970	103.500
FMNH 215441	redundant	MT149632	MT149531	MT149435	MT149335	Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215442	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215443	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215444	MT149743					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215445	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215446	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 215447	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216628	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216629	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216630	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216631	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216632	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216645	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216646	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216647	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 216648	redundant	MT149633	MT149532	MT149436	MT149336	Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 225346	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
FMNH 225747	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
FMNH 225748	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
FMNH 225749	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
FMNH 225750	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
FMNH 225751	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184726	MT149744					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184727	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184728	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184729	redundant					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184730	MT149745					Hipposideros caffer1	Kenya	-0.564	36.254
NMK 184760	redundant					Hipposideros caffer1	Kenya	-0.430	36.174
NMK 184842	redundant					Hipposideros caffer1	Kenya	-0.539	36.294
NMK 184843	redundant					Hipposideros caffer1	Kenya	-0.539	36.294
NMK 187310	MT149718					Hipposideros caffer1	Kenya	-0.539	36.294
NMK 187311	redundant					Hipposideros caffer1	Kenya	-0.539	36.294
NMK 187312	redundant					Hipposideros caffer1	Kenya	-0.539	36.294
NMK 187323	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
NMK 187324	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
NMK 187325	redundant	MT149634	MT149533	MT149437	MT149337	Hipposideros caffer1	Kenya	-0.346	36.119
NMK 187326	redundant					Hipposideros caffer1	Kenya	-0.346	36.119
EBD 23262	FJ347977					Hipposideros caffer1	Morocco	30.630	9.830
NMP	EU934449					Hipposideros caffer1	Morocco	3.801	113.785
FMNH 223196	MT149746	MT149635	MT149534	MT149438	MT149338	Hipposideros caffer1	Uganda	0.445	32.889
FMNH 223197	MT149747					Hipposideros caffer1	Uganda	0.445	32.889
NMP	EU934463					Hipposideros caffer1	Yemen	15.283	44.167
FMNH 220955	redundant	MT149639	MT149538	MT149441	MT149342	Hipposideros caffer2	Kenya	-2.203	37.714
FMNH 220956	redundant					Hipposideros caffer2	Kenya	-2.203	37.714
FMNH 220957	MT149753					Hipposideros caffer2	Kenya	-2.203	37.714
FMNH 220958	redundant					Hipposideros caffer2	Kenya	-2.203	37.714
FMNH 225347	redundant	MT149640	MT149539	MT149442	MT149343	Hipposideros caffer2	Kenya	-1.547	35.306
FMNH 225348	redundant					Hipposideros caffer2	Kenya	-1.531	35.320
FMNH 225349	MT149754					Hipposideros caffer2	Kenya	-1.531	35.320
FMNH 225350	MT149755	MT149641	MT149540	MT149443		Hipposideros caffer2	Kenya	-1.531	35.320
FMNH 225351	MT149756					Hipposideros caffer2	Kenya	-1.531	35.320
FMNH 225352	redundant					Hipposideros caffer2	Kenya	-1.531	35.320
NMK 184977	MT149749					Hipposideros caffer2	Kenya	-0.117	34.541
NMK 184978	MT149750	MT149637	MT149536	MT149440	MT149340	Hipposideros caffer2	Kenya	-0.117	34.541
NMK 184979	MT149751					Hipposideros caffer2	Kenya	-0.117	34.541
NMK 184981	redundant					Hipposideros caffer2	Kenya	-0.117	34.541
NMK 184982	MT149752	MT149638	MT149537		MT149341	Hipposideros caffer2	Kenya	-0.117	34.541
NMK 184999	MT149748	MT149636	MT149535	MT149439	MT149339	Hipposideros caffer2	Kenya	-0.555	37.388
FMNH 215914	MT149768					Hipposideros caffer3	Kenya	-3.706	38.776
FMNH 215915	redundant					Hipposideros caffer3	Kenya	-3.706	38.776
FMNH 215916	MT149769					Hipposideros caffer3	Kenya	-3.706	38.776
