Cryptic Diversity in the Genus Miniopterus with the Description of a New Species from Southern Africa

Home , Chaerephon (bat), Miniopterus

Acta Chiropterologica, 22(1): 1–19, 2020 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS doi: 10.3161/15081109ACC2020.22.1.001

Cryptic diversity in the genus Miniopterus with the description of a new species from southern Africa

ARA MONADJEM1, 2, 9, JEN GUYTON3, PIOTR NASKRECKI4, 5, LEIGH R. RICHARDS6, ANNA S. KROPFF7, and DESIRE L. DALTON7, 8

1Department of Biological Sciences, University of Eswatini, Private Bag 4, Kwaluseni, Eswatini 2Mammal Research Institute, Department of Zoology and Entomology, University of Pretoria, Private Bag 20, Hatfield 0028, Pretoria, South Africa 3Department of Ecology and Evolutionary Biology, Princeton University, 106A Guyot Hall, Princeton, NJ 08544, USA 4E.O. Wilson Biodiversity Laboratory, Gorongosa National Park, Mozambique 5Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA 6Durban Natural Science Museum, PO Box 4085, Durban 4000, South Africa 7South African National Biodiversity Institute, P.O. Box 754, Pretoria, 0001, South Africa 8Department of Zoology, University of Venda, Thohoyandou, South Africa 9Corresponding author: E-mail: [email protected]

Species richness in the genus Miniopterus has been greatly under-reported, with a large number of taxa having been discovered and described in the past two decades. Using molecular, standard morphometrics and acoustic data, we present evidence for the existence of a new species in Mozambique and neighbouring Malawi. Based on cytochrome b (cyt b) and cytochrome oxidase I (COI), the new species is sister to M. minor, from which it is readily distinguishable by its larger size (including non-overlapping forearm measurements, allowing separation in the field). It is distinguishable from sympatric M. mossambicus, itself a newly described taxon from Mozambique, by forearm measurements and a peach-orange wash to the skin around the eyes. In external appearance, it is most similar to M. fraterculus, from which it is only reliably identifiable by multivariate analysis of craniodental features and by a genetic distance of 6.4% in the cyt b gene; the two species also occupy widely differing geographic ranges. The type locality of the new species is Mount Gorongosa, and all known records are from large mountains in central and northern Mozambique and southern Malawi. Further research is required to establish its geographic range and understand its basic ecology. Considering its relatively restricted distribution to threatened montane habitats, we suggest that its global conservation status be urgently assessed.

Key words: cryptic species, Miniopteridae, cytochrome b, morphometrics, taxonomy, Mozambique

INTRODUCTION Many of the newly described taxa are cryptic species, not easily distinguishable based on external Small mammals remain poorly known in the characteristics (Monadjem et al., 2013b). The genus Afrotropics (Kingdon et al., 2013), despite this Miniopterus appears to be particularly rich in cryptic being one of the most diverse regions of the world species, clearly demonstrated by the rapid increase for mammals (Burgin et al., 2018). Indeed, it has in the number of recognized species since the appli- been predicted that a greater number of mammal cation of molecular techniques to systematic studies species remain to be described in Africa than on any of this group (Goodman et al., 2007, 2009). In 2005, other continent (Fisher et al., 2018). The number of just four species of Miniopterus were recognized on new bat taxa recognized in sub-Saharan Africa con- Madagascar (Simmons, 2005); within 10 years that tinues to rise, with 33 species having been described number had risen to 12 species (Christidis et al., or elevated to species rank in the 30 years from 1988 2014; Goodman et al., 2015). to 2018 excluding those described from Madagascar Progress on resolving the taxonomy and system- and other offshore islands (Hoffmann et al., 2009; atics of Miniopterus from mainland Africa has been Taylor et al., 2018). far slower, with only three new species having been 2 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff recently described (Monadjem et al., 2013a, 2019; distribution closely associated with the high moun- Puechmaille et al., 2014). This is unlikely to be due tains of Mozambique and Malawi. Recent studies to any inherent lack of diversity for biogeographic point to this region having a large number of en- reasons. On a single mountain in Malawi, south-cen- demic species (Bayliss et al., 2014; Conradie et al., tral Africa, four Miniopterus species were reported, 2016), many with small distributions centered on but none of them could be named with certainty one or a few mountains such as the recently de- (Curran et al., 2012); this compares well with the scribed gecko Afroedura gorongosa (Branch et al., situation in Madagascar where up to four species of 2017). Miniopterus may co-occur (Goodman et al., 2009). Indeed, a recent phylogeny based on a multilocus MATERIALS AND METHODS approach suggested the existence of up to five undescribed taxa in Africa (Demos et al., 2019). Samples The taxonomy of Miniopterus in West Africa has recently been investigated, where currently two spe - Bats were captured at Mount Gorongosa (18.48°S, 34.04°E, cies occur, both endemic to the Upper and Lower elevation 920 m above sea level [a.s.l.]) (Fig. 1), and deposited in either the Durban Natural Science Museum, Durban, South Guinea forest block (Monadjem et al., 2019). North Africa (permit information: N/Ref.133/MHN/E.27/2015; 13/ Africa also has two species, one of which was re- 1/1/30/2/0-215/08/005246; OP 3321/2015; OP3318/215) or the cently described as a cryptic species (Puechmaille et EO Wilson Laboratory based in Chitengo, Gorongosa National al., 2014). The situation in East Africa is compli- Park, Mozambique (permits PNG/DSCi/Cll/2015, PNG/DSCi/ cated by the lack of recently published surveys, but C8/2015). Tissue samples were taken from the pectoral muscles of collected specimens. at least one new species, resulting from the elevation of an existing taxon to specific status, has been un- Genomic DNA Isolation, Amplification and covered by molecular analysis (Šrámek et al., 2013). Sequencing Furthermore, the recent multi-locus phylogeny referred to above, demonstrated that nine of the Genomic DNA was isolated from tissue samples using the 13 recognized clades of African Miniopterus occur QIAamp DNA Investigator Kit (Qiagen, Germany). Tissue sam- in East Africa (Demos et al., 2019). ples were cut with a scalpel blade and were subsequently di- Three species have been traditionally recognized gested overnight (20–22 hours) in Proteinase K and ATL tissue lysis buffer. Following digestion, DNA was isolated according in southern Africa south of the Zambezi River, dif- to the manufacturer’s instructions. Primers were used to amplify ferentiated mostly on size (Stoffberg et al., 2004) regions of the mitochondrial genes cytochrome oxidase I (COI, from smallest to largest: M. fraterculus, M. nata- 520 bp) and cytochrome b (cyt b, 1140 bp). The COI gene was lensis and M. inflatus (Monadjem et al., 2010b). amplified using the universal conservative primers BatL5310 A fourth species, M. mossambicus, was recently de- and R6036R (Hebert et al., 2003). In addition, L14724 and H15915 (Xiao et al., 2001) and L15162 and H15915 (Irwin et scribed from northern Mozambique but was thought al., 1991) were used to amplify a region of cyt b. Amplification to extend further south into central Mozambique of the respective gene regions was carried out in separate PCR (Monadjem et al., 2013a) and represents the small- reactions consisting of 1 × DreamTaq Green PCR Master Mix, est Miniopterus species in the region. These four 0.4 µM of each primer, and approximately 20 ng template DNA species are not easily identifiable in the field but are in a total volume of 20 µl. The temperature profile was as follows: an initial denaturation at 95°C for 2 min, 35 cycles of clearly distinguishable on cranial and genetic char- 95°C for 30 s, 55°C for 30 s, and 72°C for 1 min, followed by acteristics (Miller-Butterworth et al., 2005; Mona- a final extension at 72°C for 10 min. Successful PCR products djem et al., 2013a), underscoring the fact that cryp- were purified with Exonuclease I and FastAP (Thermo Fisher tic species abound in this genus. Adding to the Scientific Inc.). Gene fragments were sequenced in both direc- confusion is the possibility of a fifth species occur- tions using the BigDye Terminator v3.1 Cycle Sequencing Kit and visualized on a 3500 Genetic Analyzer (Applied Biosys - ring in this region, another diminutive species, tems). Sequence chromatograms were viewed using Sequencing M. mi nor which is currently known from Tanzania, Analysis Software v.6.0 (ThermoFisher Scientific). Central African Republic and Democratic Republic of Congo (Monadjem et al., 2010b), with the west- Phylogenetic Analysis ern subspecies occidentalis probably a separate species (Juste and Ibañez, 1992). The dataset consisted of 37 cyt b sequences (Table 1), which Recent bat surveys on Mount Gorongosa contained 35 reference sequences representing the species Miniopterus fraterculus Thomas and Schwann, 1906; M. fuligi- recorded a small Miniopterus species on the mid- nosus (Hodgson, 1835), M. gleni Peterson, Eger and Mitchell, slopes of the mountain. Here, we show that this an- 1995, M. griveaudi Harrison, 1959, M. cf. inflatus, M. macroc- imal represents an undescribed taxon, with a larger neme Revilliod, 1914, M. majori Thomas, 1906, M. manavi Cryptic diversity in southern African Miniopterus 3

