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Home , Ambilobe, Chaerephon (bat), Chaerephon jobimena

A New Species of (Molossidae) from with Notes on Other Members of the Family

Authors: Goodman, Steven M., and Cardiff, Scott G. Source: Acta Chiropterologica, 6(2) : 227-248 Published By: Museum and Institute of Zoology, Polish Academy of Sciences URL: https://doi.org/10.3161/001.006.0204

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Downloaded From: https://bioone.org/journals/Acta-Chiropterologica on 13 Jul 2020 Terms of Use: https://bioone.org/terms-of-use Acta Chiropterologica, 6(2): 227–248, 2004 PL ISSN 1508-1109 © Museum and Institute of Zoology PAS

A new species of Chaerephon (Molossidae) from Madagascar with notes on other members of the family

STEVEN M. GOODMAN1 and SCOTT G. C ARDIFF2

1Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, Illinois 60605, USA, and WWF, BP 738, Antananarivo (101), Madagascar; E-mail: [email protected] 2Columbia University, Department of Ecology, Evolution, and Environmental Biology, 1200 Amsterdam Avenue, New York, NY 10027, and Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA

We describe a species of Chaerephon (Molossidae) new to science from western Madagascar. This differs from the other two Chaerephon occurring on the island and from comparably sized African and Asian Chaerephon based on measurements, pelage and wing coloration, and cranial and dental characters. Chaerephon sp. nov. occurs at three sites in the drier western portion of the island. We also provide some natural history and distributional information on other Malagasy members of this family.

Key words: Chaerephon, Molossidae, new species, distributional records, western Madagascar

INTRODUCTION coastal and inland areas and it is also re- ported from the eastern offshore island of The of Madagascar are poorly Isle Sainte Marie (Peterson et al., 1995). It known. As currently understood, seven taxa can be found roosting in monospecific col- represent the Molossidae fauna of the is- onies or sharing sites with other molossid land: jugularis, ful- bats such as M. jugularis and M. leuco- milans fulminans, Chaerephon leucogaster, stigma. Chaerephon pumilus occurs in areas C. pumilus, leucostigma, M. midas of the humid east and central highlands and miarensis, and madagascariensis the vast majority of records are associated (Peterson et al., 1995; and spell- with human habitations (Peterson et al., ing in accordance with Simmons, In press). 1995). Its range at least partially overlaps Given the difficulty of capturing members with M. jugularis (see below). of this family, few data are available about During the course of biological invento- their distribution and natural history, even ries carried out over the past decade at dif- for the common commensal species. ferent sites on the island (Fig. 1), particular- On Madagascar two species of Chae- ly the western portion, we have captured rephon are currently recognized: C. leu- a number of bats, including Molossidae. cogaster and C. pumilus. The former spe- Amongst these collections is a species of cies often occurs as a commensal, living in Chaerephon that is unknown to science. man-made shelters, along the drier western The purpose of this paper is to describe the

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43° 47° 51° 12°

Andrafiabe Cave Ankarana 3rd canyon Ambilobe Ankijabe

Andapa

Mahajanga

16° Katsepy Ampijoroa Namoroka

Antsalova

Grotte Antananarivo d’Anjohimbazimba

20° Kirindy (CFPF)

Belo-sur-Mer Betakilotra Andrambovato Morombe

Ankiloaka Parc National de I’Isalo Tsifota Parc National de Zombitse-Vohibasia Toliara Sarodrano Saint Augustin Beza Mahafaly 24° Efoetse Andohahela Manantenina (parcelle 2) Collection sites of Ankapoky new Chaerephon sp. Ambovombe Other localities Faux Cap Protected areas

FIG. 1. Map showing sites at which Chaerephon sp. nov. has been collected as well as other localities mentioned in text