FMNH 215917	redundant	MT149646	MT149545	MT149448	MT149348	Hipposideros caffer3	Kenya	-3.706	38.776
FMNH 215918	redundant					Hipposideros caffer3	Kenya	-3.706	38.776
FMNH 215921	redundant					Hipposideros caffer3	Kenya	-3.076	39.217
FMNH 215922	MT149770					Hipposideros caffer3	Kenya	-3.076	39.217
FMNH 215923	MT149771					Hipposideros caffer3	Kenya	-3.076	39.217
FMNH 215924	redundant					Hipposideros caffer3	Kenya	-3.076	39.217
FMNH 215925	MT149772	MT149647	MT149546	MT149449	MT149349	Hipposideros caffer3	Kenya	-3.076	39.217
FMNH 220648	MT149766					Hipposideros caffer3	Kenya	0.170	38.194
FMNH 220669	MT149767	MT149645	MT149544	MT149447	MT149347	Hipposideros caffer3	Kenya	0.024	38.066
FMNH 234022	MT149757					Hipposideros caffer3	Kenya	-1.019	38.326
FMNH 234023	MT149758					Hipposideros caffer3	Kenya	-0.992	38.330
NMK 184226	MT149762	MT149644	MT149543	MT149446	MT149346	Hipposideros caffer3	Kenya	2.320	37.994
NMK 184238	MT149763					Hipposideros caffer3	Kenya	2.320	37.994
NMK 184284	MT149764					Hipposideros caffer3	Kenya	2.320	37.994
NMK 184287	MT149765					Hipposideros caffer3	Kenya	2.283	37.954
NMK 184425	MT149761	MT149643	MT149542	MT149445	MT149345	Hipposideros caffer3	Kenya	0.228	37.113
NMK 185050	redundant	MT149642	MT149541	MT149444	MT149344	Hipposideros caffer3	Kenya	-0.992	38.330
NMK 185051	redundant					Hipposideros caffer3	Kenya	-0.992	38.330
NMK 185052	MT149759					Hipposideros caffer3	Kenya	-0.992	38.330
NMK 185053	redundant					Hipposideros caffer3	Kenya	-0.992	38.330
NMK 185054	MT149760					Hipposideros caffer3	Kenya	-0.992	38.330
DM 8587	KF551805					Hipposideros caffer4	Mozambique	-23.205	32.499
DM 8590	KF551810					Hipposideros caffer4	Mozambique	-12.182	37.550
TM 48051	EU934451					Hipposideros caffer4	Mozambique	-21.517	35.100
DM 11007	KF551806 redundant					Hipposideros caffer4	South Africa	-27.596	32.220
	FJ347979					Hipposideros caffer4	South Africa	-27.660	32.251
	EU934452					Hipposideros caffer4	South Africa	-23.999	31.645
DM 7920	EU934458					Hipposideros caffer4	Swaziland	-26.870	31.463
FMNH 215941	redundant					Hipposideros caffer5	Kenya	-3.300	39.995
FMNH 220176	MT149780					Hipposideros caffer5	Kenya	-4.590	39.331
FMNH 220177	MT149781					Hipposideros caffer5	Kenya	-4.590	39.331
FMNH 220178	redundant					Hipposideros caffer5	Kenya	-4.590	39.331
FMNH 220179	MT149782					Hipposideros caffer5	Kenya	-4.590	39.331
FMNH 220180	MT149783					Hipposideros caffer5	Kenya	-4.590	39.331
FMNH 220182	MT149784					Hipposideros caffer5	Kenya	-4.082	39.483
FMNH 220183	MT149785					Hipposideros caffer5	Kenya	-4.082	39.483
FMNH 220184	MT149786					Hipposideros caffer5	Kenya	-4.082	39.483
FMNH 220185	redundant					Hipposideros caffer5	Kenya	-4.082	39.483
FMNH 220186	redundant					Hipposideros caffer5	Kenya	-4.082	39.483
FMNH 220202	MT149773					Hipposideros caffer5	Kenya	-3.300	39.995
FMNH 220203	redundant					Hipposideros caffer5	Kenya	-3.300	39.995
FMNH 220204	redundant	MT149648	MT149547	MT149450	MT149350	Hipposideros caffer5	Kenya	-3.300	39.995
FMNH 220205	redundant					Hipposideros caffer5	Kenya	-3.323	40.042
FMNH 220206	MT149774					Hipposideros caffer5	Kenya	-3.323	40.042
FMNH 220207	redundant	MT149649	MT149548	MT149451	MT149351	Hipposideros caffer5	Kenya	-3.323	40.042
FMNH 220208	redundant	MT149650	MT149549	MT149452	MT149352	Hipposideros caffer5	Kenya	-3.323	40.042
FMNH 220209	MT149775					Hipposideros caffer5	Kenya	-3.323	40.042
FMNH 233985	redundant					Hipposideros caffer5	Kenya	-3.335	40.031
NMK 187199	MT149776					Hipposideros caffer5	Kenya	-3.323	40.042
NMK 187200	redundant					Hipposideros caffer5	Kenya	-3.323	40.042
NMK 187201	MT149777					Hipposideros caffer5	Kenya	-3.323	40.042
NMK 187202	MT149778					Hipposideros caffer5	Kenya	-3.323	40.042
NMK 187203	MT149779					Hipposideros caffer5	Kenya	-3.323	40.042
CM 97957	FJ347980					Hipposideros caffer5	Kenya	-4.250	39.383
FMNH 192789	MT149787	MT149651	MT149550	MT149453	MT149353	Hipposideros caffer5	Tanzania	-4.902	39.688
FMNH 192855	redundant	MT149652	MT149551	MT149454	MT149354	Hipposideros caffer5	Tanzania	-4.902	39.688
FMNH 187385	redundant	MT149653	MT149552	MT149455	MT149355	Hipposideros caffer6	Tanzania	-8.003	39.762
FMNH 187386	MT149788	MT149654	MT149553	MT149456		Hipposideros caffer6	Tanzania	-8.003	39.762
