An integrative approach to characterize Malagasy bats of the subfamily Vespertilioninae Gray, 1821, with the description of a new species of Hypsugo Author Goodman, Steven M. Author Rakotondramanana, Claude Fabienne Author Ramasindrazana, Beza Author Kearney, Teresa Author Monadjem, Ara Author Schoeman, M. Corrie Author Taylor, Peter J. Author Naughton, Kate Author Appleton, Belinda text Zoological Journal of the Linnean Society 2015 2015-04-30 173 4 988 1018 https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/zoj.12223 journal article 10.1111/zoj.12223 0024-4082 5335115 PIPISTRELLUS HESPERIDUS (TEMMINCK, 1840) Molecular genetics K2P distance within this lineage for animals obtained in dry forest habitats of western Madagascar was 0.001 ( N = 14, Table 1 ); hence, these populations show little genetic variation at the nucleotide level. No haplotype was shared in common between sequenced samples of P . hesperidus from South Africa (TM 47666) and Swaziland (TM 47738), as compared to those from Madagascar ( N = 28, Table 9 ). These lineages exhibited a divergence of 1.89% (K2P), reflecting 22 fixed mutations in the trimmed sequence. Haplotype diversity was reasonably high in the Malagasy population (Hd 0.744), in spite of a lower nucleotide diversity (Pi 0.00181), but these indices were not calculated for the African population due to limited sample size. Morphometrics Measurements presented in Table 2 . Craniodental morphology Members of this genus are easily differentiated from known Malagasy members of the genus Neoromicia by the presence of five post-canine teeth as compared with four in the latter genus, which lacks a first upper premolar (P 3 ). Bates et al . (2006) noted that the upper canine (C 1 ) was not in contact with the second upper premolar (P 4 ). On the basis of specimens sequenced herein, this character holds, although in some cases these two teeth are almost in direct contact. Bioacoustics Measurements presented in Table 6 . Bacular morphology As described by Bates et al . (2006) for Malagasy specimens of P . hesperidus , the baculum is relatively short ( 1.7–1.8 mm ); here we add an additional four specimens that range in total length 1.60–2.10 mm ( Table 8 ). The distal tip and proximal base show a bifid indentation, and the shaft is distinctly recurved ( Fig. 8C ). This is the same configuration found in African P . hesperidus ( Fig. 8D ; Kearney et al ., 2002 ), which ranges in total length from 1.7 to 2.5 mm (T. Kearney, unpublished data, N = 23). Known geographical range In Figure 1 , localities are presented of sequenced specimens of P . hesperidus , which also include the sites mentioned by Bates et al . (2006) for verified records of this species on Madagascar . Taxonomic comments Different lines of evidence, including karyological distinctions (e.g. Volleth et al ., 2001 ), have been published indicating that P . hesperidus is most likely paraphyletic and best considered a species complex. However, because of a lack of resolution in character differences of holotypes representing different proposed names currently considered synonyms of P . hesperidus , Simmons (2005) recognised several subspecies, which include P . h . fuscatus Thomas, 1901 from much of the Afrotropics and P . h . subtilis Sundevall, 1846 from southern Africa and Madagascar . Greater genetic sampling is needed from southern Africa to resolve the differences found in the current study between this portion of the continent and Madagascar , but given the subtle genetic differences and seemingly continuous character states associated with bacular morphology, we maintain Malagasy populations as P . hesperidus . Milne-Edwards (1881) described a small species of vespertilionid from Madagascar under the name Vespertilio humbloti . Peterson et al . (1995) mentioned that the type series was apparently lost and the status of this taxon could not be assessed. In the MNHN, the syntype series associated with Milne- Edwards’ description of V . humbloti was located (MNHN 1986.1074 to 1986.1082); most of the nine specimens were immature. Through the courtesy of Dr. Cécile Callou of the MNHN, the skulls from two adult specimens were extracted, cleaned, and allocated the numbers MNHN 1986.1074 and 1986.1075. Both of these specimens have a prominent first upper premolar (P 3 ), typical of the genera Pipistrellus and Hypsugo . Further, 1986.1074 has a seemingly bicuspid second upper incisor (I 3 ) and 1986.1075 an unicuspid I 3 that, in both cases, the longest portion of the I 3 reaches the length of the first upper incisor (I 2 ) cingulum; a diastema between C and second upper premolar (P 4 ), a prominent first upper premolar (P 3 ) in lateral view (similar to the illustration in Bates et al ., 2006 , Fig. 5B ); and craniodental measurements falling within the range of P . hesperidus ( Tables 3 and 4 ). On the basis of these different features, we identify the Milne-Edwards series of V . humbloti as P . hesperidus ; this awaits confirmation based on bacular structure or molecular genetics. Hence, V . humbloti is considered a junior synonym of P . hesperidus . Figure 6. Plot of principal component scores associated with the five bioacoustic variables for genotyped Malagasy vesper specimens on A, PC1 vs. PC2, B, PC1 vs. PC3 and C) PC2 vs. PC3. Information on component loadings is presented in Table 7. Two unsexed specimens held in the BMNH (28.1.24.1, 28.1.24.2) were collected by Capt. K. Parcon in June 1922 on Europa Island ( 22.3683°S , 40.3633°E ) positioned halfway across the Mozambique Channel between Mozambique and Madagascar . Using the craniodental characters outlined in Bates et al . (2006) and Monadjem et al . (2010) , the two animals are referable to Pipistrellus , and one of the specimens in relatively good shape (BMNH 28.1.24.2), based on the form of the first upper premolar (P 3 ) and the second upper incisor (I 3 ), as well as cranial shape, is identified as P . hesperidus and not Neoromicia nana , as written on the specimen label. This identification needs to be verified based on molecular data or potentially bacular morphology. In the same accession of the Pipistrellus material is a specimen of Coleura seychellensis Peters, 1868 (BMNH 28.1.24.3) obtained by the same captain on 4 October 1924 near Amirante Island, Seychelles ( 6.000°S , 53.1667°E ), indicating that Capt. Parcon navigated boats in the western Indian Ocean. The context that the Pipistrellus specimens were obtained is ambiguous and they could have been stowaways on the vessel after docking in a coastal area and then transported to Europa Island; the direction of travel from Africa towards Madagascar or vice versa is unknown. However, if they represent a natural occurrence on Europa Island, this would provide evidence of movements for this species spanning at least half the distance between southern Africa and Madagascar .