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new taxon and present new distributional mastoid processes; maxillary toothrow: length from records and natural history information for anterior alveolar border of canine to posterior alveo- 3 Malagasy molossid bats. lar border of M ; palatal length: from posterior border of hard palate to anterior edge of premaxillary bone; postorbital width: dorsal width at most constricted MATERIALS AND METHODS part of skull; post-palatal length: from posterior edge of hard palate, not including post-palatine process, to To investigate the taxonomic identity of the new the midventral lip (basion) of the foramen magnum; species of Malagasy Molossidae we have consulted upper canine height: greatest length from point im- specimens housed in several natural history muse- mediately dorsal to cingulum to end of unbroken or ums. The acronyms of these institutions are as fol- unworn tooth; upper molariform row: length from lows: BMNH — The Natural History Museum, Lon- PM4 to M3 (alveolar); upper toothrow: length from I1 don (formerly British Museum of Natural History); to M3; and width M3: greatest lateral-medial width of FMNH — Field Museum of Natural History, Chi- tooth. Further, one qualitative character associated cago; MNHN — Muséum National d’Histoire with the palatal emargination, as presented in Free- Naturelle, Paris; UADBA — Université d’Antana- man (1981: 15), was also recorded. narivo, Département de Biologie Animale, Anta- In order to help distinguish the new taxon of nanarivo; USNM — United States National Museum Chaerephon from other species in this genus of simi- of Natural History, Washington, D.C.; and UWZM — lar size, we conducted a discriminant function analy- University of Wisconsin Zoological Museum, Mad- sis followed by a canonical variates analysis in ison, Wisconsin. the statistical package SAS (PROC DISCRIM op- We recorded six external measurements in mil- tion CANONICAL; SAS, 1990). We used a non- limeters from collected specimens before preparation. parametric 3 nearest neighbor technique (option These included: total length, tail length, hind foot METHOD=NPAR K=3; SAS, 1990) for the discrimi- length (not including claw), ear length, and forearm nant function analysis because of the possible non- length. Further, we measured body mass in grams us- normal distribution of our data. We calculated Ma- ing a spring balance. For certain taxa, SMG obtained halanobis distances between mean discriminant func- these data directly from specimen labels or field cata- tion centroids and present F-statistics for these dis- logs in museum collections and not from we tances along with canonical axes loadings and eigen- measured. SMG added the additional variable of head values. We conducted separate analyses for skull, and body length from these museum sources. There body, and wing measurements and excluded some are notable differences amongst field collectors in the skull measurements because missing values would techniques that they use to measure bats; most impor- otherwise have greatly reduced sample size for the tant for molossids is the difficulty of consistent tail analyses. measurements and hind foot length (with or without Herein we follow the generic classification of the the claw). When tabulating descriptive statistics of Molossidae proposed by Freeman (1981) and subse- external measurements for certain species (Table 1), quently adopted by Simmons (In press), in which we have in some cases only used measurements of Chaerephon is given generic status rather than con- a single collector – although this reduces sample size, sidered a subgenus of Tadarida. As currently recog- it also decreases the range and variance of these meas- nized the genus Chaerephon is composed of 18 urements. SMG also took six wing measurements species (Simmons, In press), distributed across most from liquid preserved specimens: 3rd digit meta- of the Old World. Several members of this genus are carpal, 3rd digit 1st phalanx, 3rd digit 2nd phalanx, wide ranging. This includes C. pumilus, which occurs 4th digit metacarpal, 4th digit 1st phalanx, and 4th across much of Africa to the Middle East and on the digit 2nd phalanx. islands of Boiko, São Tomé, Pemba, Zanzibar, Al- SMG measured seven cranial and five dental dabra, Seychelles, and Madagascar. In contrast, other characters using digital calipers, accurate to the near- species, such as C. gallagheri, which is only known est 0.1 mm. The measurements and their definitions from the Democratic Republic of Congo, have limit- (derived largely from Freeman, 1981) are: greatest ed continental ranges. There are several remote island skull length: from posterior-most part of occipital to endemics (e.g., C. leucogaster from Madagascar, C. anterior-most point of the premaxillary bone; greatest solomonis from the Solomon Islands, and C. tomensis zygomatic breadth: width taken across zygomatic from São Tomé), which indicates that members of this arches at the widest point; lacrimal width: width genus are capable of dispersal over water and subse- across rostrum dorsally at lacrimal protuberances; quent colonization and speciation on distant oceanic mastoid breadth: greatest breadth across skull at islands.

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SYSTEMATIC DESCRIPTION limestone Ankarana Massif. The limestone in this area is largely eroded into pinnacle sp. nov. karst with solution flutes, stress fractures, (Figs. 2, 3, 4; Tables 1, 2) and bedding planes. The forest becomes wooded savanna several hundred meters Holotype — FMNH 169716, an adult Y, away from the capture site. collected on 23 January 2001 by Steven M. Diagnosis — A member of the genus Goodman, field number SMG 11932. The Chaerephon (see Freeman, 1981; Peterson specimen was prepared as a round study et al., 1995) given the following characters: skin, with associated skull and partial post- tooth behind the canine in line with the ca- cranial skeleton. The skin, skull, and post- nine; posterior upper molar with inverted cranial skeleton are in good condition. ‘N’ shaped cusp; lips wrinkled; ears joined Tissue samples were preserved in EDTA. by band of skin. Chaerephon jobimena is a Measurement – external (in mm): total relatively robust molossid with an average length 114, tail length 42, hind foot length forearm length of 46.5 mm, five upper (without claw) 7, ear length 23, forearm cheek teeth, a notably elevated mandibular length 46; body mass: 15.5 g; skull: (in mm) condyle, and a relatively constricted inci- greatest skull length 18.8, greatest zygomat- sive foramina that is not included in any ic breadth 10.9, postorbital width 3.9, mas- palatine emargination. It lacks the white toid breadth 10.3, lacrimal width 6.2, post- wing patches characteristic of several mem- palatal length 7.3, palatal length 7.8; dental bers of this genus. Wing membranes and (in mm): maxillary toothrow 6.6, upper uropatagium are dark and show no notable molariform row 5.1, upper toothrow 7.8, change in fur coloration along lower inner width M3 1.8, and upper canine height 2.9 edge of wing membrane. The dorsum and (Table 1). throat of C. jobimena is a uniform medium Etymology — The species epithet is de- chocolate brown. On the mid- to lower-ven- rived from northern dialects of Malagasy, trum hairs become more broadly tipped the region of the island the holotype was with light brownish-gray, giving an indis- collected, with joby meaning dark or black tinct grizzled gray appearance (Fig. 2). A ru- and mena red. When formed as a compound fous pelage phase also occurs. Interaural word in Malagasy the ‘y’ in joby becomes band forms complete connection between an ‘i’. The combination of these two words the ears. alludes to the two color morphs of this Paratypes — Madagascar: Province de species. Toliara, Forêt de Zombitse, 22°51’S, Type locality — Madagascar: Province 44°43’E, 870 m a.s.l., collected 18 April d’Antsiranana, Réserve Spéciale d’Ankara- 1993 by S. M. Goodman and I. H. Raso- na, 2.6 km E , in forest near An- lazaka — adult X, FMNH 151942 (SMG drafiabe Cave, 12°55.9’S, 49°03.4’E, about 5932); adult Y, FMNH 151985 (SMG 50 m a.s.l. (Fig. 1). 5936). Madagascar: Province de Fiana- Habitat — The type specimen was cap- rantsoa, Parc National de l’Isalo, 3.8 km tured in a harp trap placed on a small knoll NW Ranohira, along Namaza River, about 35 m from the principal west entrance 22°32.4’S, 45°22.8’E, 800 m a.s.l., collect- of Andrafiabe Cave. The habitat surround- ed 16–19 April 1999 by S. M. Goodman and ing the site is dry semi-deciduous forest D. Rakotondravony — adult X, FMNH growing on basaltic soils and alluvial sedi- 166075 (SMG 10990); adult Y, FMNH ments at the base of the western edge of the 166076 (SMG 10991); adult X, FMNH

