Hipposideros commersoni (Sabatier & Legendre, 1985)

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?, Zoological Journal of the Linnean Society 177 (2), pp. 428-449 : 436-442

NEOTYPE OF HIPPOSIDEROS COMMERSONI ( FIG. 6 )

As mentioned earlier, a holotype was not designated by Geoffroy Saint-Hilaire (1813) in his description of R. (H.) commersoni , which was based on illustrations and different notes made by Philibert Commerson. Following the rules of The International Code of Zoological Nomenclature we designate a neotype here. The collections of the MNHN (Paris), where a potential type would have been deposited by Geoffroy Saint- Hilaire was searched to no avail for a specimen from Madagascar that pre-dates the description of this species. Hence, we have chosen a specimen that was measured and sequenced in the context of this study as the neotype of H. commersoni .

Neotype of Hipposideros commersoni

Adult female, FMNH 175972 , collected 9.xii.2002 by S. M. Goodman (field number SMG 13401). The specimen was conserved in 12% formaldehyde and subsequently transferred to 70% ethanol. Before preservation, the skull was removed via small incisions on both sides of the mouth, conserved in 60% ethanol, and then cleaned by dermestid beetles. The skull is in excellent condition. Samples of pectoral muscle C–M 3, complete cranial toothrow, length from anterior alveolar border of canine to posterior alveolar border of third molar; PM 4 –M 3, complete molariform toothrow, length from anterior alveolar border of second premolar ( PM 4 ) to posterior alveolar border of third molar; C 1 –C 1, width across upper canines, taken across the outer alveolar borders of the canines; M 3 –M 3, width across third molars, taken across the outer alveolar borders of the third molars; i 1 –m 3, mesiodistal length of complete mandibular toothrow, length from anterior alveolar border of incisors to posterior alveolar border of third molar; c–m 3, complete canine–molar mandibular toothrow, length from anterior alveolar border of canine to posterior alveolar border of third molar.

were collected and saved in lysis buffer. The specimen, with a full adult dentition and the basisphenoid–basioccipital suture completely fused, had large mammae, and an enlarged embryo (crown– rump length of 41 mm). External measurements: total length 133 mm, tail length 34 mm, hindfoot length (without claws) 16 mm, ear length 29 mm, forearm length 82 mm, and body mass 63 g (including 14.5 g embryo) ( Table 2). Information associated with the specimen includes ‘Netted over Sahanafa River in gallery forest’.

Type locality

Madagascar: Province de Fianarantsoa, Parc National de l’Isalo, along Sahanafa River, near foot of Bevato Mountain , 28 km south-east of Berenty-Betsileo , 22°19.0′S, 45°17.6′E, 550 m a.s.l.

Original diagnosis and type locality

Geoffroy Saint-Hilaire (1813) provided the following diagnosis of his Rhinolophus Commersonii : simple nasal leaf with a rounded terminal edge: without rostrum structure: the tail is half the length of the leg and occurs on Madagascar [‘Feuille nasale simple à bord terminal arrondi: sans bourse sur la front: la queue de la moitié moins longue de la jambe... Habite Madagascar’]. The neotype designated here possesses these same characters and was obtained on Madagascar .

Saint-Hilaire specifically mentioned that Commerson’s original description was based on an animal from near ‘ fort Dauphin’ [= Tolagnaro] ( Fig. 1 ). In the context of this current study, only a single adult female was measured from this locality ( USNM 578738 ), for which no fresh tissue was available. On the basis of the PCAs of craniodental and external characters, USNM 578738 is relatively large, yet it overlaps with other large-sized specimen referred to clades B and C ( Fig. 3A, B ). It also has a relatively large HF, similar to some smaller and medium-sized specimens from clade B ( Fig. 3B ). We prefer to designate a sequenced animal as the neotype of H. commersoni , in this case from Isalo, rather than the individual from Tolagnaro .

GSKL, greatest skull length, from posterior-most part of occipital to anterior-most point of upper canines; CCL, condylocanine length, from occipital condyles to anterior-most point of upper canines; ZYGO, greatest zygomatic breadth, width taken across zygomatic arches at the widest point; POB, postorbital breadth, dorsal width at most constricted part of skull; MAST, mastoid breadth, maximum width of skull across mastoid processes; MAND, mandible length, from the posteriormost portion of the condyles to anterior-medial-most point of upper incisors; MOMARM, moment arm of the temporal, length from mandibular condyle to tip of coronoid process; C–M 3, complete cranial toothrow, length from anterior alveolar border of canine to posterior alveolar border of third upper molar; PM 4 –M 3, complete molariform toothrow, length from anterior alveolar border of second premolar (PM 4) to posterior alveolar border of third molar; C 1 –C 1, width across upper canines, taken across the outer alveolar borders of the canines; M 3 –M 3, width across third upper molars, taken across the outer alveolar borders of the third molars; i 1 –m 3, mesiodistal length of complete mandibular toothrow, length from anterior alveolar border of incisors to posterior alveolar border of third molar; c–m 3, complete canine–molar mandibular toothrow, length from anterior alveolar border of canine to posterior alveolar border of third molar; TL, total length; TaL, tail length; HF, hind foot length (not including claw); FA, forearm length.