FIG. 1. Map of southern and eastern Africa showing the distribution of the newly described species, Miniopterus sp. nov., as well as other species mentioned in this study. The location of the type locality is indicated by an arrow, and country names referred to in the text are provided. Specimens used in the genetic and/or morphological analyses are indicated separately

Thomas, 1906, M. minor Peters, 1857, M. mossambicus Mona - Monte Carlo (MCMC) analysis was conducted using the djem, Goodman, Stanley and Appleton, 2013, M. natalensis above mentioned model parameters (GTR G+I: nst = 6; rates = (A. Smith, 1833), M. orianae Thomas, 1922, and M. sororculus invgamma), default parameters for the estimations of priors, one Goodman et al., 2007, obtained from National Center for cold and three heated chains, and run for 100,000 generations Biotechnology Information (NCBI) GenBank, Barcode of Life sampled every 100 generations with a burn in of 25%. By de- Data System (BOLD) and International Barcode of Life (iBOL). fault, two simulations are conducted at the same time. A stop The COI dataset included ten sequences of which eight were rule parameter using the average standard deviation of split fre- reference sequences from M. fuliginosus, M. magnater Sanborn, quencies (SD) between the two simulations was also imple- 1931, M. minor, M. cf. natalensis, M. natalensis and M. schrei - mented to stop analysis when the threshold (SD = 0.01) was bersii (Kuhl, 1817) (Table 2). Lastly, two field isolate sequen- reached, which occurred at 750,000 generations. An SD value of ces (JAG444, JAG445) generated by South African National 0.05 is the default value used as a diagnostic determinant of Bio diversity Institute (SANBI), and Chaerephon pumilus convergence of the two simulations. An SD value approaching (Cretzs ch mar, 1826) and Scotophilus dinganii (A. Smith, 1833) 0 indicates that the two simulations become increasingly simi- were used as outgroups. lar. The generated tree file was converted to the Newick format All sequences were manually trimmed and aligned with using FigTree (v1.4.3) and annotated in MEGA7. Molecular MUSCLE (Edgar, 2004) using default parameters in MEGA 7 clock analysis was conducted in BEAST 1.7.4 (Drummond and version 7.0.26 (Kumar et al., 2016). The best model for se- Rambaut, 2007; Drummond et al., 2012) following methods de- quence evolution, General Time Reversible (GTR) model with scribed in Christidis et al. (2014). Gamma distribution (G = 1.53) and Invariable sites (I = 0.58), was determined under the Bayesian Information Criterion (BIC) Morphological Analysis using the model test function incorporated in MEGA7. Phylogenetic relationships were evaluated using the ML and NJ The specimens (listed in Appendix) on which the mor- method implemented in MEGA 7 (Kumar et al., 2016). To esti- phometric study is based are deposited in the following collec- mate support for internal nodes, 1000 bootstrap replications tions: The Natural History Museum (formerly The British were run using the same program (Felsenstein, 1985; Kumar et Museum of Natural History), London (BMNH); Durban Natural al., 2016). Sequence variation and average sequence divergence Science Museum (DM); Field Museum of Natural History, were determined by group mean distances using the p-distance Chicago (FMNH); Muséum national d’Histoire naturelle, Paris substitution model in MEGA 7 (Kumar et al., 2016). Aligned se- (MNHN); and E.O. Wilson Biodiversity Laboratory, Gorongosa quences were exported to a Fasta file format for Bayesian National Park (EOWL). Measurements of only adult specimens, Inference (BI) analysis in Mr Bayes (v3.2.7). Markov Chain identified by fully erupted adult dentition and the fusion of 4 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff

TABLE 1. Cytochrome b (cyt b) sequences of Miniopterus species, and outgroups, used in this study GenBank No. Species Locality Reference JAG444 Miniopterus sp. nov. Mozambique This study JAG445 M. sp. nov. Mozambique This study AY614744.1 M. natalensis South Africa Miller-Butterworth et al. (2005) KF709542 M. natalensis Namibia Monadjem et al. (2013a) KF709543 M. natalensis Namibia Monadjem et al. (2013a) AY614751.1 M. fraterculus South Africa Miller-Butterworth et al. (2005) AY614754.1 M. fraterculus South Africa Miller-Butterworth et al. (2005) AY614755.1 M. fraterculus South Africa Miller-Butterworth et al. (2005) DQ899760.1 M. fraterculus South Africa Goodman et al. (2007) AY614732.1 M. orianae Australia Miller-Butterworth et al. (2005) KJ535821.1 M. schreibersii Romania Puechmaille et al. (2014) KJ535822.1 M. schreibersii Spain Puechmaille et al. (2014) AY614737.1 M. cf. inflatus Malawi Miller-Butterworth et al. (2005) FJ383129 M. manavi Madagascar Goodman et al. (2009) FJ383130 M. manavi Madagascar Goodman et al. (2009) HQ619934 M. manavi Madagascar Goodman et al. (2011) FJ232797.1 M. griveaudi Anjouan Weyeneth et al. (2008) FJ232798.1 M. griveaudi Comore Weyeneth et al. (2008) FJ383136.1 M. griveaudi Madagascar Goodman et al. (2009) KF709538.1 M. mossambicus Mozambique Monadjem et al. (2013a) KF709539.1 M. mossambicus Mozambique Monadjem et al. (2013a) AY614738.1 M. mossambicus Zambia Miller-Butterworth et al. (2005) AY614739.1 M. mossambicus Zambia Miller-Butterworth et al. (2005) FJ232803.1 M. minor Tanzania Weyeneth et al. (2008) FJ232805.1 M. minor Tanzania Weyeneth et al. (2008) FJ232806.1 M. minor Tanzania Weyeneth et al. (2008) JF440236.1 M. gleni Madagascar Ramasindrazana et al. (2011) JF440237.1 M. gleni Madagascar Ramasindrazana et al. (2011) JF440238.1 M. gleni Madagascar Ramasindrazana et al. (2011) HQ619954.1 M. majori Madagascar Goodman et al. (2011) HQ619955.1 M. majori Madagascar Goodman et al. (2011) HQ619939.1 M. majori Madagascar Goodman et al. (2011) DQ899771.2 M. sororculus Madagascar Goodman et al. (2007) DQ899773.2 M. sororculus Madagascar Goodman et al. (2007) HQ619938.1 M. sororculus Madagascar Goodman et al. (2011) AY614734.1 M. macrocneme New Guinea Miller-Butterworth et al. (2005) AB085735 M. fuliginosus Japan Sakai et al. (2003) AY614756.1 Chaerephon pumilus South Africa Miller-Butterworth et al. (2005) AY614757.1 Scotophilus dinganii South Africa Miller-Butterworth et al. (2005)