Downloaded From: https://bioone.org/journals/Acta-Chiropterologica on 13 Jul 2020 Terms of Use: https://bioone.org/terms-of-use A new species of Chaerephon from Madagascar 231 and ) ) ) ) ) ) ) 9 3 12 18 13 13 18 – – C. jobimena 9.4 ± 1.53 11.9 ± 1.26 11.9 14.5 ± 1.09 14.6 ± 1.08 14.2 ± 1.26 19.6 ± 3.54 (5.6–11.0, (5.6–11.0, (12.5–16.0, (13.0–15.5, (9.5–14.0, (12.5–16.0, (15.0–28.0, ) ) (15.0–29.5, ) ) ) ) ) ) )– ) – 9 3 3 6 12 15 18 22 10 24 1 44.6 39.5 ± 0.71 (39–41, ) (45–48, ) (47–51, ) (45–51, ) (33–38, ) (42–46, ) (43–44, )– – )) (46–48, (45–47, ) ) (45–52, 5 9 3 3 7 11 12 15 20 22 24 1 20.2 -test statistical comparisons of measurements t ) (17–22, ) (18–22, ) (19–22, ) (16–21, ) (19–21, ) (21–24, ) (17–18, )) (22–24, (21–27, ) (12–19, ) (18–21, and other members of this genus. Descriptive statistics presented as 5 13 20 24 11 9 3 3 4 1 < 0.0001 12 22 P 11.2 (8–9, 8.3 ± 0.58 15.7 ± 3.51 ) (7–9, ) (10–12, ) (10–13, ) ) (6–8, ) (10–13, ) (5–7, ) (9–11, ) (9–12, )) (7–9, (7–9, 5 9 3 11 12 15 20 10 21 24 10 < 0.001, *** — 1 P Chaerephon jobimena 33.1 (32–51, (36–45, 39.9 ± 5.86 7.9 ± 0.93 23.1 ± 0.93 46.7 ± 0.87 39.3 ± 4.9339.8 ± 5.43 8.0 ± 1.00 7.9 ± 0.90 21.7 ± 0.58 22.8 ± 1.06 46.0 ± 1.0 46.5 ± 0.90 38.6 ± 2.16 ± 0.79 11.3 20.6 ± 1.05 48.7 ± 1.74 (32–51, (35–44, (30–40, (26–33, ) (29–36, ) (31–45, ) (30–44, ) (20–45, 3 11 24 10 1 – – – – – – – – – – 67.8 ) ) ) (64–68, ) (62–73, ) – (28–40, ) ) ) ) 5 9 3 11 12 20 24 10 21 ). Samples measured by a single field collector are in bold. 1 n – 72, 76 35, 38 17, 18 17, 20 40, 44 23.0, 26.2 –– 69.7 ± 3.30 (61–73, 39.1 ± 4.46 6.3 ± 0.96 18.9 ± 1.18 47.5 ± 2.37 17.0 –– 68.7 ± 1.42 (67–71, 39.1 ± 3.62 10.6 ± 0.82 20.1 ± 1.51 43.3 ± 0.58 – 110 – 40 12 17 47 – 100 Total and Head Tail Hindfoot Ear Forearm Body length body length length length length length mass 88.4 ± 3.70 – 29.9 ± 1.88 6.9 ± 0.61 17.2 ± 0.43 36.2 ± 1.26 95.3 ± 3.16 – 35.2 ± 2.66 (90–99, (80–95, 111.0 ± 3.46 111.0 112.3 ± 2.88 112.3 112.0 ± 2.92 112.0 113.0 ± 3.87 113.0 112.3 ± 6.12112.3 63.3 ± 2.08 39.3 ± 6.12 ± 6.12 11.6 19.8 ± 1.47 48.9 ± 1.13 21.0 ± 4.46 108.6 ± 3.85 – 35.6 ± 4.50 10.7 ± 1.36*** 20.0 ± 0.71*** 47, 50 – (107–117, (107–117, (109–115, (104–112, (104–112, (93–105, (106–122, (107–117, (107–117, (106–119, (106–119, YY XX XX (sexes combined for each species) were conducted: ** — + YY 1. External measurements (in mm) and body mass g) of Species ± SD (minimum–maximum, — Taken from Allen (1917) from — Taken ABLE 0 T 1 C. jobimena C. ansorgei C. bemmeleni C. bivittatus C. leucogaster C. jobensis C. johorensis C. ansorgei C. nigeriae C. russatus C. pumilus C. plicatus