Note on spelling of species name

In the original description of this taxon, Geoffroy Saint-Hilaire (1813) proposed the name ‘ Rhinolophus Commersonii ’. Over the past nearly 200 years, the species name for this bat, with few exceptions, has been presented as commersoni . Following the International Code of Zoological Nomenclature, article 33.3.1, concerning ‘Incorrect subsequent spellings’, ‘when an incorrect subsequent spelling is in prevailing usage and is attributed to the publication of the original spelling, the subsequent spelling and attribution are to be preserved and the spelling is deemed to be a correct original spelling’. Hence, we maintain the spelling H. commersoni .

Morphological characters

Amongst H. commersoni , without exception, females of clade C are on average larger than those of clade B for external ( Table 2), cranial ( Table 3), and dental ( Table 4) measurements. As H. cryptovalorona sp. nov., named below, is morphologically similar to H. commersoni , characters are presented within the new species diagnosis and description, rather than being repeated here.

Molecular variation and phylogeny

The phylogenetic analyses confirmed the paraphyly of what was previously considered H. commersoni . Two well-supported, independently evolving lineages were recovered. The first lineage (clade A) is genetically distinct from the other H. commersoni lineage; the latter is further subdivided into two geographically correlated clades (clades B and C). Clade A is morphologically similar to H. commersoni individuals in clades B and C, but based on genetic data, this lineage can MD m 1, mesiodistal length of first lower molar (m 1), taken in line with the toothrow at the level of the buccal cingulum; BL m 1, buccolingual length of first lower molar, taken perpendicular to the toothrow at the widest point; MD m 3, mesiodistal length of third lower molar, taken in line with the toothrow at the widest point; BL m 3, buccolingual length of third lower molar (m 3), taken perpendicular to the toothrow at the widest point; TL c 1, total length of lower canine, taken along the vertical axis of the tooth from buccal cingulum to distal tip; TL m 1, total length of first lower molar, taken along the vertical axis of the tooth from buccal cingulum to distal tip; TL m 3, total length of third lower molar, taken along the vertical axis of the tooth from buccal cingulum to distal tip; MD M 2, mesiodistal length of second upper molar, taken in line with the toothrow at the level of the buccal cingulum; BL M 2, buccolingual length of second upper molar, taken perpendicular to the toothrow at the widest point; BL M 3, buccolingual length of third upper molar, taken perpendicular to the toothrow at the widest point.

MD M 2, mesiodistal length of second upper molar, taken in line with the toothrow at the level of the buccal cingulum; BL M 2, buccolingual length of second upper molar, taken perpendicular to the toothrow at the widest point; BL M 3, buccolingual length of third upper molar, taken perpendicular to the toothrow at the widest point. MD m 1, mesiodistal length of first lower molar (m 1), taken in line with the toothrow at the level of the buccal cingulum; BL m 1, buccolingual length of first lower molar, taken perpendicular to the toothrow at the widest point; MD m 3, mesiodistal length of third lower molar, taken in line with the toothrow at the widest point; BL m 3, buccolingual length of third lower molar (m 3), taken perpendicular to the toothrow at the widest point; DMB c, depth of mandibular base at level of canine, taken perpendicular along the buccal surface from the dorsal edge of the cingulum to edge of the base; DMB m 1, depth of mandibular base at level of first lower molar (m 1), taken perpendicular along the buccal surface from the anterior-most portion of the cingulum to edge of the base; DMB m 3, depth of mandibular base at level of third lower molar (m 3), taken perpendicular along the buccal surface from the anterior-most portion of the cingulum to edge of the base.

be recognized as a distinct species under the evolutionary species concept ( Simpson, 1961; Wiley, 1981; Templeton, 1989), phylogenetic species concept ( Eldredge & Cracraft, 1980; Cracraft, 1983), and the genetic species concept ( Bradley & Baker, 2001). The Cyt b sequence variation observed between clade A and H. commersoni (clades B and C) exceeds that recorded for other Afro- Malagasy bats. For example, Cyt b sequence divergence separating Chaerephon pusillus and Chaerephon leucogaster from congeners ranges from only 1.3–2.3% ( Goodman et al., 2010), and species belonging to the Rhinolophus hildebrandtii complex differ by 7.7–9.0% ( Taylor et al., 2012). The 9–11% average sequence divergence separating clade A from clades B and C is thus strong evidence for clade A representing a distinct species. The 6% sequence variation between clades B and C is also not trivial and highlights the need for further fine-scale phylogeographical study to clearly identify and understand the processes underpinning the evolution of these cryptic taxa.

Figures

Figure 1. Map of Madagascar showing different sampling localities of Hipposideros used in the morphological and molecular parts of this study and the different clade representation. The map also shows other localities mentioned in the text. An overlay is used of the simplified bioclimatic regions of the island (Cornet, 1974).

Figure 3. Principal components analysis plots of: A, 12 craniodental measurements and B, five external measurements for genotyped Malagasy Hipposideros female specimens. Clade B specimens from Sahanafa are shown as shaded squares with black outlines (clade Bs). Holotype of Hipposideros cryptovalorona sp. nov. (clade A, FMNH 175970) and neotype of Hipposideros commersoni (clade B, FMNH 175972), both obtained from Sahanafa, are labelled accordingly. Information on component loadings is presented in Table 5.

Figure 6. Different views of skull and mandible of neotype of Hipposideros commersoni (FMNH 175972, female) from Province de Fianarantsoa, Parc National de l’Isalo, along Sahanafa River. Dorsal view of cranium (upper row, left), ventral view of cranium (upper row, right), and lateral view of cranium and mandible (lower row). (Photographs taken by J. Weinstein, Field Museum image number Z95239_007d.)