the basisphenoid-basioccipital suture, were included in this of the cranium; greatest mastoid breadth (MAST), greatest study. We examined type specimens in the BMNH and FMNH, breadth of cranium at mastoid processes; greatest braincase which included: M. fraterculus (BMNH 1905.5.7.18), M. infla- width (GBW), lateral braincase width taken posterior to the pos- tus (BMNH 1903.2.4.8, holotype), M. natalensis (BMNH terior insertion of the zygomatic arches; lachrymal width (LW), 1848.6.12.19, holotype), and M. mossambicus (FMNH 213651, greatest width across rostrum at lachrymal projections; and holotype). greatest mandible length (MAND), taken from the posterior- The following standard external measurements were taken in most point of the condylar processes to the anterior-most point the field: total body length, tail length, ear length, and forearm of the incisors. length. Forearm length was taken with callipers to the closest 0.1 Eight dental measurements were taken with callipers to the mm; all other measurements were at an accuracy of 1 mm. Body closest 0.01 mm following Monadjem et al. (2013b) that in- mass was taken with a Pesola spring balance to the nearest 1 g. cluded: width across the third molars (M3–M3), taken across the Eight cranial measurements were taken with callipers fol- outer-most point of the alveoli of the 3rd molars; complete lowing Monadjem et al. (2013b) that included: greatest skull upper canine-molar tooth row (C–M3), taken from the anterior- length (GSKL), from the posterior-most point of the cranium to most point of the alveolus of the canine to the posterior-most the anterior-most point of the incisors; condylo-incisive length point of the 3rd molar; complete upper tooth row (I1–M3), taken (CIL), from the occipital condyles to the anterior-most point of from the anterior-most point of the alveolus of the first incisor the incisors; greatest zygomatic breadth (ZYGO), the greatest to the posterior-most point of the 3rd molar; complete upper width across the zygomatic arches; postorbital width (POB), molar tooth row (UPMOLS), taken from the anterior-most point narrowest dorsal width posterior to the postorbital constriction of the alveolus of the anterior premolar to the posterior-most Cryptic diversity in southern African Miniopterus 5

TABLE 2. Cytochrome oxidase I (COI) sequences of Miniopterus species, and outgroups, used in this study

GenBank No. Species Locality Reference JAG444 Miniopterus sp. nov. Mozambique This study JAG445 M. sp. nov. Mozambique This study KF452626 M. natalensis South Africa S. D. McCulloch, unpublished data KR259958 M. natalensis South Africa J. Coertse, unpublished data SKBET039-07 M. cf. natalensis Ethiopia Barcode of Life Data System (BOLD) SKBET028-07 M. cf. natalensis Ethiopia Barcode of Life Data System (BOLD) HQ580335 M. fuliginosus Japan International Barcode of Life (iBOL) KP247545 M. magnater China Li et al. (2015) JF442828 M. schreibersii Russia Kruskop et al. (2012) JF442486 M. minor Kenya B. Agwanda and I. V. Kuzmin, unpublished data KF452603 Chaerephon pumilus South Africa S. D. McCulloch, unpublished data MF947528 Scotophilus dinganii South Africa M. Geldenhuys et al., unpublished data

point of the 3rd molar; width across upper canines (C–C), taken new species is sister to M. minor. Genetic distances across the outer-most points of the alveoli of the canines; com- between the various species mentioned in this phy- plete mandibular molar tooth row (LWMOLS), taken from the anterior-most point of the alveolus of the anterior premolar to logeny were 5–16% (Table 3). The new species the posterior-most point of the 3rd molar; and complete lower M. sp. nov. differs from M. minor between 4.1 and tooth row (i1–m3), taken from the anterior-most point of the 5.7% (average = 4.9%). Divergence based on the alveolus of the first incisor to the posterior-most point of the 3rd Beast analysis (Fig. 3) between M. minor and M. sp. molar. Tooth abbreviations are as follows: I = incisor, C = ca- nov. is estimated to have occurred between 1 to nine, P = premolar, M = molar; with upper teeth presented in upper case and lower teeth in lower case. 2 Mya. A principal components analysis (PCA) of standardized values of the above craniodental measurements was conducted on Morphometric Analyses the variance-covariance matrix in package ‘vegan’ (Oksanen et al., 2007) in R version 3.4.4 (R Core Team, 2019), to compare A PCA ordination based on craniodental meas- the morphology of the various taxa measured in this study. Due urements showed that southern African Miniopterus to the lack of sexual dimorphism (Monadjem et al., 2010b), the sexes were combined for all analyses. species mostly occupied distinct morphospace (Fig. 4A). The first two principal components accounted Acoustic Analysis for 92% of the variation, and hence are shown here. The first principal component represented a size gra- Echolocation calls were recorded from hand-released indi- dient with highest loadings on GSKL and ZYGO viduals and individuals flying in a large free-flight tent using (0.313 and 0.311, respectively), with all other load- a full-spectrum recorder Pettersson D1000X at a sampling fre- ings also being positively associated with it (Table quency of 300 kHz. Calls were analysed using BatSound Pro 4). As a result, the species are distributed along this software (version 3.20 — Pettersson Elektronik, Uppsala, Sweden), and the peak frequency was recorded (Monadjem et axis based on size with the smallest species al., 2010b). (M. minor) occurring on the left and the largest species (M. cf. inflatus) on the right of the ordina- RESULTS tion. The second principal component represented differences in shape with high positive and nega- Molecular Analyses tive loadings, the largest being with POS and C–M3 (0.440 and -0.493, respectively) (Table 4). The phylogenies based on cyt b and COI genes Similar sized species were typically separated on produced similar tree topologies for both Bayes- this axis such as M. fraterculus and M. mossambicus ian and neighbor-joining methods (Fig. 2). Basal (Fig. 4A). Only two species did not separate in mor- nodes are poorly supported. However, several phospace: M. mossambicus and the new species clades are apparent. Miniopterus natalensis is sister M. sp. nov. which overlapped somewhat. How- to the Malagasy species; embedded within the ever, when these two species (M. mossambicus latter is M. mossambicus. Miniopterus cf. inflatus and M. sp. nov.) were considered separately, they (from Malawi) is sister to a clade that includes occupied non-overlapping areas of morphospace M. fraterculus, M. minor, and the new species (Fig. 4B), illustrating their distinctiveness in cranial from Mount Gorongosa Miniopterus sp. nov. The and dental features. Based on these molecular and 6 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff

FIG. 2. Phylogenetic position of Miniopterus sp. nov. from Mount Gorongosa based on: A — the cyt b gene produced using Bayesian analysis with posterior probabilities indicated followed by ML bootstrap support; and B — the COI gene produced using NJ analysis with ML bootstrap support indicated. Labels include accession number, species, and area of collection. * indicates samples sequenced in this study TABLE 3. Average genetic distances within and between clades of Miniopterus species as determined with cyt b in MEGA 7 (Tamura et al., 2013); values in bold refer to Miniopterus sp. southern in African diversity Cryptic nov. Values in parentheses indicate within-species distance