Downloaded From: https://bioone.org/journals/Acta-Chiropterologica on 13 Jul 2020 Terms of Use: https://bioone.org/terms-of-use 232 S. M. Goodman and S. G. Cardiff ) ) ) ) ) ) 6 9 5 14 18 18) 10 11, 11, – 12, 13, 14, 14, 11, 11, 11, 11, – – – – – – – )(9 )(11 )(11 )(7 ) (10 ) (10 )(11 6 9 5 18 18 14 10 16, 17, 18, 12, 16, 18, 16, – – – – – – – ) (10 ) (15 ) (13 ) (14 ) (13 ) (15 ) (16 6 9 5 18 16 18 10 42, 45, 46, 34, 46, 45, 42, – – – – – – – ) (41 ) (29 ) (41 ) (39 ) (38 ) (44 ) (41 6 5 9 14 18 18 10 18, 21, 19, 21, 14, 21, 18, – – – – – – – ) (13 ) (18 ) (16 ) (18 ) (15 ) (16 ) (19 6 9 5 18 16 10 18 20, 20, 24, 15, 23, 20, 21, – – – – – – – ) (13 ) (17 ) (18 ) (18 ) (17 ) (18 ) (21 8 9 5 18 16 10 18 46, 46, 50, 42 18 17 40 15 10 41 19 16 40 16 9 34, 48, 45, 46, – – – – – – – 37, 38 15, 15 15, 15 34, 36 12, 13 9, 10 3rd digit 3rd digit 3rd digit digit 4th digit 4th digit 4th (40 (42 (45 metacarpal phalanx 1st 2nd phalanx metacarpal phalanx 1st 2nd phalanx 32.6 ± 1.33 13.7 ± 0.57 13.6 ± 0.51 31.3 ± 1.57 ± 0.55 11.2 9.4 ± 0.62 42.7 ± 2.66 18.2 ± 1.17 16.5 ± 1.38 40.3 ± 1.86 14.3 ± 1.03 ± 0.75 11.2 45.9 ± 1.77 19.8 ± 1.42 19.1 ± 0.87 43.7 ± 1.35 16.1 ± 0.86 12.4 ± 1.01 42.5 ± 1.35 19.6 ± 0.84 16.8 ± 0.63 40.3 ± 1.06 15.4 ± 0.70 9.2 ± 1.35 47.2 ± 1.92 22.4 ± 1.14 20.0 ± 1.00 44.8 ± 0.84 17.2 ± 0.84 12.0 ± 0.71 43.8 ± 1.44 19.3 ± 0.83 19.2 ± 0.99 42.1 ± 1.13 15.1 ± 0.83 12.2 ± 1.00 (30 (42 (41 (41 44.0 ± 1.58** 19.6 ± 0.73 18.0 ± 0.79** 42.2 ± 1.48** 15.8 ± 0.67 10.6 ± 0.53** YY XX 1. Extended Species ABLE C. leucogaster C. johorensis T C. nigeriae C. jobimena C. plicatus––––– russatus–––––– C. C. pumilus C. C. ansorgei C. bivittatus C. bemmeleni C. jobensis

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166077 (SMG 10998); adult Y, UADBA Province d’Antsiranana, Ankarana National 11501 (SMG 10999); adult Y, UADBA Park [sic], west side of park, Grotte 11502 (SMG 11000); adult Y, FMNH d’Andrafiabe, collected 7 June 1995 by J. 166100 (SMG 10968). Madagascar: Provi- M. Hutcheon — adult Y, UWZM 27555 nce de Fianarantsoa, Parc National de l’Isa- (JMH 107); adult Y, UWZM 27556 (JMH lo, 3.8 km NW Ranohira, along Namaza 108); UWZM 27557 (JMH 109). All of River, 22°32.4’S, 45°22.8’E, 800 m a.s.l., these specimens were preserved in fluid and collected 1 December 2002 by S. M. Good- the skulls removed and cleaned, with the man and H. V. Razakarivony — adult Y, exceptions of FMNH 151985 that the com- FMNH 175992 (SMG 13344); adult Y, plete cadaver was conserved in fluid and FMNH 175993 (SMG 13345). Madagascar: FMNH 151942 that was prepared as a round Province d’Antsiranana, Réserve Spéciale skin with associated skull and skeleton. d’Ankarana, Andrafiabe cave in 3rd canyon Description — In most adult individuals next to Andôkotôkana River (access to the of C. jobimena the dorsal pelage is a dense Fitsangatsanganan’Ilay Olona Tôkana pas- and velvety rich medium chocolate brown sage), 3.5 km SE Andrafiabe village, with lightly tipped hairs, particularly on the 12°56.5’S, 49°03.3’E, ca. 50 m a.s.l., col- mid-dorsum to rump, which gives an etched lected 15 May 2003 by SMG and SGC — or slightly iridescent appearance. Basally adult Y, FMNH 176329 (SMG 13706). the dorsal hairs are beige-white. In the holo- Madagascar: Province d’Antsiranana, An- type two small groups of white hair occur karana National Park [sic], west side of symmetrically on the dorsum posterior-lat- park, ‘Grotte de Cathedral’, collected 5 June erally to the ear insertions; these patches are 1995 by J. M. Hutcheon — adult X, not apparent in the other specimens refer- UWZM 27547 (JMH 99). Madagascar: able to this taxon. Ventrally, the throat is the