FMNH 187387	MT149789					Hipposideros caffer6	Tanzania	-7.993	39.792
FMNH 187388	redundant					Hipposideros caffer6	Tanzania	-7.993	39.792
FMNH 187417	MT149790					Hipposideros caffer6	Tanzania	-7.891	39.843
FMNH 187418	MT149791					Hipposideros caffer6	Tanzania	-7.891	39.843
FMNH 187426	MT149792					Hipposideros caffer6	Tanzania	-7.891	39.843
FMNH 187428	redundant					Hipposideros caffer6	Tanzania	-7.993	39.792
FMNH 198066	MT149793	MT149655	MT149554	MT149457	MT149356	Hipposideros caffer6	Tanzania	-5.878	39.311
FMNH 198067	MT149794					Hipposideros caffer6	Tanzania	-5.878	39.311
FMNH 198072	redundant	MT149656	MT149555	MT149458	MT149357	Hipposideros caffer6	Tanzania	-6.244	39.320
FMNH 198073	redundant					Hipposideros caffer6	Tanzania	-6.244	39.320
FMNH 198074	MT149795	MT149657	MT149556	MT149459	MT149358	Hipposideros caffer6	Tanzania	-6.244	39.320
FMNH 198075	MT149796					Hipposideros caffer6	Tanzania	-6.244	39.320
FMNH 198076	redundant					Hipposideros caffer6	Tanzania	-6.244	39.320
FMNH 198082	redundant					Hipposideros caffer6	Tanzania	-6.280	39.451
FMNH 198083	MT149797					Hipposideros caffer6	Tanzania	-6.280	39.451
FMNH 198084	MT149798					Hipposideros caffer6	Tanzania	-6.280	39.451
FMNH 198131	redundant					Hipposideros caffer6	Tanzania	-5.878	39.311
FMNH 198132	MT149799					Hipposideros caffer6	Tanzania	-5.878	39.311
FMNH 198133	MT149800					Hipposideros caffer6	Tanzania	-5.878	39.311
NMP	EU934460					Hipposideros caffer6	Tanzania	-5.998	39.187
NMP	EU934477					Hipposideros caffer7	Malawi	-16.033	35.500
DM 8528	KF551817					Hipposideros caffer7	Mozambique	-13.401	34.870
DM 8550	KF551816 redundant					Hipposideros caffer7	Mozambique	-13.401	34.870
FMNH 155554	MT149801	MT149658	MT149557	MT149460	MT149359	Hipposideros caffer7	Tanzania	-8.519	35.904
FMNH 192790	redundant	MT149659	MT149558	MT149461	MT149360	Hipposideros caffer7	Tanzania	-4.902	39.688
FMNH 192792	MT149802					Hipposideros caffer7	Tanzania	-5.367	39.645
FMNH 192793	redundant					Hipposideros caffer7	Tanzania	-5.367	39.645
FMNH 192794	redundant					Hipposideros caffer7	Tanzania	-5.367	39.645
FMNH 192795	redundant					Hipposideros caffer7	Tanzania	-5.367	39.645
FMNH 192796	MT149803					Hipposideros caffer7	Tanzania	-5.367	39.645
FMNH 192849	MT149804	MT149660	MT149559	MT149462	MT149361	Hipposideros caffer7	Tanzania	-4.902	39.688
FMNH 187140	MT149805	MT149661	MT149560	MT149463	MT149362	Hipposideros caffer8	Tanzania	-3.798	36.069
FMNH 219065	MT149806	MT149662	MT149561	MT149464	MT149363	Hipposideros caffer8	Tanzania	-8.037	34.502
FMNH 219241	MT149807					Hipposideros caffer8	Tanzania	-8.037	34.502
FMNH 219242	MT149808					Hipposideros caffer8	Tanzania	-7.707	34.031
FMNH 232868	redundant					Hipposideros caffer8	Uganda	2.240	31.688
FMNH 232869	MT149809	MT149663	MT149562	MT149465	MT149364	Hipposideros caffer8	Uganda	2.240	31.688
FMNH 232874	redundant	MT149664	MT149563	MT149416	MT149365	Hipposideros caffer8	Uganda	2.240	31.688
FMNH 232875	MT149810	MT149665	MT149564		MT149366	Hipposideros caffer8	Uganda	2.240	31.688
LSUMZ MT-4480	redundant	MT149666		MT149466	MT149367	Hipposideros cervinus	Malaysia	1.970	103.500
LSUMZ MT-4481	MT149811	MT149667	MT149565	MT149467	MT149368	Hipposideros cervinus	Malaysia	1.970	103.500
LSUMZ MT-4500	MT149812	MT149668	MT149566	MT149468	MT149369	Hipposideros cervinus	Malaysia	1.970	103.500
UNIMAS 787	EF108144					Hipposideros cervinus	Malaysia	3.316	113.125
UNIMAS 788	EF108146					Hipposideros cervinus	Malaysia	3.316	113.125
LSUMZ MT-4495	MT149813	MT149669		MT149469	MT149370	Hipposideros cf. bicolor	Malaysia	1.970	103.500
UNIMAS 1459	EF108142					Hipposideros cf. bicolor	Malaysia	1.716	110.467
UNIMAS 1474	EF108143					Hipposideros cf. bicolor	Malaysia	1.716	110.467
FMNH 235856	MT149814	MT149670	MT149567	MT149470	MT149371	Hipposideros cf. cervinus	Solomon Islands	-10.569	161.913
FMNH 235857	MT149815	MT149671	MT149568	MT149471	MT149372	Hipposideros cf. cervinus	Solomon Islands	-10.569	161.913
NMP 91848	EU934474					Hipposideros cf. lamottei	Benin	7.783	2.267
NMP 91849	EU934475					Hipposideros cf. lamottei	Benin	7.783	2.267
IVB S862	EU934453					Hipposideros cf. lamottei	Senegal	12.350	-12.317
IVB PV56	HQ343266					Hipposideros cf. ruber	Ghana	7.668	-1.962
DM 12598	KF551812					Hipposideros cf. ruber	Liberia	7.553	-8.492