Species natalensis sororculus fraterculus majori minor mossambicus griveaudi fuliginosus manavi sp. nov. schreibersii gleni cf. inflatus macrocneme natalensis (0.0118) sororculus 0.096 (0.0039) fraterculus 0.105 0.132 (0.0083) majori 0.097 0.078 0.116 (0.0098) minor 0.108 0.105 0.080 0.095 (0.0039) mossambicus 0.104 0.096 0.103 0.078 0.113 (0.0223) griveaudi 0.095 0.095 0.115 0.087 0.100 0.106 (0.0078) fuliginosus 0.147 0.133 0.165 0.150 0.156 0.155 0.155 (-) manavi 0.087 0.084 0.107 0.067 0.106 0.082 0.103 0.140 (0.0078) sp. nov. 0.103 0.109 0.064 0.103 0.049 0.108 0.110 0.146 0.105 (0.0029) schreibersii 0.102 0.123 0.122 0.126 0.128 0.135 0.122 0.118 0.112 0.118 (0.0882) gleni 0.100 0.094 0.118 0.066 0.101 0.090 0.115 0.148 0.069 0.109 0.120 (0.0098) cf. inflatus 0.104 0.131 0.093 0.097 0.117 0.104 0.118 0.147 0.115 0.102 0.127 0.101 (-) macrocneme 0.114 0.111 0.118 0.122 0.132 0.128 0.128 0.089 0.113 0.102 0.080 0.125 0.121 (-) T D Miniopterus morphometric data,wedescribeanewspeciesof nov., craniodental measurements(seeMaterialsandMethods)of analysis (PCA)forPC1,PC2,andPC3basedonstandardized uuaiettl 859. 94.4 -0.517 -0.265 0.404 -0.098 92.1 0.146 -0.446 0.291 88.5 0.303 0.297 Cumulative total LWMOLS i–m MAND M C–C type isillustratedinFig.5B. ratory ZoologicalMuseum.Photographofthepara - posited intothecollectionsofE.O. Wilson Labo - collected onthesamedayatsiteandde- Paratypes cultural fields. est fragmentadjacenttomontanegrasslandandagri- It wasnettedoverariverinmontaneriverinefor- gosa (18.48309°S,34.04485°W)at920ma.s.l. was collectedon22 April 2018onMountGoron - National Park,MountGorongosa(Fig.1). The bat Type locality Fig. 5A anditsskullin Fig.6. External featuresoftheholotypeareillustratedin ethanol. The skullhasbeenextractedandcleaned. Mozambique. The specimenwaspreservedin70% Laboratory atChitengo,GorongosaNationalPark, lections oftheE.O. Wilson LaboratoryBiodiversity Guyton on22 April 2018,anddepositedintothecol- Holotype S .0 .2 0.424 0.444 0.176 0.121 0.310 0.121 0.440 0.085 0.306 -0.016 0.304 0.290 -0.203 0.311 0.313 C–M GSW MAST POS ZYGO GSKL variation explained (%) ABLE SRPINOF ESCRIPTION 3 –M 3 A singlefemale(JAG 445)wasalsocapturedand Mozambique, SofalaProvince,Gorongosa JAG 444,anadultmale,wascollectedbyJen 3 M. minor 3 4. Eigenvectorloadingsoftheprincipalcomponents hrce C C PC3 PC2 PC1 Character Family MiniopteridaeDobson1875 Genus Miniopterus Miniopterus wilsoni Wilson’s Long-fingeredBat , for Africa. M. fraterculus Miniopterus N EW S PECIES .0 .0 -0.321 0.206 -0.112 -0.163 -0.259 0.301 -0.493 0.307 0.294 , M. natalensis Bonaparte 1837 sp. nov. and M. inflatus M. sp. 7 8 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff and S. P.Richards, A. Guyton, R. J. L. Monadjem, Naskrecki, A. 8

FIG. 3. BEAST molecular clock analysis based on cyt b sequences. Analysis included a Yule tree model under uniform speciation prior based on a relaxed uncorrelated lognormal molecular clock and a variable rate of 2% sequence evolution per lineage per million years Cryptic diversity in southern African Miniopterus 9

AB PC 2

PC 1 PC 1

FIG. 4. PCA ordination based on craniodental measurements of: A — all Miniopterus species known to occur in southern Africa; and B — M. wilsoni sp. nov. and M. mossambicus only. For the latter ordination, forearm length was also included

Etymology 6.4%) and on being larger in dental features (Table This species is named after Edward O. Wilson 6). The two species also occupy non-overlapping who has supported and facilitated biodiversity re- geographic ranges. Based on the cyt b gene, the sis- search at Gorongosa National Park over the past ter species to M. wilsoni sp. nov. is M. minor, from decade. which it can be readily distinguished by its larger, non-overlapping size (forearm length) and most Diagnosis craniodental features (Tables 5 and 6). This is a small-sized Miniopterus currently confirmed by molecular analysis from central Mozam - Description bique, and by morphometric analysis from northern External characters. — Miniopterus wilsoni sp. Mozambique and southern Malawi. Mean forearm nov. has the typical features characteristic of the length for the species is 44.1 mm (n = 8). The small genus including a rounded head, an elongated sec- size of this bat readily distinguishes it from medium- ond phalanx of the third digit, rounded ears, and and large-sized Miniopterus in Africa, particularly a relatively long and straight tragus (Fig. 5). The tail the M. inflatus/M. africanus and the M. natalensis is almost exactly half that of the total length. The groups (Happold 2013a, 2013b). In general, M. wil - pelage is medium brown above and only slightly soni sp. nov. is similar in size to M. fraterculus and paler below. Individual hairs are unicoloured. The M. mossambicus, but significantly larger than the species is small-sized for a Miniopterus, being tiny M. minor (Tables 5 and 6); however, in multi- larger than M. minor, M. occidentalis and M. mos - dimensional morphospace based on craniodental sam bicus and smaller than M. natalensis and M. cf. measurements, it occupies a distinct morphospace inflatus; it does not overlap with any of these four (Fig. 4A and 4B). Furthermore, it can be readily dis- species based on forearm length (Table 3). How- tinguished from sympatric M. mossambicus by fore- ever, it is similar in size and colouring to that of arm length (FA > 43 mm in M. wilsoni sp. nov., < 43 M. fraterculus. It shares the distinct wash across mm in M. mossambicus); the two species do not the face but is peach-orange in colour compared overlap in this measurement (Table 3). Finally, all with a yellowish wash in M. fraterculus (Fig. 5B). specimens of M. wilsoni sp. nov. that have been ex- How ever, it is not known whether this is a character amined, have a peach-orange wash to the skin of the specific to these two species, nor whether all indi- face under the fur around the eyes, which is absent viduals of these two species have it. Miniopte- in M. mossambicus. The only other southern African rus wilsoni sp. nov. has a tragus that is mostly equal- Miniopterus species with this feature that we are ly wide along its entire length, except for a slight aware of is M. fraterculus, which has a yellowish constriction or inflection near its midpoint (Fig. 5B). wash. Morphologically, M. wilsoni sp. nov. most Unlike M. mossambicus, the tragus of M. wilsoni sp. resembles M. fraterculus, from which it differs on nov. lacks the slightly constricted base (also evident molecular grounds (K2P pairwise genetic distance in M. fraterculus) and has a rounded tip that is 10 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff

FIG. 5. Photographs of M. wilsoni sp. nov.: A — in flight showing typical features of the genus; the peach-orange wash to the face of this species is not obvious in this photograph (holotype, JAG 444); and B — showing the peach-orange wash to the face, and the relatively long and narrow tragus typical of the genus (paratype, JAG 445) (photographs by Piotr Naskrecki) broader than the rest of the tragus (Monadjem et species are widely distributed within Gorongosa al., 2013a). National Park at lower elevations (< 300 m a.s.l.), Craniodental characters. — The skull is gracile M. mossambicus and M.cf. inflatus. for a Miniopterus species. The rostrum is broad, and Echolocation calls of specimens captured and re- the braincase is rounded and high, typical for the leased at the same site as the holotype and paratype, genus of Miniopterus. The dentition of M. wilsoni and believed to represent the same species, had sp. nov. is I 2/3, C 1/1, P 2/3, M 3/3, which is typical mean peak frequency (± SD) of 56.8 kHz ± 0.96 of the genus. In the upper tooth row, the inner inci- kHz (range: 56–58 kHz, n = 4) which is slightly sor is larger than the outer one, and the anterior pre- higher than that reported for M. mossambicus from molar is relatively well developed (Fig. 7). The cra- Mount Namuli (Monadjem et al., 2013a), however nial and dental measurements of the holotype the recordings reported for the latter species were compared with a sample of other southern African taken from a location that is now included in the dis- Miniopterus species are shown in Table 6. tribution of M. wilsoni sp. nov. and with a different bat detector (Anabat). The peak frequency of two Distribution M. mossambicus in Gorongosa National Park were Currently, the only genetically confirmed speci- 60 and 61 kHz, while a single individual of the mens of this new species come from Mount Goron - larger M. cf. inflatus had a peak frequency of 49 gosa, central Mozambique. However, a number of kHz. Based on these limited recordings, it would ap- specimens collected in northern Mozambique and pear that the three species of Miniopterus in central southern Malawi without genetic identification are Mozambique differ in their echolocation calls. morphologically similar and therefore they are likely to be conspecific with M. wilsoni sp. nov. All DISCUSSION these specimens were collected on sizeable mountains, including Mount Gorongosa (elevation: 1,863 In this paper, we present new information on m a.s.l.), Mount Namuli (2,419 m a.s.l.), and Mount a population of Miniopterus from Mozambique Mabu (> 1,600 m a.s.l.) in Mozambique, and Zomba showing that it represents a new species to science. Massif (2,047 m a.s.l.) in Malawi (Fig. 1). This is the second new Miniopterus species to be described from this country in the past decade and fol- Biology lows a trend of new discoveries in Madagascar and Little is known about the biology of this species. It West Africa (Christidis et al., 2014; Goodman et al., appears to be associated with montane habitats, with 2015; Monadjem et al., 2019). Considering the re- most records taken from around 1,000 m a.s.l. Based cent phylogeny for Miniopterus by Demos et al. on our limited sampling, it appears to be the only (2019), it would appear that these new species de- Miniopterus species at higher elevations (> 900 m) on scriptions are just the tip of the iceberg, as predicted Mount Gorongosa. However, two other Miniopterus by Monadjem et al. (2013a). Cryptic diversity in southern African Miniopterus 11

FIG. 6. The cranium of M. wilsoni sp. nov. (holotype, JAG 444) showing dorsal, ventral, and lateral views of the neurocranium; and lateral view of the mandible (photographs by Piotr Naskrecki). The white grid lines are 1 mm apart 12 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff

28 The description of M. wilsoni sp. nov. brings the 12 18 6 8 36 total number of Miniopterus species known from Mozambique to four (Monadjem et al., 2010a,

Miniopterus 2013a). Within Mozambique, these four species appear to occupy different geographical regions or habitats. Miniopterus wilsoni sp. nov. is only known from large mountain ranges where it occurs at mid- 7.6 ± 0.72, 6.5–8.5, 14.3 ± 1.51, 11.0–18.0, 14.3 ± 1.51, 11.0–18.0, 10.8 ± 0.97, 8.3–11.8, 10.8 ± 0.97, 8.3–11.8, 7.6 ± 0.92, 6.0–10.0, 8.1 ± 0.94, 6.5–9.1, 5.0 ± 0.38, 4.1–5.6, and perhaps higher elevations. In contrast, M. mos - 8 30 16 18 15 sambicus and M. cf. inflatus appear to occupy lower

36 elevations, whereas M. natalensis is only known from the southern parts of the country. All four

± SD, range and sample size (in italics). species also occur beyond the borders of Mozam - 0 bique: with M. natalensis and M. cf. inflatus wide- spread in the region to the south and west of the country (Monadjem et al., 2010b); M. mossambicus on morphometrics, and other species of 44.1 ± 0.46, 43.5–44.9, 47.7 ± 1.04, 46.0–50.0, 45.5 ± 1.24, 43.4–47.8, 41.2 ± 0.77, 39.8–42.9, 43.4 ± 0.87, 41.4–44.2, 38.0 ± 1.00, 36–40, occurs further north into Tanzania and Kenya 3

5 (Demos et al., 2019); and, based on our morpholog- 15 11 13 36 ical analyses, M. wilsoni sp. nov. appears to also occur at Zomba Massif, Malawi. Its distribution, however, is not yet fully known and it may occur at other mountains in the region such as Mount Mulanje (Curran et al., 2012) or perhaps west into Zambia. 10.4 ± 1.20, 8.8–12.0, 12.5 ± 0.95, 11.6–13.5, 12.5 ± 0.95, 11.6–13.5, 9.6 ± 1.20, 7.5–12.0, 8.8 ± 1.35, 7.7–12.0, 9.8 ± 0.93, 8.0–11.0, 9.8 ± 0.93, 8.0–11.0, 10.3 ± 0.47, 10–11, 10.3 ± 0.47, 10–11, Based on the cyt b gene, M. wilsoni sp. nov. ap- 15

5 pears to have diverged from its closest living rela- 6 13 tive, M. minor, over the past one to two million 36 years. Divergence dates for Miniopterus species from other regions of continental African have not yet been reported, thwarting any attempts at recon- 1 structing the biogeography of this genus on the continent. However, in Madagascar, there was a pulse of 8.7 ± 0.58, 7.7–12.0, 10.5, 10.5 ± 0.99, 9.0–13.0, 8.4 ± 0.81, 7.0–9.5, 7.8 ± 0.99, 6.0–9.0, 7.8 ± 0.65, 7–9, diversification between 2–3 Mya, with some taxa 4 3 15 10 12 36 having diverged within the past million years

sp. nov. from Mount Gorongosa, Mozambique. Measurements presented as sp. nov. (Christidis et al., 2014). The appearance of M. wil - soni sp. nov., therefore, fits in with the observed trend of speciation in Malagasy Miniopterus. It is worth noting that M. cf. inflatus needs nam- M. wilsoni ing as it is not conspecific with M. inflatus from cen- 47.5 ± 3.51, 44–51, 54.0 ± 1.00, 53–55, 49.6 ± 4.91, 42–57, 47.0 ± 6.13, 38–56, 49.8 ± 1.80, 47–52, 41.3 ± 2.11, 37–46, 41.3 ± 2.11, tral Africa; in fact, these two taxa belong to different 5 16 13 13 clades (Monadjem et al., 2019). This species was 6 36 previously thought to be widely distributed across Africa, but recent genetic evidence suggests that it is restricted to Cameroon and Gabon, although populations in neighboring Congo, Democratic Republic of Congo and Central African Republic may also refer to this species. The West African population has 112.4 ± 6.46, 105–120, 112.4 109.5 ± 4.20, 102–116, 109.5 ± 4.20, 102–116, 102.5 ± 2.96, 97–108, 100.7 ± 5.68, 89–110, 100.7 ± 5.68, 89–110, 90.6 ± 3.21, 83–99, a restricted distribution centered on Mount Nimba (Liberia/Guinea) and surrounding uplands (Mona- djem et al., 2019), while the systematics of the pop-

sp. nov. ulations in southern and eastern Africa that were previously referred to as ‘M. inflatus’ needs resolv- 5. External measurements (mm) and mass (g) of inflatus ing. Furthermore, based on our morphometric analy- cf. Specimen or taxon length Total length Tail Hindfoot length Ear length Forearm length Body mass ABLE Holotype JAG 444Paratype JAG 445 95 92 44 45 9 9 10 12 44.2 43.7 7.8 7.5 All putative spec. 98.7 ± 4.41, 92–104, M. wilsoni M. T M. natalensis Measurements of the holotype and paratype new species, other specimens identified as belonging to species based Africa are shown for comparative purposes occurring in southern M. mossambicus M. fraterculus M. minor ses, M. minor minor from Tanzania is distinguishable TABLE 6. Cranial and dental measurements (mm) of specimens of M. wilsoni sp. nov. from Mount Gorongosa, Mozambique. Measurements presented as 0 ± SD, range and sample size southern in African diversity Cryptic (in italics). Measurements of the holotype and paratype of the new species, other specimens identified as belonging to the new species based on morphometrics, and other species of Miniopterus occurring in southern Africa are shown for comparative purposes