FIG. 2. Ventral view of the holotype of C. jobimena (FMNH 169716) obtained on 23 January 2001 near Andrafiabe Cave in the Réserve Spéciale d’Ankarana. (Photograph taken by Harald Schütz)

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FIG. 3. Drawings of the head of C. jobimena. Upper: lateral view and lower: dorsal view. (Drawing made by Rebecca Kramer)

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FIG. 4. Different views of cranium of C. jobimena (holotype FMNH 169716). Top — dorsal, middle — ventral, and bottom — lateral. (Photograph taken by John Weinstein, image number Z94380_04d)

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same color as the dorsum, but at the level of relatively well developed and the supraoc- the upper thoracic region there is a relative- cipital and frontal areas slightly inflated, ly abrupt change in pigmentation and the giving the braincase a bulbous but not over- hairs become more broadly tipped light ly rounded appearance. No sagittal crest is brownish-gray (Fig. 2). This transition is present and the interorbital region has a slightly diffused and does not form a dis- slight depression. Lacrimal protuberances tinctive throat collar. There is no notable and infraorbital foramina are relatively well change in fur coloration along the lower in- developed, and nasals are not inflated. ner edge of the wing membrane. In three of Paired basisphenoid pits are present and the 12 adult specimens of C. jobimena the moderately deep. Incisive foramina are in pelage coloration is distinctly rufous and the form of two small holes directly behind this species shows two chromatic phases. the canines and with a narrow septum sepa- Of 89 individuals captured for which color rating them and there is not a pronounced morph was recorded at Ankarana, 17 were emargination of the palate (condition 1 of rufous and 72 were dark; rufous males (0 = Taylor, 1999). 45.6 mm) had a shorter mean forearm than Distribution — This species is currently gray-colored males (0 = 46.0). This species known from the northern end of Mada- lacks the conspicuous postaural crest, inter- gascar in the Réserve Spéciale d’Ankarana aural pocket, or glands at the base of the tail and in the southern central area from the found in some other members of this genus Parc National d’Isalo and the Parc National (e.g., Allen, 1917; Hill, 1974; Kingdon, de Zombitse-Vohibasia (Fig. 1). 1974; Harrison, 1975). The wing membrane and uropatagium are dark brownish-black. COMPARISONS The muzzle is relatively blunt. The upper lips have approximately 5–6 wrinkles per Of the 18 species of Chaerephon recog- side and a regular, but not dense, covering nized by Simmons (In press) the following of hairs (Fig. 3). species (generalized distribution in paren- The ears are notably longer in C. jo- theses) fall within the approximate size bimena than in the two other species of range of C. jobimena (based on forearm Chaerephon occurring on Madagascar, but length) and were measured and examined falling within the general range of several for comparison with this taxon: C. ansorgei African and Asiatic species with compara- (broad range in Africa), C. bemmeleni ble forearm lengths (Table 1). The pinnae of (broad range in Africa), C. bivittatus (east- C. jobimena are relatively erect and partial- ern and southern Africa), C. jobensis (Mo- ly supported by a well-developed anterior luccas, Indonesia, New Guinea, and Aus- border (Fig. 3). There is a broad membrane tralia), C. johorensis (Malaya and Sumatra), connecting the ears, the center of which C. nigeriae (broad range in Africa), C. pli- forms a shallow ‘V’-shape. The antitragus is catus (broad range in Asia and the lobe-shaped and slightly asymmetric — Philippines, and has been recorded on measuring 5 mm (width) by 3 mm (height) Keeling Island), C. pumilus (broad range — and the tragus is a minute spike-like in Africa, and islands of Comoros, Mada- structure hidden behind the antitragus. gascar, and Seychelles), and C. russa- The skull of C. jobimena is relatively tus (broad range in Africa). Other species robust, with a short broad rostrum, expand- were not included in these analyses because ed braincase, and tapered postorbital con- they are smaller than C. jobimena (C. chapi- striction (Fig. 4). The lambdoidal crests are ni, C. gallagheri, C. major, C. shortridgei)