DM 12620	KF551811					Hipposideros cf. ruber	Liberia	7.553	-8.492
IVB S119	EU934478					Hipposideros cf. ruber	Senegal	13.050	-13.083
IVB S132	HQ343242					Hipposideros cf. ruber	Senegal	14.071	-12.572
IVB S1374	EU934479					Hipposideros cf. ruber	Senegal	13.250	-13.217
IVB S8	HQ343240					Hipposideros cf. ruber	Senegal	12.884	-12.755
LSUMZ MT-4423	DQ054809					Hipposideros cineraceus	Malaysia	3.717	102.167
FMNH 190042	JQ915701					Hipposideros coronatus	Philippines	9.097	125.705
FMNH 202631	JQ915702					Hipposideros coronatus	Philippines	9.764	124.266
KU 166444		MT149672		MT149472	MT149373	Hipposideros coronatus	Philippines	11.813	125.278
	EF108148					Hipposideros coxi	Malaysia	1.378	110.120
	EF108147 redundant					Hipposideros coxi	Malaysia	1.378	110.120
UNIMAS 1424	EF108149					Hipposideros diadema	Malaysia	5.531	118.072
KU 164028	MT149816	MT149673	MT149569	MT149473	MT149374	Hipposideros diadema	Philippines	19.331	121.439
KU 164029	MT149817	MT149674	MT149570	MT149474	MT149375	Hipposideros diadema	Philippines	19.331	121.439
KU 164245	MT149818	MT149675	MT149571	MT149475		Hipposideros diadema	Philippines	13.796	120.159
	FJ460489					Hipposideros doriae	Malaysia	1.117	110.217
NHMOU.CHI MP4.2016	KY176014					Hipposideros durgadasi	India	23.317	78.414
UNIMAS 312	EF108150					Hipposideros dyacorum	Malaysia	4.401	117.889
UNIMAS 556	EF108151					Hipposideros dyacorum	Malaysia	3.316	113.125
	EU934468					Hipposideros fuliginosus	Guinea Bissau	11.117	-14.933
	EU934467					Hipposideros fuliginosus	Guinea Bissau	11.333	-13.900
	JX849198					Hipposideros griffini	Vietnam
CVVD AG	200700214	JN247005					Hipposideros halophyllus	Thailand
NMP 91842	EU934471					Hipposideros jonesi	Benin	7.783	2.267
IVB S804	EU934472					Hipposideros jonesi	Senegal	12.350	-12.317
	EU934473					Hipposideros jonesi	Senegal	12.350	-12.317
EBD 23514	DQ054816					Hipposideros khaokhouayensis	Laos	18.433	102.950
	KF551824					Hipposideros lamottei	Guinea	7.570	-8.471
	KF551823					Hipposideros lamottei	Guinea	7.570	-8.471
NHMOU.CHI MP15.2016	KY176015					Hipposideros lankadiva	India	23.317	78.414
LSUMZ MT-4478	MT149819	MT149676	MT149572	MT149476	MT149376	Hipposideros larvatus	Malaysia	1.970	103.500
LSUMZ MT-4479	redundant	MT149677	MT149573		MT149377	Hipposideros larvatus	Malaysia	1.970	103.500
LSUMZ MT-4488	MT149820	MT149678	MT149574	MT149477	MT149378	Hipposideros larvatus	Malaysia	1.970	103.500
UNIMAS 1485	EF108152 redundant					Hipposideros larvatus	Malaysia	1.717	110.467
UNIMAS 1501	EF108153					Hipposideros larvatus	Malaysia	1.717	110.467
FMNH 195507	JQ915904					Hipposideros lekaguli	Philippines	16.314	121.394
	KR908661					Hipposideros lylei	China	25.603	99.752
DM 12607	KF551825					Hipposideros marisae	Liberia	7.553	-8.492
FMNH 140601	JQ915906					Hipposideros obscurus	Philippines	13.767	124.350
KU 165040	redundant	MT149679	MT149575		MT149379	Hipposideros obscurus	Philippines	11.434	122.079
KU 165041	MT149821	MT149680	MT149576	MT149478	MT149380	Hipposideros obscurus	Philippines	11.434	122.079
KU 165226	MT149717		MT149577	MT149479	MT149381	Hipposideros obscurus	Philippines	13.447	120.426
PSUZC MM2006.129	JN247029					Hipposideros pendelburyi	Thailand	7.565	99.624
	DQ054810					Hipposideros pomona	Laos	18.250	104.517
	EU434952					Hipposideros pratti	China	27.729	115.734
FMNH 190070	JQ915992					Hipposideros pygmaeus	Philippines	9.097	125.705
KU 164542	MT149822	MT149681	MT149578	MT149480	MT149382	Hipposideros pygmaeus	Philippines	14.823	121.968
KU 164543	redundant	MT149682	MT149579	MT149481	MT149383	Hipposideros pygmaeus	Philippines	14.823	121.968
KU 164544	MT149716	MT149683	MT149580	MT149482	MT149384	Hipposideros pygmaeus	Philippines	14.823	121.968
LSUMZ MT-4425	MT149715	MT149684	MT149581	MT149483	MT149385	Hipposideros ridleyi	Malaysia	3.557	102.761
LSUMZ MT-4477	MT149823		MT149582	MT149484	MT149386	Hipposideros ridleyi	Malaysia	3.557	102.761
SMF 83828	DQ054811					Hipposideros ridleyi	Malaysia	3.717	102.167
	DQ054813					Hipposideros rotalis	Laos	18.250	104.517
	FJ347996					Hipposideros ruber1	Cameroon	3.150	13.000
	FJ347995					Hipposideros ruber1	Cameroon	4.451	11.571
	FJ347993					Hipposideros ruber1	Cameroon	3.564	13.408
	FJ347992					Hipposideros ruber1	Cameroon	3.564	13.408
	FJ347989					Hipposideros ruber1	Cameroon	5.385	11.688