Specimen or taxon GSKL ZYGO POB MAST GBW MAND M. wilsoni sp. nov. Holotype JAG 444 15.10 8.23 3.88 8.23 7.69 10.61 Paratype JAG 445 14.86 8.00 3.67 8.28 7.58 10.54 All specimens 14.69 ± 0.25, 14.30–15.10, 13 8.01 ± 0.28, 7.50–8.58, 13 3.73 ± 0.16, 3.45–4.04, 13 88.06 ± 0.25, 7.57–8.43, 13 7.52 ± 0.19, 7.30–7.76, 13 10.66 ± 0.31, 10.02–11.16, 12 M. cf. inflatus 16.47 ± 0.17, 16.27–16.71, 5 9.01 ± 0.18, 8.80–9.19, 5 4.17 ± 0.11, 4.08–4.36, 5 8.88 ± 0.11, 8.76–8.97, 5 8.26 ± 0.20, 8.04–8.51, 5 12.20 ± 0.41, 11.53–12.63, 5 M. natalensis 15.44 ± 0.19, 15.03–15.69, 16 8.56 ± 0.14, 8.23–8.77, 16 4.09 ± 0.12, 3.83–4.29, 16 8.57 ± 0.15, 8.28–8.90, 16 8.00 ± 0.15, 7.72–8.21, 16 11.41 ± 0.25, 10.74–11.74, 16 M. mossambicus 14.71 ± 0.24, 14.38–15.20, 17 8.06 ± 0.18, 7.85–8.40, 8 3.77 ± 0.13, 3.60–4.14, 17 8.09 ± 0.20, 7.60–8.50, 17 7.56 ± 0.18, 7.20–8.00, 17 10.84 ± 0.29, 10.01–11.22, 17 M. fraterculus 14.37 ± 0.13, 14.05–14.60, 15 7.97 ± 0.13, 7.79–8.20, 15 3.75 ± 0.10, 3.47–3.87, 15 8.12 ± 0.10, 7.95–8.34, 15 7.46 ± 0.14, 7.24–7.70, 15 10.39 ± 0.25, 10.03–10.73, 15 M. minor 14.07 ± 0.25, 13.60–14.57, 21 7.62 ± 0.16, 7.35–7.94, 21 3.48 ± 0.08, 3.32–3.62, 21 7.69 ± 0.17, 7.33–8.03, 21 7.15 ± 0.15, 6.89–7.42, 21 9.48 ± 0.20, 9.05–9.80, 21 M. occidentalis 13.73 ± 0.17, 13.45–14.00, 8 7.46 ± 0.15, 7.30–7.80, 8 3.59 ± 0.11, 3.50–3.75, 8 7.43 ± 0.14, 7.30–7.70, 8 7.04 ± 0.14, 6.85–7.30, 8 10.03 ± 0.15, 9.70–10.20, 8

TABLE 6. Extended

Specimen or taxon C–M3 C–C M3–M3 i–m3 LWMOLS

M. wilsoni sp. nov. Miniopterus Holotype JAG 444 5.63 4.16 6.00 6.95 5.24 Paratype JAG 445 5.57 4.03 6.00 6.90 5.19 All specimens 5.55 ± 0.19, 5.20–5.84, 13 4.09 ± 0.14, 3.90–4.34, 13 5.90 ± 0.19, 5.45–6.13, 13 6.91 ± 0.19, 6.62–7.31, 13 5.27 ± 0.19, 4.94–5.54, 13 M. cf. inflatus 6.39 ± 0.09, 6.28–6.52, 5 4.78 ± 0.14, 4.69–5.02, 5 6.76 ± 0.07, 6.72–6.89, 5 7.87 ± 0.18, 7.70–8.12, 5 6.06 ± 0.21, 5.90–6.35, 5 M. natalensis 5.74 ± 0.10, 5.58–5.93, 16 4.42 ± 0.14, 4.08–4.61, 16 6.29 ± 0.15, 5.87–6.45, 16 7.54 ± 0.17, 7.26–7.93, 16 5.97 ± 0.18, 5.68–6.23, 16 M. mossambicus 5.52 ± 0.16, 5.27–5.87, 17 4.11 ± 0.13, 3.94–4.40, 17 6.00 ± 0.13, 5.84–6.21, 16 7.10 ± 0.19, 6.90–7.40, 7 5.65 ± 0.18, 5.40–5.80, 7 M. fraterculus 5.30 ± 0.09, 5.10–5.43, 15 3.90 ± 0.08, 3.78–4.07, 15 5.65 ± 0.11, 5.50–5.85, 15 6.95 ± 0.16, 6.70–7.24, 15 5.44 ± 0.16, 5.10–5.66, 15 M. minor 5.25 ± 0.09, 5.04–5.40, 21 3.95 ± 0.09, 3.79–4.09, 21 5.90 ± 0.19, 5.45–6.13, 13 6.52 ± 0.13, 6.14–6.88, 21 4.97 ± 0.09, 4.74–5.14, 21 M. occidentalis 5.16 ± 0.09, 5.00–5.30, 8 3.72 ± 0.14, 3.50–3.95, 8 6.76 ± 0.07, 6.72–6.89, 5 6.57 ± 0.25, 6.35–7.00, 8 5.06 ± 0.21, 4.90–5.50, 8 13 14 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff

FIG. 7. Upper teeth of M. wilsoni sp. nov. (holotype, JAG 444) showing: A — the long outer incisors which are almost the same length as the inner incisors; and B — the relatively large anterior premolar (photographs Piotr Naskrecki)

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Received 07 December.2019, accepted 31 May 2020

