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Joffrin, Léa, Goodman, Steven M., Wilkinson, David A., Ramasindrazana, Beza, Lagadec, Erwan, Gomard, Yann, Minter, Gildas Le, Santos, Andréa Dos, Schoeman, M. Corrie, Sookhareea, Rajendraprasad, Tortosa, Pablo, Julienne, Simon, Gudo, Eduardo S., Mavingui, Patrick, Lebarbenchon, Camille (2020): Supplementary information: Bat coronavirus phylogeography in the Western Indian Ocean. Scientific Reports (supplement) 10 (6873): 92-104, DOI: http://doi.org/10.5281/zenodo.3768642, URL: https://static-content.springer.com/esm/art%3A10.1038%2Fs41598-020-63799-7/MediaObjects/41598_2020_63799_MOESM1_ESM.pdf, {"references":["1. Melade J, McCulloch S, Ramasindrazana B, Lagadec E, Turpin M, Pascalis H, et al. Serological Evidence of Lyssaviruses among Bats on Southwestern Indian Ocean Islands. PLoS One. 2016;11(8):e0160553.","2. Wilkinson DA, Melade J, Dietrich M, Ramasindrazana B, Soarimalala V, Lagadec E, et al. Highly Diverse Morbillivirus-Related Paramyxoviruses in Wild Fauna of the Southwestern Indian Ocean Islands: Evidence of Exchange between Introduced and Endemic Small Mammals. J Virol. 2014;88(15):8268-77.","3. Gomard Y, Dietrich M, Wieseke N, Ramasindrazana B, Lagadec E, Goodman SM, et al. Malagasy bats shelter a considerable genetic diversity of pathogenic Leptospira suggesting notable host-specificity patterns. FEMS Microbiol Ecol. 2016;92(4):fiw037.","4. Melade J, Wieseke N, Ramasindrazana B, Flores O, Lagadec E, Gomard Y, et al. An ecoepidemiological study of Morbilli-related paramyxovirus infection in Madagascar bats reveals host-switching as the dominant macro-evolutionary mechanism. Sci Rep. 2016;12(6):23752.","5. Hoarau F, Minter G Le, Joffrin L, Schoeman MC, Lagadec E, Ramasindrazana B, et al. Bat Astrovirus in Mozambique. Virol J. 2018;15(104):1-5."]}

Joffrin, Léa, Goodman, Steven M., Wilkinson, David A., Ramasindrazana, Beza, Lagadec, Erwan, Minter, Gildas Le, Santos, Andréa Dos, Schoeman, M. Corrie, Sookhareea, Rajendraprasad, Tortosa, Pablo, Julienne, Simon, Gudo, Eduardo S., Mavingui, Patrick, Lebarbenchon, Camille (2020): Bat coronavirus phylogeography in the Western Indian Ocean. Scientific Reports 10 (6873): 1-11, DOI: 10.1038/s41598-020-63799-7

RHINOLOPHUS LOBATUS PETERS, 1852 PETERS’ S HORSESHOE BAT Synonyms: R. l. angolensis (Seabra, 1898), Angola, Hahna. Type locality: Mozambique, Tete (= Tete Province), Sena village, south bank of the Zambesi River, - 15.679° S, 33.809° E. Syntypes: ZMB 375 (female, poor skin only), ZMB 2496 (female, skin in alcohol, skull extracted, broken), ZMB 24922 (adult male, complete skeleton), ZMB 24927 (female, complete skeleton), collected between 1843 and 1847 by W. C. H. Peters from Mozambique, Tette (= Tete Province), Sena on the south bank of the Zambesi River, - 15.679° S, 33.809° E. Referred specimens having molecular identifications: FMNH 228936 (SMG 18959), male, 228937 (SMG 18988), female, 228938 (SMG 18989), male, 228939 (SMG 18990), male, 228940 (SMG 18991), female, 228941 (SMG 18992), male, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Chihalatan Cave, 38.2 km Eof Inhassoro, - 21,671° S, 34,864° E. FMNH: 228947 (SMG 19022), female, 228954 (SMG 19029), female, 228965 (SMG 19040), female, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Mozambique, Inhambane Province, Malashane Cave, 39.1 km Eof Inhassoro, - 21,668° S, 34,847° E, and situated Referred specimens having non - molecular identifications: DM 13894 (female), 13927 (male), 13938 (unknown sex), 13942 (unknown sex), 13943 (unknown sex), 13944 (unknown sex), 13945 (male), 14529 (male), 14531 (male), 14532 (female), collected by A. Monadjem, G. le Minter and E. Lagadec in July 2015 from Mozambique, Sofala Province, Gorongosa National Park. TMSA 49114 (female), TMSA 49117 (male), collected from Mozambique, Sofala