FMNH 195085	MT149824	MT149685	MT149583	MT149485	MT149387	Hipposideros ruber1	D. R. Congo	-4.991	29.080
FMNH 215448	MT149826					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215449	MT149827					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215450	redundant					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215451	redundant					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215452	redundant					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215453	redundant					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215476	MT149828					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215477	redundant					Hipposideros ruber1	Kenya	1.036	34.753
FMNH 215478	redundant					Hipposideros ruber1	Kenya	1.036	34.753
NMK 184904	MT149825	MT149686	MT149584	MT149486		Hipposideros ruber1	Kenya	0.212	34.899
NMK 184905	redundant					Hipposideros ruber1	Kenya	0.212	34.899
NMK 187407	redundant					Hipposideros ruber1	Kenya	1.036	34.753
NMK 187408	MT149829					Hipposideros ruber1	Kenya	1.036	34.753
NMK 187409	MT149830	MT149687	MT149585	MT149487	MT149388	Hipposideros ruber1	Kenya	1.036	34.753
NMK 187410	redundant					Hipposideros ruber1	Kenya	1.036	34.753
NMK 187412	MT149831					Hipposideros ruber1	Kenya	1.036	34.753
DM 12603	KF551819					Hipposideros ruber1	Liberia	7.553	-8.492
DM 13245	KF551815					Hipposideros ruber1	Liberia	7.553	-8.492
DM 13246	KF551820					Hipposideros ruber1	Liberia	7.553	-8.492
FMNH 225201	MT149832	MT149688	MT149586	MT149488	MT149389	Hipposideros ruber1	Rwanda	-2.485	29.199
FMNH 225202	MT149833					Hipposideros ruber1	Rwanda	-1.504	29.613
FMNH 225203	redundant					Hipposideros ruber1	Rwanda	-1.504	29.613
FMNH 225204	redundant					Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 225205	redundant					Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 225206	MT149834					Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 225207	redundant					Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 225208	MT149835	MT149689	MT149587	MT149489	MT149390	Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 225209	redundant					Hipposideros ruber1	Rwanda	-1.506	29.615
FMNH 192935	MT149836	MT149690	MT149588	MT149490	MT149391	Hipposideros ruber1	Tanzania	-1.094	31.515
FMNH 137629	FJ347987					Hipposideros ruber1	Uganda	32.283	-0.005
FMNH 160358	MT149837					Hipposideros ruber1	Uganda	-0.989	29.614
FMNH 160359	MT149838	MT149691	MT149589	MT149491	MT149392	Hipposideros ruber1	Uganda	-0.989	29.614
FMNH 160361	MT149839					Hipposideros ruber1	Uganda	-1.041	29.580
FMNH 161040	MT149840					Hipposideros ruber1	Uganda	-0.245	29.819
FMNH 223866	redundant	MT149692	MT149590	MT149492	MT149393	Hipposideros ruber1	Uganda	-0.342	31.966
FMNH 223867	MT149841					Hipposideros ruber1	Uganda	-0.342	31.966
FMNH 227415	MT149842					Hipposideros ruber2	Central African Republic	3.033	16.410
FMNH 227416	MT149843					Hipposideros ruber2	Central African Republic	3.033	16.410
FMNH 227417	MT149844	MT149693	MT149591		MT149394	Hipposideros ruber2	Central African Republic	3.033	16.410
FMNH 149408	FJ347986	MT149694	MT149592	MT149493	MT149395	Hipposideros ruber2	D. R. Congo	-1.417	28.583
FMNH 149409	redundant					Hipposideros ruber2	D. R. Congo	-1.417	28.583
FMNH 149410	redundant	MT149695	MT149593	MT149494	MT149396	Hipposideros ruber2	D. R. Congo	-1.417	28.583
FMNH 149412	redundant					Hipposideros ruber2	D. R. Congo	-1.417	28.583
ROM 100546	FJ347978					Hipposideros ruber2	Ivory Coast	6.930	-7.217
FMNH 215479	redundant					Hipposideros ruber2	Kenya	0.244	34.907
FMNH 215480	MT149845					Hipposideros ruber2	Kenya	0.244	34.907
FMNH 215481	MT149846					Hipposideros ruber2	Kenya	0.244	34.907
FMNH 215482	redundant	MT149696	MT149594	MT149495	MT149397	Hipposideros ruber2	Kenya	0.244	34.907
FMNH 215483	redundant	MT149697	MT149595	MT149496	MT149398	Hipposideros ruber2	Kenya	0.244	34.907
NMK 184878	MT149847					Hipposideros ruber2	Kenya	0.248	34.906
NMK 184880	MT149848					Hipposideros ruber2	Kenya	0.248	34.906
NMK 184882	MT149849					Hipposideros ruber2	Kenya	0.248	34.906
NMK 184883	MT149850					Hipposideros ruber2	Kenya	0.248	34.906
NMK 184884	MT149851					Hipposideros ruber2	Kenya	0.248	34.906
NMK 187383	redundant					Hipposideros ruber2	Kenya	0.212	34.899
NMK 187384	MT149852					Hipposideros ruber2	Kenya	0.212	34.899