Associate Editor: Wiesław Bogdanowicz Cryptic diversity in southern African Miniopterus 17 13.9 16.05 9.05 3.99 8.71 8.06 6.10 4.52 6.37 11.83 7.40 5.73 44.0 7.0 14.42 8.09 3.85 8.12 7.24 5.42 3.89 5.52 10.03 6.91 5.37 44.0 6.5 14.37 8.15 3.81 8.25 7.62 5.34 3.83 5.54 10.62 6.90 5.32 11.0 11.0 47.5 16.15 9.15 4.11 8.90 8.40 6.38 4.86 6.54 12.09 8.02 6.17 10.0 10.0 46.4 15.83 9.05 3.98 8.76 8.37 6.21 4.63 6.71 11.95 7.76 5.92 90 37 8 10 39 4.9 14.3 7.69 3.56 7.79 7.33 5.31 4.09 5.66 9.61 6.58 5.10 99 46 8 10 40 5.6 14.38 7.72 3.47 7.59 7.18 5.40 4.02 5.67 9.72 6.68 5.05 96 45 9.0 8.0 43.5 14.25 7.84 3.73 7.97 7.36 5.39 3.92 5.71 10.14 6.97 5.32 88 38 7 11 37 5.3 13.98 7.35 3.47 7.60 7.04 5.28 3.82 5.36 9.50 6.46 4.91 95 40 8.0 8.0 41.4 14.38 8.06 3.79 8.14 7.45 5.33 4.00 5.78 10.23 7.08 5.66 84 38 7 10 36 4.4 13.60 7.46 3.50 7.53 6.89 5.10 3.79 5.37 9.05 6.14 4.87 93 43 7 11 38 5.2 13.94 7.39 3.45 7.33 7.06 5.18 3.83 5.53 9.35 6.45 4.96 98 44 8.0 8.0 41.9 14.05 7.81 3.87 7.95 7.42 5.26 3.83 5.51 10.32 6.85 5.63 89 43 7 11 39 5 13.89 7.47 3.42 7.56 7.15 5.28 3.86 5.49 9.39 6.55 4.97 89 35 7.0 44.0 14.37 7.84 3.69 8.17 7.33 5.34 3.85 5.68 10.73 6.77 5.43 89 43 8 10 39 4.9 14.29 7.73 3.37 7.76 7.07 5.32 4.06 5.68 9.76 6.65 5.05 96 40 7 10 39 5.2 96 43 9 11 38 4.8 97 38 8.9 8.1 43.6 8.6 14.30 7.88 3.67 8.11 7.42 5.34 4.00 5.72 10.59 7.21 5.50 94 41 8 10 37 4.7 14.16 7.76 3.57 7.81 7.22 5.21 4.04 5.57 9.60 6.58 4.86 93 44 8 10 39 5 14.21 7.68 3.50 7.89 7.32 5.24 3.99 5.63 9.51 6.55 4.99 93 45 8 10 40 4.8 14.57 7.74 3.43 8.03 7.42 5.38 4.06 5.76 9.80 6.67 5.14 96 45 8 10 38 5.1 14.40 7.79 3.60 7.90 7.03 5.25 4.02 5.66 9.61 6.62 4.97 89 44 7 10 37 4.6 13.77 7.57 3.32 7.62 7.02 5.14 3.95 5.36 9.38 6.36 4.91 94 42 8 10 37 4.7 14.26 7.59 3.56 7.77 7.12 5.35 3.93 5.53 9.65 6.63 4.99 111 55 10.5 13.5 46.9 14.0 16.71 9.11 4.36 8.97 8.42 6.52 5.02 6.72 12.63 8.12 6.35 118 46.8 16.0 16.44 9.12 4.16 8.96 8.51 6.39 4.71 6.73 12.24 7.98 5.92 110 53 12.0 43.5 14.50 8.10 3.80 8.15 7.60 5.15 3.90 5.60 10.50 6.90 5.20 105 53 11.6 46.6 16.27 8.80 4.18 8.76 8.15 6.35 4.69 6.73 11.53 7.74 5.90 108 54 12.5 45.7 16.56 9.19 4.08 8.94 8.20 6.42 4.74 6.89 12.42 7.70 6.21 120 47.3 15.5 16.39 8.83 4.09 8.77 8.04 6.28 4.75 6.72 12.20 7.82 5.93 102 44.0 8.5 14.32 7.86 3.81 8.07 7.40 5.26 3.78 5.60 10.16 6.79 5.40 103 44.1 8.5 14.52 7.96 3.69 7.98 7.32 5.34 3.97 5.61 10.15 7.24 5.52 100 46 8.0 9.5 43.4 14.40 7.96 3.75 8.09 7.45 5.43 4.07 5.85 10.09 6.92 5.42 106 52 9.0 43.5 14.60 8.20 3.70 8.15 7.70 5.10 3.80 5.50 10.70 6.70 5.10 103 53 9.5 8.0 44.0 7.9 14.41 7.97 3.84 8.34 7.69 5.25 3.91 5.80 10.63 6.96 5.51 107 43 9.0 10.0 44.2 9.0 14.32 8.01 3.75 8.20 7.55 5.30 3.86 5.72 10.67 7.18 5.61 102 56 7.7 42.2 9.1 14.27 7.79 3.47 8.15 7.34 5.32 3.86 5.65 10.36 6.94 5.62 Y Y X Y Y Y X X Y Y X X Y Y X Y Y X X X X Y X Y Y Y Y Y Y Y Y Y Y Y Y Y X Y specimens present in southern Africa and used in this study specimens present in southern Miniopterus FM 198004 Tanzania FM 198003 Tanzania DM 8366Africa South DM 7091Africa South DM 7090Africa South DM 10140Africa South DM 9847 Mozambique DM 8779 Mozambique DM 8556 Mozambique DM 8527 Mozambique DM 8383Africa South DM 8359Africa South DM 8358Africa South FM 198100 Tanzania DM 7034Africa South FM 198099 Tanzania DM 7033Africa South FM 198098 Tanzania DM 7032Africa South FM 198097 Tanzania DM 5604Africa South FM 198035 Tanzania DM 14047Africa South FM 198034 Tanzania DM 14046Africa South FM 198011FM 198033 Tanzania Tanzania – 14.35 7.88 3.52 7.92 7.29 5.26 3.92 5.70 9.76 6.56 5.00 DM 10034Africa South FM 198010 Tanzania BM 1905.5.7.22Africa South FM 198009 Tanzania BM 1905.5.7.18 HolotypeAfrica South FM 198008 Tanzania DM 8050 Eswatini FM 198007 Tanzania DM 8048 Eswatini FM 198006 Tanzania DM 5785 Eswatini FM 198005 Tanzania Taxon Museum No Type Country Sex Total Tail HF/cu Ear FA Mass GSKL ZYGO POB MAST GBW C-M3 C1-C1 M3-M3 MAND i1-m3 LWMOLS External and craniodental measurements of PPENDIX minor minor inflatus inflatus inflatus inflatus inflatus inflatus inflatus inflatus fraterculus fraterculus fraterculus minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor fraterculus minor A 18 A. Monadjem, J. Guyton, P. Naskrecki, L. R. Richards, and A. S. Kropff 15.20 8.40 3.80 8.50 8.00 5.40 4.00 5.90 10.80 7.10 5.40 14.50 7.85 3.70 7.60 7.40 5.50 3.95 5.85 10.70 6.90 5.50 14.80 8.10 3.70 8.00 7.25 5.60 4.40 6.10 11.20 7.40 5.80 14.70 7.90 3.60 7.80 7.20 5.60 4.10 6.00 11.20 7.00 5.80 15.00 8.10 3.70 8.00 7.60 5.50 4.20 6.10 11.00 7.00 5.80 90 41 8 11 39 5.3 14.13 7.69 3.49 7.58 6.92 5.23 3.96 5.59 9.51 6.55 4.99 9089 4090 41 891 41 891 43 10 791 42 10 37 988 41 11 5.2 37 889 42 10 5.5 13.82 37 783 41 11 4.5 13.86 39 7.46 790 41 11 4.8 38 7.50 3.46 891 41 11 4.8 13.86 38 3.37 7.72 788 41 10 4.9 13.88 37 7.59 7.44 8 7.1789 39 10 5.2 14.05 39 7.46 3.52 8 6.96 5.1588 37 11 4.6 36 7.94 3.50 7.76 8 5.04 3.9690 39 10 4.1 38 3.62 7.61 8 7.07 3.98 5.5287 41 10 5.5 13.87 38 7.76 7 7.20 5.25 5.3892 40 10 5.4 37 7.59 9.31 7 7.37 5.18 3.9591 41 10 5.1 38 9.13 3.41 6.42 7 5.24 3.80 5.5690 41 11 5.5 39 6.36 7.54 8 4.00 5.5291 41 10 4.89 5.3 38 9.47 8 7.02 5.65 41 10 4.74 5.3 38 6.55 9.3 9 5.38 10 5.1 38 9.30 9 4.02 6.46 10 5.00 4.8 38 6.50 5.51 10 4.7 38 5.06 4.96 4.3 39 9.42 4.2 6.60 5.00 97 48 7.0 9.0 40.6 7.0 14.47 8.10 