Province, Gorongosa National Park by A. J. Guyton in July 2015 from Chitengo Camp and Cheringoma Plateau respectively. TMSA 14653 (female), 14655 (unknown sex), 14656 (female), 14657 (female), 14662 (male), 14663 (male), 14664 (female), collected 28 – 29 July 1964 from Mozambique, Tete Province, Muchena, - 15.679° S, 33.809° E. Incertae sedis: DM 8575 (male) from Mozambique, Cabo Delgado Province, Pemba Island (- 13.006° S, 40.524° E) and DM 8574 (female), from Mozambique, Sofala Province, Chinizuia Forest (- 18.977° S, 35.052° E), both collected by A. Monadjem in June 2006. TMSA 49115 (male), TMSA 49118 (male), collected from Mozambique, Sofala Province, Gorongosa National Park by A. J. Guyton in July 2015 from Chitengo Camp and Cheringoma Plateau, respectively. Etymology: The Latin word lobatus means lobed, perhaps referring to the general shape of the noseleaf (Fig. 13). Re-diagnosis and comparisons: This taxon is clearly phylogenetically distinct on Cytb gene sequences (see also: Dool et al., 2016) from topotypic landeri from West Africa and seems to be more closely affiliated with the capensis group of Csorba et al. (2003), i.e. R. capensis, R. denti, R. simulator and R. swinnyi (Fig. 2). Specimens from Chihalatan and Malashane (N = 4) had a mean echolocation call peak frequency of 106.8 ± 0.4 kHz (Table 2), close to the 107 kHz generally reported for southern Africa ‘ landeri ’ (Monadjem et al., 2010). This contrasts with a mean maximum frequency of 104.3 ± 0.42 kHz (N = 2) recorded for topotypic West African (Liberian) R. landeri, making a peak frequency of c.107 kHz a possible diagnostic criterion for R. lobatus. However, a wider range (102–108 kHz) and slightly lower mean frequency of 105 kHz was reported for 20 individuals at Gorongosa (Table 2). This variation could be indicative of further undetected cryptic speciation in this taxon, as also indicated by the very wide variation in baculum and noseleaf characters, discussed in the following paragraph. Although not easily distinguished on body or skull size (Table 2; Fig. 4), this taxon differs from topotypic R. landeri by displaying a prognathic rostrum (LMs 1–3), a shortened braincase that extends posteriorly (LMs 10–14) and a broader nasal inflation (LMs 16–18) (Figs 5, 6). Specimens of cf. lobatus from Pemba (Mozambique) had a shorter foramen magnum (LMs 9–10), a narrower jugal process (LM 3–4) and a slightly broader nasal inflation (LMs 16–18), than other Mozambique animals. The same individual from Pemba possessed a distinctive robust baculum (Fig. 8) and noseleaf (Fig. 9) having a markedly hastate lancet and pronounced posterior lobe of the connecting process. However, pending molecular data, we here refer them to lobatus incertae sedis. Although genetically clearly assigned to lobatus, individuals from Gorongosa National Park are somewhat smaller in external (but not cranial) measurements (Table 2), and also display unique noseleaf and bacular morphologies. The noseleaf of a Gorongosa individual has a small hastate lancet and bluntly pointed tip, and a relatively small and more erect connecting process with semi-symmetrical lobe (Figs 9, 13), while its baculum is highly divergent from all other forms considered in our study (Fig. 8), having a remarkable short baculum with wide shaft and very small base compared with landeri from West Africa, as well as cf. lobatus from Malashane and Chihalatan Caves. Given the clear evidence for close genetic identity in our study between Malashane and Chihalatan Caves and Gorongosa National Park for animals referred to R. lobatus, we provisionally regard this variation in noseleaf and baculum structure to represent polymorphic traits in this species. It is possible that these morphological types represent good species between which introgression has occurred. Further analysis with