FMNH 165161	redundant					Hipposideros ruber2	Uganda	1.733	31.467
FMNH 165162	MT149853					Hipposideros ruber2	Uganda	1.683	31.533
FMNH 165163	redundant	MT149698	MT149596	MT149497	MT149399	Hipposideros ruber2	Uganda	1.683	31.533
FMNH 165164	MT149854					Hipposideros ruber2	Uganda	1.683	31.533
FMNH 165165	MT149855					Hipposideros ruber2	Uganda	1.683	31.533
FMNH 165166	redundant					Hipposideros ruber2	Uganda	1.750	31.583
FMNH 165167	redundant	MT149699	MT149597	MT149498	MT149400	Hipposideros ruber2	Uganda	1.733	31.467
FMNH 224069	redundant					Hipposideros ruber2	Uganda	0.501	30.426
FMNH 224071	redundant					Hipposideros ruber2	Uganda	0.501	30.426
FMNH 224074	MT149856					Hipposideros ruber2	Uganda	0.501	30.426
FMNH 224075	MT149857					Hipposideros ruber2	Uganda	0.501	30.426
	FJ347994					Hipposideros ruber3	Cameroon	3.564	13.408
	FJ347991					Hipposideros ruber3	Cameroon	2.941	9.911
	FJ347990					Hipposideros ruber3	Cameroon	2.941	9.911
	FJ347988					Hipposideros ruber3	Cameroon	4.913	9.241
EBD 18240	FJ347984					Hipposideros ruber3	Equatorial Guinea	1.889	9.793
EBD 18266	FJ347985					Hipposideros ruber3	Equatorial Guinea	1.889	9.793
EBD 18511	FJ347983					Hipposideros ruber3	Equatorial Guinea	3.747	8.750
EBD 18942	FJ347981					Hipposideros ruber3	Principe	1.615	7.404
EBD 18926	FJ347982					Hipposideros ruber3	São Tomé	0.219	6.727
FMNH 219477	MT149858	MT149700	MT149598	MT149499	MT149401	Hipposideros ruber4	D. R. Congo	-5.290	14.871
FMNH 169707	KT583815					Macronycteris commersoni	Madagascar	-12.932	49.057
FMNH 175777	KT583822					Macronycteris commersoni	Madagascar	-16.380	45.345
FMNH 175966	KT583823					Macronycteris commersoni	Madagascar	-22.486	45.392
FMNH 175974	MT149859					Macronycteris commersoni	Madagascar	-22.317	45.293
FMNH 175975	MT149860					Macronycteris commersoni	Madagascar	-22.317	45.293
FMNH 176155	KT583824	MT149701	MT149599	MT149500	MT149402	Macronycteris commersoni	Madagascar	-22.778	43.523
FMNH 176158	MT149861					Macronycteris commersoni	Madagascar	-22.217	43.330
FMNH 176277	redundant					Macronycteris commersoni	Madagascar	-12.942	49.055
FMNH 177302	KT583825					Macronycteris commersoni	Madagascar	-16.315	46.810
FMNH 178803	redundant					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178806	KT583816					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178808	KT583817 redundant					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178809	KT583818					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178810	KT583819					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178811	KT583820					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178812	KT583826					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 178815	KT583821 redundant					Macronycteris commersoni	Madagascar	-12.712	49.474
FMNH 179201	MT149862					Macronycteris commersoni	Madagascar	-17.280	49.420
FMNH 183934	KT583827					Macronycteris commersoni	Madagascar	-24.050	43.750
FMNH 183980	KT583813 redundant					Macronycteris commersoni	Madagascar	-12.337	49.385
FMNH 184030	KT583812					Macronycteris commersoni	Madagascar	-14.966	47.308
FMNH 184170	KT583828					Macronycteris commersoni	Madagascar	-24.650	43.963
FMNH 184887	MT149863					Macronycteris commersoni	Madagascar	-15.904	46.598
FMNH 209236	MT149864					Macronycteris commersoni	Madagascar	-18.063	44.541
FMNH 217940	KT583831 redundant					Macronycteris commersoni	Madagascar	-22.632	45.338
FMNH 221308	KT583829					Macronycteris commersoni	Madagascar	-12.932	49.057
FMNH 231862	MT149865	MT149702	MT149600	MT149501	MT149315	Macronycteris commersoni	Madagascar	-17.889	49.203
UADBA 32916	KT583830					Macronycteris commersoni	Madagascar	15.538	46.886
UADBA 32987	KT583814					Macronycteris commersoni	Madagascar	-13.932	49.057
UADBA 32989	KR606333					Macronycteris commersoni	Madagascar	-12.917	49.143
FMNH 175970	MT149866	MT149703	MT149601	MT149502	MT149403	Macronycteris cryptovalorona	Madagascar	-22.317	45.293
FMNH 184173	MT149867	MT149704	MT149602	MT149503	MT149404	Macronycteris cryptovalorona	Madagascar	-24.650	43.963
AMNH 269871	KT583801					Macronycteris gigas	Central African Republic
FMNH 219602	MT149868	MT149705	MT149603	MT149504	MT149405	Macronycteris gigas	D. R. Congo	0.241	20.883