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or larger (C. aloysiisabaudiae) (Hayman C. russatus is a uniform russet-brown to and Hill, 1971; Kingdon, 1974; Koopman, cinnamon-brown. In C. jobensis the dorsal 1994; Taylor, 1999, 2000; Fenton and Eger, fur is notably short and mottled brown with 2002). A few taxa were not included in these a white grizzled appearance. Chaerephon comparisons because of their insular ranges bemmeleni has nearly transparent wing tips, that are distant from Madagascar (C. bre- bands of white fur at the base of the lower gullae, C. solomonis, C. tomensis). Even wings, and well-developed glands at the though smaller than C. jobimena, the Mal- base of the tail. The three species that show agasy endemic C. leucogaster is included pelage characteristics that approach those of herein for comparative purposes. C. jobimena are C. ansorgei from Africa, Other Malagasy members of the genus and C. johorensis and C. plicatus from Asia. Chaerephon can be distinguished easily The latter two taxa are apparently closely from C. jobimena. This latter species is no- related and are members of the same species tably larger in external and cranial charac- group (Simmons, In press). All four species ters than C. leucogaster (Tables 1 and 2), show dichromatism in pelage coloration has a relatively uniform dark pelage, and with rufous and dark brown morphs. The dark wings. Chaerephon pumilus is smaller throat hairs of C. ansorgei are generally than C. jobimena in numerous external and elongated and dark, forming a sort of cranial measurements (Tables 1 and 2) and ‘beard’, although in some cases this pelage has several pelage characters, such as the character appears to be rather variable band of white hairs on undersides of wings, (Rautenbach et al., 1984), and the wing that readily differentiate these species. In membranes are light in coloration and contrast to east African populations, C. pu- translucent. In dark individuals of C. plica- milus on Madagascar do not have distinc- tus the dorsum is generally a blackish- tive white wing membranes. In addition, brown and the ventrum notably paler — the nonparametric discriminant function analy- lower portion is edged with a gray wash. sis followed by canonical variates analysis The dorsum in dark morphs of C. jobimena indicate that C. pumilus differs from C. jo- is a medium chocolate brown and notably bimena based on skull, body, and wing lighter than in dark individuals of C. plica- measurement (Tables 3 and 4, Fig. 5). tus. On the basis of specimens that we have Chaerephon jobimena can be distin- examined, individuals of each morph type guished from most extralimital African and tend to be lighter in C. jobimena than in the Asiatic members of this genus with approx- comparable morph of C. johorensis and C. imately the same forearm length (Table 1) plicatus. by pelage characters (Hill, 1974; Kingdon, Hill (1974) described the well-devel- 1974; Bates and Harrison, 1997; Bonac- oped interaural band and pocket in C. jo- corso, 1998; Bouchard, 1998; Taylor, 1999, horensis, which is similarly developed in C. 2000; Willis et al., 2002). Chaerephon nige- plicatus. In this species complex the pocket riae and C. russatus have well-developed can be closed by an extension of the band postaural crests, which are not present in C. that joins inner-anterior margins of the ears. jobimena. Further, C. nigeriae has long The base of the pocket is largely without fur white hairs on the back and flanks, at least with the exception of a few long hairs on the in males, and whitish wing and interfemoral anterior portion. Chaerephon jobimena does membranes. Chaerephon bivittatus has dis- not possess this structure (Fig. 3). tinct white spots or short stripes on the head The skulls of C. jobensis, C. bivittatus, and shoulders. The pelage coloration in C. johorensis, and C. ansorgei are notably