additional nuclear sequences is necessary to test this hypothesis. There is also the possibility that these could represent subadults with bacula that are not fully ossified, as detailed in R. adami described by Kock et al. (2000). Male R. lobatus from Chihalatan and Malashane Caves, as well as Gorongosa National Park, typically possessed a dark brown apical tuft of stiff hairs characteristic of R. landeri (Fig. 14; Csorba et al., 2003; Monadjem et al., 2010). This convergent character could be one of the contributing factors that have led to the historical misidentification of the species with R. landeri. Distribution and biology: As anticipated by Monadjem et al. (2010), who considered that the southern (and possibly eastern) African savannah-occurring lobatus might prove to be distinct from the West African forest-occurring landeri, we here refer all southern African specimens to lobatus (Fig. 11) [see appendix of Monadjem et al. (2010) for a full list of localities]. We agree with Monadjem et al. (2010) that R. angolensis Seabra, 1989 from western Angola may merit specific status. Nevertheless, in the absence of the type series that was destroyed in the Lisbon fire of 1978, resolution of its status must await a detailed revision, including molecular evidence with designation of new type material. Likewise, the status of east and central African populations must await further studies, including molecular data. Until further evidence becomes available, we suggest that it is prudent to use the name lobatus for all savannah populations in southern and east Africa., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1270-1272, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Dool SE, Puechmaille SJ, Foley NM, Allegrini B, Bastian A, Mutumi GL, Maluleke TG, Odendaal LJ, Teeling EC, Jacobs DS. 2016. Nuclear introns outperform mitochondrial DNA in inter-specific phylogenetic reconstruction: lessons from horseshoe bats (Rhinolophidae: Chiroptera). Molecular Phylogenetics and EVolution 97: 196 - 212.","Csorba G, Ujhelyip P, Thomas N. 2003. Horseshoe bats of the aeorld (Chiroptera: Rhinolophidae). Shropshire: Alana books.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of Southern and South-central Africa: a biogeographic and taxonomic synthesis. Johannesburg: Wits University Press.","Kock D, Csorba G, Howell KM. 2000. Rhinolophus maendeleo n. sp. from Tanzania, a horseshoe bat noteworthy for its systematics and biogeography. Senckenbergiana biologica 80: 233 - 239."]}

RHINOLOPHUS RHODESIAE ROBERTS, 1946 ROBERTS' S HORSESHOE BAT Synonyms: None. Holotype: TMSA 1325, adult female, collected by A. Roberts on 16 August 1913. Type locality: 'Southern Rhodesia' (= Zimbabwe), Bezwe River, tributary of 'Wanetsi' (= Nuanetsi) River, −21.500° S, 31.167° E. Referred specimens having molecular identifications: FMNH 228942 (SMG 19017), female, 228943 (SMG 19018), male, 228944 (SMG 19019), female, 228945 (SMG 19020), male, 228946 (SMG 19021), female, 228948 (SMG 19023), male, 228949 (SMG 19024), female, 228950 (SMG 19025), female, 228951 (SMG 19026), female, 228952 (SMG 19027), male, 228953 (SMG 19028), male, 228955 (SMG 19030), male, 228957 (SMG 19032), female, 228958 (SMG 19033), female, 228959 (SMG 19034), female, 228960 (SMG 19035), female, 228961 (SMG 19036), male, 228962 (SMG 19037), male, 228964 (SMG 19039), female, all collected on 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Inhambane Province, Malashane Cave, 39.1 km Efrom Inhassoro, −21,668° S, 34,847° E, and situated Referred specimens having only morphological identifications: DM 7080, adult male, KwaZulu-Natal Province, Hlabeni Forest Reserve, −29.933° S, 29.766° E collected by D. Forbes on 29 July 2000; DM 12007 (adult male); DM 14034 (adult male), collected by M. C. Schoeman from KwaZulu-Natal Province, Pietermaritzburg, Ferncliff