FMNH 219682	MT149869	MT149706	MT149604	MT149415	MT149406	Macronycteris gigas	D. R. Congo	0.241	20.883
FMNH 220226	redundant	MT149707	MT149605	MT149505	MT149407	Macronycteris gigas	Kenya	-4.647	39.380
FMNH 220227	redundant	MT149708	MT149606	MT149506	MT149408	Macronycteris gigas	Kenya	-4.647	39.380
FMNH 220239	MT149870					Macronycteris gigas	Kenya	-4.215	39.451
DM 12602	KF551826					Macronycteris gigas	Liberia	7.553	-8.492
IVB S1032	EU934469					Macronycteris gigas	Senegal	12.350	-13.217
IVB S1044	EU934470					Macronycteris gigas	Senegal	12.350	-13.217
AMNH 269879	KT583802					Macronycteris vittata	Central African Republic	3.500	16.000
FMNH 215942	MT149871	MT149709	MT149607	MT149507	MT149409	Macronycteris vittata	Kenya	-3.300	39.995
FMNH 215943	redundant					Macronycteris vittata	Kenya	-3.300	39.995
FMNH 215944	redundant					Macronycteris vittata	Kenya	-3.300	39.995
FMNH 215945	redundant					Macronycteris vittata	Kenya	-3.300	39.995
FMNH 215946	MT149872					Macronycteris vittata	Kenya	-3.287	39.982
FMNH 215947	redundant					Macronycteris vittata	Kenya	-3.287	39.982
FMNH 215948	MT149873					Macronycteris vittata	Kenya	-3.287	39.982
FMNH 215949	redundant					Macronycteris vittata	Kenya	-3.287	39.982
FMNH 215950	MT149874					Macronycteris vittata	Kenya	-3.287	39.982
FMNH 215951	MT149875					Macronycteris vittata	Kenya	-3.282	39.971
FMNH 215952	MT149876					Macronycteris vittata	Kenya	-3.282	39.971
FMNH 215953	redundant	MT149710	MT149608		MT149410	Macronycteris vittata	Kenya	-3.282	39.971
FMNH 215954	MT149877					Macronycteris vittata	Kenya	-3.282	39.971
FMNH 215955	redundant					Macronycteris vittata	Kenya	-3.282	39.971
FMNH 215959	MT149878					Macronycteris vittata	Kenya	-3.309	40.018
FMNH 215960	MT149879					Macronycteris vittata	Kenya	-3.309	40.018
FMNH 215961	MT149880					Macronycteris vittata	Kenya	-3.309	40.018
FMNH 215962	MT149881					Macronycteris vittata	Kenya	-3.309	40.018
FMNH 215963	MT149882					Macronycteris vittata	Kenya	-3.309	40.018
FMNH 215967	MT149883					Macronycteris vittata	Kenya	-3.303	39.999
FMNH 215968	redundant					Macronycteris vittata	Kenya	-3.305	39.937
FMNH 215969	redundant					Macronycteris vittata	Kenya	-3.305	39.937
FMNH 220224	redundant					Macronycteris vittata	Kenya	-4.647	39.378
FMNH 220225	MT149885					Macronycteris vittata	Kenya	-4.647	39.380
FMNH 220228	MT149886					Macronycteris vittata	Kenya	-4.647	39.380
FMNH 220229	MT149887	MT149711	MT149609	MT149508	MT149411	Macronycteris vittata	Kenya	-4.647	39.380
FMNH 220231	MT149888					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220232	MT149889					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220233	MT149890					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220234	redundant					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220235	MT149891	MT149712	MT149610	MT149509	MT149412	Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220236	redundant					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220237	redundant					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220238	redundant					Macronycteris vittata	Kenya	-4.614	39.354
FMNH 220240	redundant					Macronycteris vittata	Kenya	-3.300	39.995
NMK 187219	redundant					Macronycteris vittata	Kenya	-3.335	40.031
NMK 187220	MT149884					Macronycteris vittata	Kenya	-3.335	40.031
NMK 187221	redundant					Macronycteris vittata	Kenya	-3.335	40.031
NMK 187222	redundant					Macronycteris vittata	Kenya	-3.335	40.031
DM 8645	KF551828 redundant					Macronycteris vittata	Mozambique	-18.565	32.220
DM 11510	KF551827					Macronycteris vittata	Mozambique	-18.978	34.176
FMNH 192800	redundant	MT149713	MT149611	MT149510	MT149413	Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192801	MT149892					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192857	redundant	MT149714	MT149612	MT149511	MT149414	Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192858	redundant					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192859	MT149893					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192860	KT583807					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192865	KT583808 redundant					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 192866	KT583809					Macronycteris vittata	Tanzania	-4.902	39.688
FMNH 220268		MT149614	MT149512	MT149417	MT149316	Triaenops afer	Kenya	-4.590	39.331