3.64 8.10 7.63 5.50 4.03 5.89 11.00 6.82 5.20 99 47 9.0 10.0 43.0 9.0 14.67 8.09 3.71 8.21 7.60 5.87 4.16 6.16 11.07 6.98 5.47 111 47 10.0 8.0 43.9 11.0 15.45 8.50 4.25 8.58 8.05 5.69 4.52 6.30 11.37 7.50 6.04 112 52115 53115 9.0 57 11.0 10.0 45.5 10.0 11.0 11.4112 45.4 11.0 15.69 8.0 54112 15.51 45.9 8.77 55 10.0 9.6 8.66 9.0 11.0 3.97 15.50 44.5 4.22 10.0 11.5 46.6 8.90 8.54 10.5 8.58 15.53 8.20 15.45 4.01 8.60 8.17 5.78 8.71 8.68 5.62 4.29 4.51 4.19 8.21 4.61 8.74 6.28 5.93 8.68 6.45 8.15 4.46 11.62 8.06 5.85 11.54 7.56 6.31 5.73 4.43 7.48 4.33 11.60 6.19 6.45 6.11 6.17 7.79 11.60 11.00 7.50 6.23 7.31 5.94 5.82 100 47103 50 6.0 10.0 6.0 41.0 10.0 7.6 41.0 6.7 15.12 7.92 3.73 8.18 7.53 5.78 4.29 5.96 11.22 7.34 5.77 101 52108 8.0 50 10.0 41.7 8.0 7.0 14.55 8.0 42.0 8.29 8.0 3.74 14.77 8.03 8.08 7.76 3.65 5.62 8.32 4.04 7.64 5.94 5.56 4.00 10.91 6.67 5.84 10.48 5.31 6.71 5.44 109 56 11.0 10.0 45.8 8.3 15.25 8.58 4.10108 49106 8.50 44 11.0 8.07 10.0 10.0 45.7 5.85 11.0 11.0 43.4 4.32 15.43 11.0 15.21 6.29 8.70 8.50 4.08 11.49 4.05 7.59 8.53 8.40 7.91 5.74 7.80 5.68 5.76 4.52 4.50 6.43 6.34 11.24 7.54 11.33 7.70 5.85 5.82 X Y Y Y X X Y X X X Y X X Y X X Y Y X Y X Y Y Y X X X Y X X X Y X Y Y Y Y Y X X Y FM 198101 Tanzania FM 198102 Tanzania FM 198103FM 198105FM 198106FM 198107FM 198108FM 198159FM 198160 Tanzania FM 198161 Tanzania FM 198162 Tanzania FM 198163 Tanzania FM 198164 Tanzania FM 198165 Tanzania FM 198166 Tanzania FM 198167 Tanzania FM 198168 Tanzania FM 198169 Tanzania FM 198170 Tanzania FM 198171 Tanzania FM 198172 Tanzania BM 1987.1178 Tanzania Tanzania Tanzania Tanzania Tanzania Malawi Tanzania FMNH Paratype Mozambique FMNH 213651BM 1968.1014 Holotype Mozambique Zambia DM 13913DM 13914DM 13936 Mozambique Mozambique Mozambique BM 1968.1015 Zambia BM 1968.1017 Zambia BM 1968.1018 Zambia DM 8028DM 8038DM 8051DM 8433DM 8436 Eswatini Eswatini Eswatini Eswatini Eswatini DM 8430 Eswatini DM 3687DM 7190DM 7917 Zimbabwe Eswatini DM 8438 Eswatini Eswatini . Continued Taxon Museum No Type Country Sex Total Tail HF/cu Ear FA Mass GSKL ZYGO POB MAST GBW C-M3 C1-C1 M3-M3 MAND i1-m3 LWMOLS PPENDIX minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor minor mossambicus mossambicus mossambicus mossambicus mossambicus mossambicus mossambicus mossambicus mossambicus mossambicus natalensis natalensis natalensis natalensis natalensis natalensis mossambicus natalensis natalensis natalensis A Cryptic diversity in southern African Miniopterus 19 13.6013.70 7.3013.45 7.40 3.60 7.40 3.50 7.3013.70 3.70 7.40 7.0013.90 7.50 7.30 6.90 5.1514.00 7.50 3.50 6.85 5.25 3.7013.70 7.80 3.70 7.40 5.10 3.70 5.6014.30 7.40 3.75 7.50 7.00 3.70 5.40 10.0014.70 7.50 3.50 7.70 7.15 5.10 5.50 6.35 10.10 8.05 3.70 7.30 7.30 5.15 3.75 6.40 10.20 3.80 7.80 7.00 5.20 3.95 4.90 5.60 6.70 8.00 7.30 5.00 3.85 4.90 5.60 9.70 7.60 5.30 3.60 5.00 5.50 10.15 6.35 5.20 3.90 5.40 6.50 10.10 4.10 5.90 6.85 10.00 5.10 5.50 5.90 7.00 4.90 10.80 5.00 7.10 6.90 5.40 5.30 44.9 14.70 7.84 3.62 8.02 7.58 5.69 4.12 5.87 10.82 6.78 5.10 7.7 8.8 43.7 14.37 7.89 3.69 7.95 7.30 5.60 4.09 5.94 10.02 6.80 4.94 92 45 9.0 12.0 43.7 7.5 14.86 8.00 3.67 8.28 7.58 5.57 4.03 6.00 10.54 6.90 5.19 95 44 9.0 10.0 44.2 7.8 15.10 8.23 3.88 8.23 7.69 5.63 4.16 6.00 10.61 6.95 5.24 99 51 9.0 10.0 44.0 6.5 14.68 8.10 3.80 8.11 7.41 5.49 4.03 6.13 10.79 6.77 5.45 111 47 13.0 12.0 47.5 15.57 8.68 3.99 8.61 8.12 5.85 4.54 6.40 11.57 7.65 6.11 116 45.4 10.6 15.33 8.39 4.09 8.62 7.87 5.74 4.45 6.35 11.40 7.37 5.75 104 43.5 8.0 14.43 7.88 3.45 7.57 7.30 5.55 3.90 5.45 10.50 6.62 5.08 101 44.5 8.5 14.90 8.58 4.04 8.43 7.76 5.84 4.34 6.07 11.16 7.31 5.43 101 50 9.0 11.0 44.0 7.0 14.83 8.05 3.70 8.21 7.76 5.67 4.25 5.78 10.67 6.95 5.54 102 45 11.0 9.0 45.8 15.17 8.46 4.12 8.28 7.96 5.60 4.16 6.11 11.30 7.49 6.11 105 43 10.0 10.0 46.0 15.53 8.44 4.18 8.52 7.87 5.74 4.41 6.19 11.52 7.26 5.68 107 48 11.0 9.0 47.8 11.5 15.66 8.63 4.11 8.63 8.01 5.81 4.50 6.39 11.74 7.45 6.18 104 42 9.0 7.5 43.5 15.03 8.23 3.83 8.38 7.72 5.58 4.08 5.87 10.74 7.93 5.88 106 52 10.0 10.6 45.8 11.8 15.66 8.55 4.02 8.50 7.85 5.70 4.44 6.37 11.56 7.49 6.00 X Y Y Y Y Y Y Y Y Y Y X Y X Y X X Y Y X X Y Y Y MNHN 1985-628 MNHN 1985-629MNHN 1985-631MNHN 1985-637MNHN 1985-647 CAR MNHN 1985-649 CAR MNHN 1985-651 CAR BM 1954.866 CAR CAR CAR CAR – DRC 13.75 7.40 3.50 7.50 7.10 5.30 3.50 5.10 10.00 6.40 5.20 DM 8370Africa South DM 8369Africa South DM 6803Africa South DM 5729Africa South DM 5529Africa South DM 4998Africa South BM 1848.6.12.19 HolotypeDM 13288Africa South –Africa South 45.7 5.60 3.90 5.95 10.60 6.90 6.00 . Continued sp. nov. JAG445 Paratype Mozambique sp. nov. JAG444 Holotype Mozambique sp. nov. DM 9840 Mozambique sp. nov. DM 8520 Mozambique sp. nov. DM 8484 Mozambique sp. nov. DM 14852 Mozambique sp. nov. DM 14847 Mozambique sp. nov. BM 1987.1156sp. nov. BM 1987.1177sp. nov. DM 10836 Malawi Malawi Mozambique Taxon Museum No Type Country Sex Total Tail HF/cu Ear FA Mass GSKL ZYGO POB MAST GBW C-M3 C1-C1 M3-M3 MAND i1-m3 LWMOLS PPENDIX wilsoni wilsoni wilsoni wilsoni wilsoni wilsoni wilsoni occidentalis occidentalis occidentalis occidentalis occidentalis occidentalis occidentalis occidentalis wilsoni wilsoni wilsoni natalensis natalensis natalensis natalensis natalensis natalensis natalensis natalensis A