Nature Reserve, Ferncliff Cave, −29.550° S, 30.320° E; DM 11270 (female), 11271 (male), 11272 (female), 11273 (female), 11275 (male), all collected by S. Stoffberg from Chihalatan Cave, 38.2 km Eof Inhassoro, Inhambane Province, Mozambique −21.671° S, 34.864° E;. DM 11275 collected on 8 August 2006 while the other specimens were collected on 3 September 2007; FMNH 228956 (SMG 19031), collected 2 May 2015 by S. M. Goodman, M. C. Schoeman and G. le Minter from Mozambique, Inhambane Province, Malashane Cave, 39.1 km E from Inhassoro, −21,668° S, 34,847° E; DM 13450 (female), DM13451 (female), collected on 8 May 2012 by J. Bayliss at Mozambique, Niassa Province, Mount Mecula, −12.068° S, 37.662° E. Etymology: The name refers to the location in Southern Rhodesia (now Zimbabwe) where the type specimen was collected. Re-diagnosis and description: Roberts (1946) described this subspecies as being slightly smaller than the nominate R. s. swinnyi based on slightly smaller body size, slightly longer tail, smaller ears and its bright ochraceous colour. Since the last-mentioned character is known to be an environmentally induced effect in many cave-dwelling bat species, it does not serve as a diagnostic character. In our analysis, molecular evidence closely matched our series from Chihalatan and Malashane Caves with Genbank sequences from the extreme northern South Africa (Pafuri), Zimbabwe (Dambanzara) and Zambia (Kalenda and Shimalala Caves). Since these localities encompass the type locality of rhodesiae (Bezwe River in Zimbabwe), and Pafuri is only 100 km south of Bezwe, we are confident to use this available name for this widespread taxon. The species can be further diagnosed by having echolocation peak frequencies around 100 kHz (99–102 kHz, N = 8 from Malashane Cave; Table 2), which are quite distinct from typical swinnyi (105–107kHz, N = 6; Table 2) as well as the new Gorongosa National Park taxon R. gorongosae sp. nov. (104–108 kHz, N = 16; Table 2). Noseleaf structure is distinctive, being characterised by a hastate lancet, not as concave as true swinnyi, less erect, low, rounded connecting process and more pronounced posterior lobe. Bacular structure of rhodesiae is clearly distinct from other swinnyi -like animals (see Figs 7, 8), being characterized by a distinctly longer tapered baculum with a distinctly broader base and shallow notchalong the lower portion of the shaft that isvisible in the lateral profile. Traditional morphometrics (Table 2; Fig. 4) do not differentiate rhodesiae from swinnyi proper; however, rhodesiae is clearly distinguished with minimal overlap from the distinctly smaller gorongosae sp. nov. Although quite small, the R. rhodesiae holotype falls within the range of variation of specimens assigned to R. rhodesiae from Mozambique, Zimbabwe, Zambia, northern South Africa (Pafuri, Limpopo Province), and Zanzibar (Table 2; Fig. 4). It falls clearly outside (larger than) the range of variation of the smaller gorongosae sp. nov. taxon (Fig. 4). Geometric morphometric results result in a better separation between rhodesiae and swinnyi s.s. with only minimal overlap (Fig. 5). Once again, the rhodesiae holotype from Bezwe River clusters within the range of variation of the rhodesiae taxon and outside the gorongosae sp. nov. or swinnyi taxa, thus validating the use of this name for this taxon. Distribution and biology: Combined molecular and morphometric data suggest the widespread distribution of this taxon from central and northern South Africa through Zimbabwe, Zambia and Mozambique extending to Zanzibar (Fig. 11). Based on specimen assignments on morphological grounds, the species co-occurs with R. swinnyi in central KwaZulu-Natal at Ferncliff Cave, as well as occurring in close proximity in northern KwaZulu-Natal, recorded at Hlabisi Forest close to Ngome Forest where