The Definition of Harpiola (Vespertilionidae: Murininae) and the Description of a New Species from Taiwan
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Acta Chiropterologica, 8(1): 11–19, 2006 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS The definition of Harpiola (Vespertilionidae: Murininae) and the description of a new species from Taiwan HAO-CHI KUO1, YIN-PING FANG2, GÁBOR CSORBA3, and LING-LING LEE1, 4 1Graduate Institute of Ecology and Evolutionary Biology, National Taiwan University, 1, Sec 4. Roosevelt Road, Taipei, Taiwan R.O.C. 2Department of Biological Resources, National Chiayi University, 300 University Road, Chiayi, Taiwan R.O.C. 3Department of Zoology, Hungarian Natural History Museum, H-1083 Budapest, Ludovika tér 2, Hungary 4Corresponding author: E-mail: [email protected] A new species of Harpiola from Taiwan is described based on 11 specimens collected between 1998 and 2004. Careful examination of these specimens and those of the genus Murina, revealed the valid characters distinguishing Harpiola from Murina, including the enlarged upper incisors, the well developed first premolars in both toothrows with their bulk subequal to canines and the second premolars in the corresponding toothrow, and the strongly bifid lower canine. The new species from Taiwan can be distinguished from Harpiola grisea in India by having different shape of second upper premolar and different structure of first upper molar. Key words: Harpiola sp. nov., Taiwan, taxonomy INTRODUCTION The description of the genus Harpiola was based on a single specimen collected Within the subfamily Murininae two from Northwest India and both the wet genera are accepted generally, namely specimen and its extracted skull are in Murina and Harpiocephalus (Corbet and a very bad condition. As Corbet and Hill Hill, 1992; Koopman, 1993, 1994; Pav- (1992: 151) stated “the features upon which linov, 2003; Simmons, 2005). The name Thomas based Harpiola do not appear to be Harpiola was introduced by Thomas (1915) artificial, but to some extent the status of as a separate genus with Murina grisea grisea must remain uncertain until further Peters, 1872 as its type species. Although material is obtained.” Recently, alongside Tate (1941) accepted this opinion and pro- the description of the second known speci- vided a brief diagnosis of the genus (char- men of H. grisea from Mizoram, India, acterized by the attachment point of the Bhattacharyya (2002) re-evaluated Har- wing and by dental features), Ellerman piola and raised once again the taxon to and Morrison-Scott (1951) listed this form generic rank. as a subgenus of Murina, a taxonomic During a series of bat surveys carried arrangement which was followed subse- out between 1998 and 2004 in mountain quently. areas in Taiwan, 11 Harpiola bats were 12 H.-C. Kuo, Y.-P. Fang, G. Csorba, and L.-L. Lee obtained that contributed to the richest col- — taken across the outer borders of upper canines; lection of specimens of this genus in the upper molar width — taken across the outer crowns of world. In this paper, detailed comparisons the last upper molars; zygomatic width — the great- est width of the skull across the zygomatic arches; of these specimens with those of H. grisea mastoid width — the greatest distance accross the and Murina spp. were made to reveal their mastoid region; postorbital width — the least width of taxonomic status and to re-define the diag- the postorbital constriction; maxillary toothrow nostic features of the genus Harpiola. length — from the front of upper canine to the back Meanwhile, the special zoogeography of of the crown of the third molar; upper canine–premo- Harpiola as well as other mammalian gen- lar length — the largest distance from the front of the upper canine to the back of the crown of the posterior era was also considered. premolar; length of mandible — from the anterior rim of the alveolus of the first lower incisor to the most MATERIALS AND METHODS posterior part of the condyle; mandibular toothrow length — from the front of the lower canine to the The following comparative material was used: back of the crown of the third lower molar; lower ca- Harpiola grisea: India, BM(NH) 79.11.21.117 (holo- nine–premolar length — the greatest distance from type); Murina aenea: Malaysia, BM(NH) 64.770 the front of the lower canine to the back of the crown (holotype); M. aurata: China, MNHN CG1870-590a of the posterior premolar; height of the coronoid (paratype); M. cyclotis cyclotis: India, BM(NH) process — taken perpendicularly from the extremity 9.4.4.4 (cotype); M. cyclotis peninsularis: Malaysia, of the coronoid process to the ramus mandibulae. BM(NH) 64.771 (holotype); M. florium: Indonesia, BM(NH) 63.12.26.14 (holotype); M. hilgendorfi: Ja- pan, HZM 1.2974; M. huttoni: India, BM(NH) EVALUATION OF THE HARPIOLA CHARACTERS 79.11.21.685 (holotype); M. leucogaster rubex: India, BM(NH) 16.3.25.111 (holotype); M. puta: