Doryrhina Peters, 1871

Foley, Nicole M., Goodman, Steven M., Whelan, Conor V., Puechmaille, Sebastien J. & Teeling, Emma, 2017, Towards Navigating the Minotaur's Labyrinth: Cryptic Diversity and Taxonomic Revision within the Speciose Genus Hipposideros (Hipposideridae), Acta Chiropterologica 19 (1), pp. 1-18 : 12-13

Genus Doryrhina Peters, 1871 (p. 314)

New combination — Doryrhina cyclops ( Peters, 1871) .

Synonyms

Phyllorrhina cyclops Temminck, 1853 .

Phyllorrhina cyclops Temminck, 1853 = Doryrhina cyclops (Temminck, 1853) , see Peters (1871).

Rhinolophus micaceus de Winton, 1897 .

Hipposideros cyclops de Winton, 1899 .

Hipposideros langi J. A. Allen, 1917 .

Doryrhina Peters, 1871

Description of the Genus Doryrhina

Morphological characters

Peters (1871) in his naming of the subgenus Doryrhina presented different characters associated with the noseleaf structure, and the type species for the subgenus was Phyllorhina (Doryrhina) cyclops ; a tropical African taxon ( Fahr, 2013). Here we elevate the subgenus Doryrhina to the level of genus for cyclops and expand Peter’s diagnosis.

The single species we place herein in the genus Doryrhina , D. cyclops , has a frontal sac on the forehead ( Hill, 1963; Fahr, 2013), in a central position behind the posterior noseleaf, and opening as a relatively small vertical slit ( Fig. 4 ). Within the slit are stiff white hairs that when everted form a distinct hair tuft ( Rosevear, 1965). The noseleaf of Doryrhina shows several particular aspects which differ from other hipposiderids, including Macronycteris . Members of the genus Doryrhina have two lateral fleshy leaflets ( Fig. 4 ), a common configuration in Hipposideros , although several members of this genus, as well as Macronycteris , have four leaflets ( Andersen, 1905, 1906; Payne et al., 1985; Strahan, 1995; Flannery, 1995 a, 1995 b; Bates and Harrison, 1997; Francis, 2008; Thong et al., 2012 a, 2012 b; Happold, 2013 c). The 2nd lateral leaflet in D. cyclops extends posteriorly and forms a continuous extension of the posterior leaf ( Fig. 4 ). Further, this species has a club-shaped process projecting from the posterior portion of the noseleaf, which distinguishes it from all African members of the genus Hipposideros , as well as Macronycteris . Other aspects of the noseleaf of D. cyclops include: 1) anterior portion is broad and with a club-like structure commencing at anterior margin of lateral leaflet and extending posteriorly to middle portion of noseleaf, internarial septa form relatively large and sculpted structures that partially obscure the deep-set nasal passages, and distinct lappets surrounded the nasal passages; 2) middle portion of noseleaf is absent; 3) posterior portion with a vertical medial septum (that extends as the club-shaped process mentioned above) and two prominent vertical lateral septa on either side of the vertical medial septum, which divide the structure into 6 separate cells, the lateral-most cell merging with the 2nd lateral leaflet, and posterior margin is a thin structure with little expansion. The different aspects of the noseleaf described herein are unique to Doryrhina and not found in another genus of the family Hipposideridae ( Rosevear, 1965; Payne et al., 1985; Flannery, 1995 a, 1995 b; Bates and Harrison, 1997; Francis, 2008; Monadjem et al., 2010; Happold, 2013 c). The exception is the African species camerunensis , apparently closely related to cyclops ( Hill, 1963) . Further, Tate (1941), Hill (1963) and Koopman (1994) based on anatomical characters considered the H. cyclops group to be distinct from other groups in this genus and composed of the African species H. cyclops and H. camerunensis , and the Australian-New Guinea species H. muscinus , H. wollastoni , H. corynophyllus , H. semoni and H. stenotis . With the exception of cyclops , these other species are not represented in our molecular dataset and are in need of study to determine if they are best placed in the genus Doryrhina or should be retained in Hipposideros .

The separated ears of D. cyclops are long, narrow and terminating with pointed tips. The average ear length in this species is 33.5 mm (sexes combined, n = 125) ( Fahr, 2013). Antitragus not present. Other measurements of D. cyclops , which show sexual dimorphism, include: mean forearm length, 65.3 mm (♂♂, n = 53) and 68.3 mm (♀♀, n = 45); tail length, 26.5 mm (♂♂, n = 42) and 29.2 mm (♀♀, n = 77); and hindfoot length (with claws), 19.8 mm (♂♂, n = 41) and 20.5 mm (♀♀, n = 73) ( Decher and Fahr, 2005). Doryrhina cyclops has dense, long and woolly pelage, on the dorsum generally blackish-brown and often with a frosted tint, while the ventrum is lighter and not frosted. No rufus or orange colour morphs are known. The wing and interfemoral membranes are blackish-brown and skin on forearm, wing digits and tibia are paler reddishbrown.

The skull of D. cyclops , is proportionately large, with a lengthened braincase, and low sagittal crest. The internarial septum is not enlarged. Rostrum is distinctly broad. Zygomatic arches are slender and form the widest portion of the skull. Premaxillae posteriorly wide and in broad contact with the palate. Anterior palatal foramina enclosed. Cochleae greatly expanded.

Dental formula — premolars 1/2, canines 1/1, premolars 2/2 and molars 3/3. The upper incisors and associated cusps are not well developed. The upper canines lack well-defined cusps, but have distinct cingula. The upper anterior premolar is reduced, visible in lateral view, and in contact with the canine and posterior premolar. Anterior lower premolar is distinctly small.

Karyological characters

On the basis of karyological information from the literature, further evidence can be found to support the resurrection of Doryrhina for a species previously placed in the genus Hipposideros , H. cyclops . Hipposideros shows extreme karyological conservatism, with nearly all species examined to date possessing a 2 n complement of 32 ( Harada et al., 1982; Hood et al., 1988; Rautenbach et al., 1993; Bogdanowicz and Owen, 1998; Koubínová et al., 2010). However, in the case of H. cyclops , 2 n = 36 ( Koubínová et al., 2010), which is also different from Macronycteris , 2 n = 52 ( Rautenbach et al., 1993; Koubínová et al., 2010).

Figures

FIG. 4. Morphological comparisons of the frontal sacs and noseleaves in A — Macronycteris commersonii (FMNH 213588, ♀), Madagascar, Province d’Antsiranana, Réserve Spéciale d’Ankarana [now Parc National], near Andrafiabe Cave, 31 October 2010; B — M. gigas (FMNH 128212, ♀), Senegal, Casmance, Diabane, 12 km SW of Adeane, 15 January 1983; C — M. vittatus (FMNH 192800, ♀, sequenced for Cyt-b), Tanzania, Pemba Island, Kaskazini Region, Micheweni District, Kilijini Village, 3 August 2006; and D — Doryrhina cyclops (FMNH 164973), Uganda, Masindi District, Budongo Forest, 25 June 1998. The form of M. cryptovalorona is similar to M. commersonii and no comparative specimen material was available for M. thomensis. Drawing by Velizar Simeonovski