Downloaded From: https://bioone.org/journals/Acta-Chiropterologica on 13 Jul 2020 Terms of Use: https://bioone.org/terms-of-use 238 S. M. Goodman and S. G. Cardiff ± SD ) ) ) ) ) ) ) ) ) ) ) 0 7 7 3 5 11 13 18 14 10 16 27 ) (7.7–9.0, ) (6.5–7.8, ) (6.9–8.0, ) (7.9–9.0, ) (5.3–6.8, ) (7.4–8.5, ) (8.2–9.2, ) (7.3–8.1, ) (7.3–8.0, ) (6.9–7.7, 7 7 7 3 11 18 13 10 15 24 < 0.0001 P -test statistical comparisons of measurements t ) (6.4–7.5, ) (7.1–8.1, ) (7.0–8.0, ) (5.8–6.6, ) (6.3–6.9, ) (6.8–7.1, ) (7.2–8.0, ) (6.8–7.9, ) (7.0–8.0, ) (7.0–7.3, ) – (7.1–7.9, 7 7 3 6 11 11 11 13 16 10 26 < 0.001, *** — P ) (6.1–6.3, ) (6.9–8.2, ) (5.7–6.4, ) (6.1–6.3, ) (6.1–6.3, ) (6.0–7.8, ) (6.3–7.3, ) (6.3–7.7, ) (6.0–7.1, ) (6.4–7.5, ) (5.1–5.9, 7 3 5 9 8 6 10 15 11 14 24 < 0.05, ** — P and other members of this genus. Descriptive statistics presented as ) (9.7–10.4, ) (9.6–10.3, ) (10.5–11.8, ) (10.2–10.7, ) (10.6–12.5, ) (10.7–11.2, ) (10.2–10.7, ) (10.3–10.4, ) (10.5–11.1, ) (10.1–12.3, 7 3 8 7 14 15 14 18 10 12 ––––– C. jobimena ) (3.5–3.9, ) (3.4–4.1, ) (3.8–4.3, ) 3.9, 4.2 10.3, 10.6 6.1, 6.1 7.1, 7.7 7.2, 7.3 ) (3.8–4.4, ) (3.8–4.1, ) (3.6–4.2, ) (4.3–4.7, ) (3.2–3.6, 27) (8.7–9.6, ) (3.8–4.1, ) (3.8–3.9, ) (3.9–4.2, 5 7 3 7 11 14 12 16 23 18 10 25 1 ) (10.3–10.9, ) (11.3–13.4, ) (10.5–11,4, ) (11.0–12.7, ) (10.7–11.6, ) (9.0–10.2, ) (10.1–11.4, ) (11.2–11.8, ) (10.4–13.3, ) (10.8–11.6, ) (10.7–11.0, ) (11.4–12.4, 6 5 7 3 8 11 14 15 23 18 10 23 (sexes combined for each species) were conducted: * — 18.5 11.0 Greatest Greatest Postorbital Mastoid Lacrimal Post-palatal Palatal 18.3, 18.8 11.3 11.1, 4.0, 4.1 10.4, 11.0 6.9, 6.9 6.5, 7.0 7.6, 8.0 17.4 ± 0.32 10.6 ± 0.21 3.7 ± 0.13 10.0 ± 0.20 6.1 ± 0.22 6.6 ± 0.18 7.0 ± 0.38 20.8 ± 0.68 12.1 ± 0.56 4.1 ± 0.14 ± 0.61 11.3 7.4 ± 0.38 8.0 ± 0.40 8.3 ± 0.32 19.1 ± 0.50 10.9 ± 0.36 3.7 ± 0.19 10.1 ± 0.26 6.8 ± 0.38 7.0 ± 0.32 7.9 ± 0.31 19.9 ± 0.57 12.1 ± 0.53 4.1 ± 0.20 ± 0.47 11.3 7.0 ± 0.51 7.6 ± 0.23 8.4 ± 0.33 18.6 ± 0.41 ± 0.26 11.0 3.9 ± 0.09 10.4 ± 0.15 6.2 ± 0.08 7.3 ± 0.29 7.6 ± 0.32 20.0 ± 1.17 ± 1.36 11.7 4.5 ± 0.17 ± 1.06 11.1 6.7 ± 0.82 6.7 7.5 ± 0.29 18.8 ± 0.33 ± 0.28 11.1 3.9 ± 0.10 10.5 ± 0.17 6.2 ± 0.08 7.4 ± 0.33 7.7 ± 0.23 18.2 ± 0.26 10.9 ± 0.15 3.9 ± 0.06 10.3 ± 0.06 6.2 ± 0.14 7.2 ± 0.15 7.3 ± 0.40 15.7 ± 0.39 9.6 ± 0.28 3.4 ± 0.10 9.1 ± 0.23 5.5 ± 0.23 6.2 ± 0.21 6.2 ± 0.30 19.3 ± 0.43 12.0 ± 0.30 3.9 ± 0.18 10.9 ± 0.18 6.8 ± 0.27 7.3 ± 0.34 8.5 ± 0.32 skull length zygomatic breadth width width width length length 19.3 ± 0.34** ± 0.27** 11.4 4.0 ± 0.10* 10.8 ± 0.20*** 6.6 ± 0.35** 7.5 ± 0.35 7.8 ± 0.29 (18.8–19.7, (18.8–21.4, (18.4–19.4, (18.0–18.2, (18.7–20.0, ). Standard deviation is not presented for palate emargination, which is a qualitative character. which is a qualitative character. deviation is not presented for palate emargination, ). Standard (18.1–19.6, (16.9–17.9, (19.3–22.0, (17.8–19.4, (18.8–20.6, (18.0–19.4, (14.9–16.3, n C. ansorgei and XX YY XX + YY 2. Cranial and dental measurements (in mm) of Species C. jobimena ABLE — Taken from Allen (1917) from — Taken C. pumilus C. jobensis C. plicatus C. russatus (minimum–maximum, C. bemmeleni C. bivittatus C. ansorgei T 1 C. johorensis of C. jobimena C. leucogaster C. nigeriae

Downloaded From: https://bioone.org/journals/Acta-Chiropterologica on 13 Jul 2020 Terms of Use: https://bioone.org/terms-of-use A new species of Chaerephon from Madagascar 239 ) ) ) ) ) ) ) ) 9 7 9 5 13 18 10 27 ) (1–2, ) (2–2, )– )) (0–2, (0–2, ) (0–1, ) (1–2, ) (1–2, )– )– ) (1–2, 6 3 3 7 11 15 18 10 14 14 27 upper canine emargination )) (2.7–3.7, (2.7–4.1, ) (2.3–2.9, )) (2.3–3.7, (2.4–3.0, ) (2.9–3.6, ) (1.9–2.8, ) (2.4–3.0, ) (2.7–3.3, ) (2.3–2.5, ) (2.5–2.9, 7 8 3 7 18 16 10 14 14 13 27 3 )) (2.0–2.3, (1.9–2.8, ) (1.8–2.0, )) (1.7–2.2, (1.5–2.2, ) (2.0–2.3, ) (1.9–2.2, ) (2.0–2.2, ) (1.8–2.0, ) (1.8–2.0, 5 7 3 7 18 15 10 14 14 12 )) (8.2–9.2, (8.1–9.4, )) (7.5–8.6, (7.1–7.7, ) (8.3–9.0, ) (7.4–8.3, ) – (1.3–1.9, ) (7.3–7.6, ) (8.0–8.6, ) (7.4–7.7, ) (7.8–8.3, 7 9 3 7 18 16 14 14 12 10 27 )) (5.0–5.7, (5.1–6.4, ) (4.6–5.4, ) (4.7–5.3, ) (4.7–5.3, ) (5.2–5.7, ) (4.1–4.9, ) (5.1–6.0, ) (5.2–5.9, ) (4.7–5.1, ) (4.8–5.3, 7 9 3 7 18 16 14 10 14 13 27 5.0, 5.0 6.7, 6.7 7.8, 8.1 2.0, 2.1 2.3, 2.5 2, 2 6.6, 6.7 4.8, 5.4 7.7, 7.8 1.9, 2.0 2.4, 2.7 1, 1 toothrow molariform row toothrow M Maxillary Upper Upper Width Height Palate 6.9 ± 0.40 5.4 ± 0.34 7.6 ± 0.22 2.1 ± 0.11 2.6 ± 0.22 1.2 7.6 ± 0.28 5.9 ± 0.45 8.8 ± 0.33 2.3 ± 0.20 3.2 ± 0.37 1.3 6.3 ± 0.19 5.0 ± 0.21 7.4 ± 0.22 1.8 ± 0.17 2.7 ± 0.22 0.9 7.4 ± 0.23 5.4 ± 0.21 8.7 ± 0.31 2.2 ± 0.10 3.0 ± 0.29 2.0 6.7 ± 0.23 5.0 ± 0.18 7.8 ± 0.25 1.9 ± 0.08 2.6 ± 0.21 – 6.7 ± 0.19 5.0 ± 0.13 8.0 ± 0.28 2.0 ± 0.15 2.9 ± 0.41 1.1 7.3 ± 0.66 5.4 ± 0.18 8.6 ± 0.22 2.2 ± 0.10 3.2 ± 0.23 1.9 5.6 ± 0.19 4.6 ± 0.23 – 1.7 ± 0.11 2.3 ± 0.20 0.9 6.5 ± 0.06 4.9 ± 0.21 7.5 ± 0.15 1.9 ± 0.10 2.4 ± 0.16 – 6.8 ± 0.20 5.0 ± 0.16 7.9 ± 0.16 1.9 ± 0.08 2.7 ± 0.17 – (6.5–7.6, (6.8–7.5, (6.4–6.5, (6.6–7.2, (7.0–7.7, (6.9–8.0, (6.4–7.1, (5.9–6.5, (6.4–7.2, (5.2–7.8, (5.3–6.0, 7.1 ± 0.21*** 5.5 ± 0.25*** 8.2 ± 0.23** 2.2 ± 0.07*** 2.9 ± 0.22* 1.7 XX XX YY + YY 2. Extended Species ABLE C. johorensis C. jobensis C. pumilus C. russatus C. bemmeleni C. bivittatus C. ansorgei C. plicatus C. nigeriae C. leucogaster C. jobimena T