swinnyi was recorded (Fig. 11). The widespread extent of this taxon and its occurrence in northern South Africa is confirmed by the widespread occurrence of a hitherto unidentified 100 kHz acoustic type recorded in the Soutpansberg (Taylor et al., 2013), and Pafuri Region of northern Kruger National Park (Taylor & Parker, unpublished data)., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1265-1269, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Taylor PJ, Sowler S, Schoeman MC, Monadjem A. 2013. Diversity of bats in the Soutpansberg and Blouberg Mountains of northern South Africa: complimentarity of acoustic and non-acoustic survey methods. South African Journal of Wildlife Research 43: 12 - 26."]}

Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor, Richards, Leigh R. (2018): Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus). Zoological Journal of the Linnean Society 184: 1249-1276, DOI: https://doi.org/10.1093/zoolinnean/zly024

RHINOLOPHUS SWINNYI GOUGH, 1908 SWINNY’ S HORSESHOE BAT Synonyms: Rhinolophus swinnyi piriensis Hewitt, 1913, South Africa, Eastern Cape Province, Pirie, near King Williams Town. Types: TMSA 1021 (holotype, male), 1022 (cotype, male), collected on 22–23 February 1908 by H. H. Swinny. Type locality: South Africa, Eastern Cape Province, Pondoland, Ngqeleni District, - 31.667° S, 29,033° E. Referred specimens having molecular identifications: None. Referred specimens having only morphological identifications: All specimens originate from South Africa. DM 7084, adult male collected by P. J. Taylor on 3 March 2001 and DM 13250, 13252 and 13254 (all of unknown sex), collected by E. J. Richardson on 2 November 2010 from the Eastern Cape Province, Insizwe Mine, (- 30.804° S, 29.281° E); DM 14036 (male), collected by M. C. Schoeman on 9 May 2012 from the KwaZulu-Natal Province, Pietermaritzburg, Ferncliff Nature Reserve, (- 29.550° S, 30.320° E); DM 14291 (female) and 14292 (female) collected by L. R. Richards on 1 April 2014 from the KwaZulu-Natal Province, Eshowe, Entumeni Nature Reserve, (- 28.886° S, 31.376° E); DM 14441 (female) and 14440 (male) collected by S. Stoffberg and M. C. Schoeman on 23 July 2004 from Eastern Cape Province, Kokstad Mine, (- 30.810° S, 29.280° E); DM 15018, adult female collected by L. R. Richards on 7 April 2016 from Eastern Cape Province, Sandile’s Rest Trout and Forest Country Estate, (- 32.661° S, 27.298° E); TMSA 39848, adult male collected by G. Bronner on 14 March 1988 from KwaZulu-Natal Province, Ngome Forest Reserve, (- 27.833° S, 31.413° E). Re-diagnosis and comparisons: Gough’s (1908) original description emphasized inter alia, the very small size of the species, the ‘mouse = grey’ colour, the position of the anterior premolar within the toothrow, the medium-sized ears, the shape of the connecting process (‘forming a marked projection, rounded terminally’), the parallel-sided edges of the sella and the moderate lancet with strongly concave edges. Rhinolopus swinny s.s., as defined here, confined to the Eastern Cape, KwaZulu-Natal and possibly Mpumalanga Provinces, South Africa, is clearly distinct on molecular grounds from R. rhodesiae and R. gorongosae sp. nov., and has as its closest relative, R. capensis with which it overlaps in range in the Eastern Cape. The echolocation peak frequency of R. swinnyi s.s. calls recorded in the Eastern Cape [mean 106.6 ± 0.4 kHz, N = 10; Schoeman & Jacobs (2008; Table S1)] and KwaZulu-Natal (mean 105.6 ± 0.76 kHz; this study) Provinces, South Africa overlap considerably, but differ clearly from the 100 kHz calls of R. rhodesiae from central and northern South Africa and Mozambique (and probably more broadly). Although R. swinnyi s.s. overlaps in external and cranial characters with R. rhodesiae, it can be distinguished from the latter taxon based on both baculum and