Taiwan, One of the main diagnostic features at- ZMNTU 1998.7.3; M. rozendaali: Malaysia, tributed to Harpiola is the attachment point BM(NH) 83.360 (holotype); M. silvatica: Japan, of the plagiopatagium. According to Tate HNHM 2001.38.1; M. suilla: Java, HNHM 2000.13.2; M. tubinaris: Pakistan, HNHM 99.14.6; (1941), Corbet and Hill (1992), Koopman M. ussuriensis: Russian Federation, ZMMU 96368 (1994), and Bhattacharyya (2002), it is at- (paralectotype). tached to the base of the first toe in Harpio- The museum acronyms mentioned in the text are la whereas it is attached to the base of the as follows: BM(NH) — Natural History Museum, claw of the first digit (the distal phalanx) in London, formerly British Museum (Natural History); ESRI — Endemic Species Research Institute, Nan- Murina. However, when describing M. hil- tou; HNHM — Hungarian Natural History Museum, gendorfi, Peters (1880) noted and figured Budapest; HZM — Harrison Institute, Sevenoaks, the attachment of the wing as being close to formerly Harrison Zoological Museum; MNHN — the base of the first toe. This view was sup- Museum National d’Histoire Naturelle, Paris; NMNS ported subsequently by Wallin (1969) and — National Museum of Natural Science, Taichung; ZMNTU — Zoological Museum of National Taiwan Yoshiyuki (1989). The latter described the University, Taipei; THU — Tunghai University, Tai- Japanese population of M. hilgendorfi as chung; ZMMU — Zoological Museum of Moscow having a “plagiopatagium attached to basal State University, Moscow. portion of first phalanxes of the first toe” The forearm (FA) measurements were taken from (Yoshiyuki, 1989: 216). Ognev (1928) not- dry or alcohol preserved museum specimens with 0.1 mm accuracy. The following craniodental measure- ed a similar point of attachment for M. og- ments were taken to the nearest 0.01 mm by the au- nevi and M. sibirica (these latter taxa are thors with digital calipers under a stereomicroscope: considered to be synonyms of M. hilgen- total length of skull — from the anterior rim of alve- dorfi — see Simmons, 2005). However, this olus of the first upper incisor to the most projecting point of the occipital region; condylobasal length — character was overlooked by subsequent au- from the exoccipital condyle to the posterior rim of thors and curiously Tate (1941: 579, 580), alveolus of the first upper incisor; upper canine width although citing Ognev’s (1928) statements, A new species of Harpiola from Taiwan 13 erroneously listed the attachment point as in M. aurata. In conclusion, the degree of a differentiating character of Harpiola. reduction of canines in both upper and low- For dental characters, the main distin- er toothrows can not serve as diagnostic guishing feature of Harpiola cited by Tate characters distinguishing Harpiola from (1941), Corbet and Hill (1992), and Koop- Murina. man (1994) was the reduction of the meta- The characters of Harpiola (Figs. 1 and cones of the anterior and middle upper 2) which do support its generic distinction molars. However, these authors were only are: 1) the upper incisors (I2 and I3) are ap- able to investigate the holotype of H. grisea proximately two-third that of the C1 in (the single known specimen of the genus at height; the basal area of second upper inci- that time) the teeth of which are extremely sor (I3) is more than two-third that of C1 (in worn. Bhattacharyya (2002) in his descrip- Murina, the height and crown area of I2 and tion of a second specimen of this species did I3 are at most half that of the C1); 2) the not mention the cusp pattern of the molars. basal area of C1, P2 and P4 are subequal, Nevertheless, in Harpiola from Taiwan (de- their height are gradually decreasing; the scribed herein as new species), the meta- corresponding teeth are similar in bulk and cones of the first (M1) and second (M2) mo- height in the lower toothrow (in Murina the lars are well developed and clearly higher canine always greatly exceeds the first pre- than the paracones. Furthermore, Thomas molar in height in both toothrows; P2 almost (1915), Corbet and Hill (1992), and Bhat- always, clearly smaller than P4 in height); 3) tacharyya (2002) all mentioned the high- C1 is strongly bifid, and the additional cusp ly reduced canines of Harpiola, which are is well developed (in Murina only a small in contrast with the usual specialized ca- secondary cingular cusp is present). These nines seen in Murina. However, even high- diagnostic characters can also be seen in er degree of reduction in the bulk of upper the type of H. grisea; however, the cusp canine can be observed in M. aurata (Mae- structure of C1 can hardly be detected due to da, 1980; Corbet and Hill, 1992). Although the very bad condition of the specimen. quantitative comparison and statistical analysis were not applicable because very SYSTEMATIC DESCRIPTION few specimens of relevant species are avail- able in the museum collections, detailed Harpiola isodon sp.