Citation

Patterson BD, Webala PW, Lavery TH, Agwanda BR, Goodman SM, Kerbis Peterhans JC, Demos TC (2020) Evolutionary relationships and population genetics of the Afrotropical leaf-nosed bats (Chiroptera, Hipposideridae). ZooKeys 929: 117–161. https://doi.org/10.3897/zookeys.929.50240

Funding Statement

JRS Biodiversity Foundation; Field Museum’s Council on Africa, Marshall Field III Fund, and Barbara E. Brown Fund for Mammal Research; The John D. and Catherine T. MacArthur Foundation, Fulbright Program of US Department of State, Wildlife Conservation Society, and the Centers for Disease Control and Prevention; WWF Gabon; MoZaR-Fond Européen de Développement Régional, Programme Opérationnel de Coopération Territoriale.

Supplementary materials

Supplementary material 1

Figure S1. Geographic distribution of voucher specimens used in this analysis

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Bruce D. Patterson, Paul W. Webala, Tyrone H. Lavery, Bernard R. Agwanda, Steven M. Goodman, Julian C. Kerbis Peterhans, Terrence C. Demos

Data type

occurrence

Supplementary material 2

Figure S2. Phylogeny of Hipposideridae based on maximum likelihood analysis of cyt-b based on 452 individuals

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Bruce D. Patterson, Paul W. Webala, Tyrone H. Lavery, Bernard R. Agwanda, Steven M. Goodman, Julian C. Kerbis Peterhans, Terrence C. Demos

Data type

phylogenetic tree

Supplementary material 3

Figure S3. Phylogeny of Hipposideridae based on Bayesian inference analysis of cyt-b based on 452 individuals

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Bruce D. Patterson, Paul W. Webala, Tyrone H. Lavery, Bernard R. Agwanda, Steven M. Goodman, Julian C. Kerbis Peterhans, Terrence C. Demos

Data type

phylogenetic tree