noseleaf characters, as well as on detailed cranial shape analysis (see above). It can be differentiated from R. gorongosae sp. nov. on its larger size (e.g. condylecanine skull length 14.8–15.2 mm in R. gorongosae sp. nov., 14.3–16.0 mm in R. swinnyi; zygomatic width 8.13–8.56 mm in R. gorongosae sp. nov., 8.7–9.2 mm in R. swinnyi; Table 2), as well as on noseleaf, bacular and subtle cranial shape characters (see above). Distribution and biology: We here restrict the distribution of R. swinnyi s.s. to South Africa, including the Eastern Cape and KwaZulu-Natal Provinces and possibly Mpumalanga Province (Fig. 11). However, a recent specimen collected from the montane western region of Swaziland has tentatively been assigned to R. rhodesiae based on morphology and acoustics (maximum frequency = 102 kHz), but a molecular analysis has yet to be conducted (A. Monadjem, unpublished data); this suggests that R. rhodesiae (and not R. swinnyi) occurs to the north in Mpumalanga Province. Specimens from Limpopo Province in the north of South Africa are referred to R. rhodesiae. As pointed out above, swinnyi and rhodesiae co-occur at at least one locality, Ferncliff Cave, in central KwaZulu-Natal., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on page 1273, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Schoeman MC, Jacobs DS. 2008. The Relative Influence of Competition and Prey Defenses on the Phenotypic Structure of Insectivorous Bat Ensembles in Southern Africa. PLoS ONE 3: e 3715. doi: 10.1371 / journal. pone. 0003715"]}

RHINOLOPHUS GORONGOSAE SP. NOV. LEAST HORSESHOE BAT Holotype: Durban Natural Science Museum (DM) No. 14820 (field number JAG196), is an adult male, preserved in 70% ethyl alcohol, originally part of series of specimens collected by J. A. Guyton on 25 April 2015. The cranium and baculum has been extracted and examined for this study. The specimen has been included in both morphometric analyses. Type locality: Bunga Inselberg, Gorongosa National Park, Sofala Province, Mozambique − 18.599° S, 34.343° E, 212 m. Paratypes: Eight specimens collected 24–25 April 2015 (DM 14815–14819), 2 May 2015 (DM 14828), 5 November 2015 (DM 14843) and 22 July 2015 (DM 14865). Referred specimens having molecular identification: DM 14815 (JAG 188) a female specimen collected by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, Bunga Inselberg, - 18.599° S, 34.343° E, 212 m; DM 14843 (JAG 228) a female specimen collected by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, - 18.694° S, 34.208° E, 308 m. Referred specimens having only morphological identification: TMSA 49116 (JAG 31), adult male, collected on 21 April 2013 by J. A. Guyton from Mozambique, Sofala Province, Gorongosa National Park, Cheringoma Plateau, Gorge Rim, Site 1, - 18.635° S, 34.808° E, 213 m. Incertae sedis: DM 14864 adult female collected by J. A. Guyton on 21 July 2015 from Mozambique, Sofala Province, Gorongosa National Park, - 18.465° S, 34.052° E, 1150 m; DM 11482, adult female collected on 1 May 2009 by J. Bayliss from Mozambique, Nampula Province, Mount Inago Forest Camp, - 15.045° S, 37.396° E 1478 m. Etymology: The species derives its name from the Gorongosa district of Mozambique, in particular Gorongosa National Park, a biologically diverse region of southern Africa. Diagnosis: The species can be clearly distinguished from both R. swinnyi s.s. and R. rhodesiae on molecular grounds (Figs 2, 3) as well as by its smaller size (Fig. 4; Table 2), distinct cranial shape (Figs 5, 6), echolocation call peakfrequency (Table 2), baculum (Figs 7, 8) and noseleaf (Figs 9, 12) characteristics. Although some measurements overlap, there is minimal overlap in condylocanine skull length and zygomatic skull width between this species (14.8–15.2 mm; 8.13–8.56 mm) and R. swinnyi (14.3–16.0 mm; 8.7–9.2 mm) and R. rhodesiae (15.1–15.7; 8.46–8.96 mm). The small size of this form makes it even smaller than denti (regarded by Csorba et al., 2003 as the smallest species in the Ethiopian region), therefore making this new species Africa’s smallest horseshoe bat. Comparing means for R. gorongosae sp. nov. (Table 2) and denti (Monadjem et al., 2010): forearm 41.3 mm cf. 43.1 mm; mass 5.6 g cf. 7.0 g. Description: The genetically-distinct R. gorongosae sp. nov. is similar in pelage colour but distinctly smaller in external (mean total length 68 mm, mean forearm length = 41 mm) and cranial (mean condylocanine length 15.1 mm) measurements (Table 2) than both rhodesiae (mean totallength 75 mm, forearmlength 44 mm, condyolocanine length 15.4 mm) and swinnyi s.s. from South Africa (mean totallength 73 mm, forearm length 44 mm, condyolocanine length 15.4 mm). Based on geometric morphometric results, the diminutive R. gorongosae sp. nov. has a reduced foramen magnum, a noticeable depression along the parietal region, low set bullae, a narrow braincase and a narrow and high set nasal inflation with a sharp slope from the nasal inflation to the maxillae (Fig. 6). Lancet is subtriangular with straight to slightly concave sides, and a high, rounded connecting process; the sella has a diminutive, pointed tip (Fig. 12). Baculum short with reduced base, with a characteristic notch on one side of the shaft tip (position variable). The slightly narrower baculum shaft with narrower tip distinguishes R. gorongosae sp. nov. from R. swinnyi (Table 5; Figs 7, 8). Maxillary toothrow with minute and laterally displaced PM 1 with a distinct gap between the alveolar borders of canine and PM 2 (Fig. 10). Mean peak echolocation CF frequency is 106 kHz (104–108 kHz; Table 2). Distribution and biology: So far, it appears that this tiny species is restricted in its distribution to Gorongosa National Park in Mozambique (Fig. 11), although we provisionally refer a very small adult individual from Mount Inago in Mozambique to this taxon. Molecular sequences are required from a wider range of localities to determine the range of this taxon. Given that the two individuals sequenced from Gorongosa National Park were distinct from each other, the possibility exists that more than one cryptic species may be present., Published as part of Taylor, Peter J., Macdonald, Angus, Goodman, Steven M., Kearney, Teresa, Cotterill, Fenton P. D., Stoffberg, Sam, Monadjem, Ara, Schoeman, M. Corrie, Guyton, Jennifer, Naskrecki, Pitor & Richards, Leigh R., 2018, Integrative taxonomy resolves three new cryptic species of small southern African horseshoe bats (Rhinolophus), pp. 1249-1276 in Zoological Journal of the Linnean Society 184 on pages 1262-1265, DOI: 10.1093/zoolinnean/zly024, http://zenodo.org/record/4253044, {"references":["Csorba G, Ujhelyip P, Thomas N. 2003. Horseshoe bats of the aeorld (Chiroptera: Rhinolophidae). Shropshire: Alana books.","Monadjem A, Taylor PJ, Cotterill FPD, Schoeman MC. 2010. Bats of Southern and South-central Africa: a biogeographic and taxonomic synthesis. Johannesburg: Wits University Press."]}

Reardon, Shannon, Schoeman, M. Corrie (2017): Species richness, functional diversity and assemblage structure of insectivorous bats along an elevational gradient in tropical West Africa. Acta Chiropterologica 19 (2): 273-285, DOI: 10.3161/15081109ACC2017.19.2.005

Our extension of the phylogenetic study of Taylor et al. (2009) on a larger, more geographically representative sample confirmed their finding of genetically distinct sympatric lineages of bats currently referred to as Chaerephon pumilus sensu lato (s.l.) (family Molossidae) in southeastern Africa. Chaerephon pumilus s.l. comprised two cytochrome b lineages separated by a mean genetic distance of 0.7% (0.1–1.4%), consistent with intraspecific variability. The C. pumilus s.l. clade was paraphyletic, containing a nested C. leucogaster (Madagascar) clade. As well as the expected four mitochondrial control region lineages, we identified a new strongly-supported clade from the Durban area. Indices of diversity and neutrality, combined with a ragged multimodal mismatch distribution, are inconsistent with demographic expansion of a single C. pumilus s.l. population in southeastern Africa, and suggest that the control region lineages are stable units at demographic equilibrium. Dating analyses suggest that these lineages were established during the late Pleistocene, between 60 000 and 13 000 years ago. We found no evidence to support our hypothesis that the sympatric genetic lineages of C. pumilus s.l. are associated with distinct sonotypes, as discriminant function analysis based on four echolocation parameters could not discriminate between the four clades. We hypothesise that the different genetic lineages may be distinguished by differences in social communication and behaviour.Key words: Chaerephon pumilus, mitochondrial DNA, differentiation, cryptic species, echolocation.

Although urbanization is perhaps the most damaging, persistent, and rapidly expanding form of anthropogenic pressure on natural ecosystems, data on the patterns and processes of sympatric bat species in urban landscapes are relatively scant. We quantified the packing and dispersion of sympatric animalivorous bats based on flight and echolocation parameters at two urban rivers in Durban, South Africa. We used null models to test if the observed phenotypic patterns deviated significantly from the random patterns expected in the absence of competition or prey defences at ensemble and functional group scales. As we predicted, species packing increased in the species-rich ensemble comprising many morphologically similar species that grouped together, with a few morphologically distinct species dispersed further away. Furthermore, we found evidence that competition influenced flight patterns of the open-air bats at the species-rich river ensemble, and prey defences influenced echolocation patterns of coexisting bats at the ensemble and functional group scales. However, non-deterministic processes such as habitat structure and the physics of sound and flight probably mediated these deterministic processes to influence the phenotypic structure of sympatric bats in urban landscapes.Key words: bats, urban landscape, rivers, competition, prey defences, null models.

HIPPOSIDEROS CRYPTOVALORONA SP. NOV. (FIGS 2, 7)

SYN. HIPPOSIDEROS C. COMMERSONI HILL, 1963, IN PART

SYN. HIPPOSIDEROS COMMERSONI SIMMONS, 2003, IN PART, Published as part of Goodman, Steven M., Schoeman, M. Corrie, Rakotoarivelo, Andrinajoro & Willows-Munro, Sandi, 2016, How many species of Hipposideros have occurred on Madagascar since the Late Pleistocene?, pp. 428-449 in Zoological Journal of the Linnean Society 177 (2) on page 443, DOI: 10.1111/zoj.12368, http://zenodo.org/record/5364289

Goodman, Steven M., Rakotondramanana, Claude Fabienne, Ramasindrazana, Beza, Kearney, Teresa, Monadjem, Ara, Schoeman, M. Corrie, Taylor, Peter J., Naughton, Kate, Appleton, Belinda (2015): An integrative approach to characterize Malagasy bats of the subfamily Vespertilioninae Gray, 1821, with the description of a new species of Hypsugo. Zoological Journal of the Linnean Society 173 (4): 988-1018, DOI: 10.1111/zoj.12223, URL: https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12223

Corman, Victor Max, Ithete, Ndapewa Laudika, Richards, Leigh Rosanne, Schoeman, M. Corrie, Preiser, Wolfgang, Drosten, Christian, Drexler, Jan Felix (2014): Rooting the Phylogenetic Tree of Middle East Respiratory Syndrome Coronavirus by Characterization of a Conspecific Virus from an African Bat. Journal of Virology 88 (19): 11297-11303, DOI: 10.1128/JVI.01498-14