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longer and more robust that C. jobimena (Tables 3 and 4, Fig. 5). Further, adult male (Table 2). Further, the first four species have and female C. jobimena are lighter in body prominent anterior and posterior sagittal mass than C. ansorgei — in the former crests, and in C. jobimena only the anterior species (Table 1) males weigh on average crest is slightly developed. The interorbital 14.6 g (range 12.5–16.0, n = 9) and females depression is absent in C. jobensis and C. 14.2 (range 13.0–15.5, n = 3), and in the lat- bivittatus, is slightly developed in C. ansor- ter species (Eger and Peterson, 1979) males gei and C. johorensis, and more prominent weigh on average 16.5 g (range 12.5–22.0, in C. jobimena. A lambdoidal crest, slightly n = 24) and females 15.8 (range 9.0–22.0, n inflated frontals, prominent lacrimal protu- = 46). Thus, based on these characters, C. berances, and moderately developed ba- jobimena and C. ansorgei are demonstra- sisphenoid pits, the latter of which are deep- tively different from one another. er in C. russatus, are found in all members of this genus. The supraoccipital region of NATURAL HISTORY C. jobimena is more inflated that in any oth- er species of this genus, with the exception Chaerephon jobimena is currently of C. russatus and C. bemmeleni. known from three sites on Madagascar, all On the basis of general size and pelage of which are in zones with various types of coloration, C. jobimena is similar in meas- dry deciduous forest. Further, two of the urements and pelage coloration (see above) three sites are in areas with large rock out- to the species C. ansorgei, which occurs in crops, filled with crevices and caves poten- the southern portion of Africa south of 12°N tially suitable as bat roosts. latitude (Eger and Peterson, 1979). A num- The northernmost site, near the extreme ber of morphological characters clearly dis- north of the country, is the Réserve Spéciale tinguish these two species, including exter- d’Ankarana, which encompasses a massif nal, cranial, and dental measurements of middle Jurassic limestone. This karstic (Tables 1 and 2). In addition, results of non- zone measuring about 25 km long and 8 km parametric discriminant function analysis wide is a myriad of caves, pinnacle karst, followed by canonical variates analysis in- and canyons (Cardiff and Befourouack, dicate that C. ansorgei differs from C. jobi- 2003). We have inventoried numerous caves mena based on body and wing measure- across the central portion of the massif and ments and possibly on skull measurements C. jobimena has only been found at three

TABLE 3. Morphological differences between C. jobimena, C. ansorgei, and C. pumilus based on nonparametric discriminant function analysis. Mahalanobis distances between discriminant function centroids differ according to the given F-statistics and probabilities (P). Differences in data availability for given specimens required the use of only select skull measurements and of separate discriminant analyses for skull, body, and wing measures. Skull characters used in the discriminant function analysis are listed in Table 4. Probabilities are approximate given small sample sizes

Analysis category F-statistics by species matrices C. ansorgei C. jobimena Skull (d.f. = 29) C. jobimena 2.12 (P < 0.09) – C. pumilus 4.73 (P < 0.01) 3.62 (P < 0.01) Body (d.f. = 16) C. jobimena 3.61 (P < 0.04) – C. pumilus 16.51 (P < 0.001) 37.25 (P < 0.001) Wing (d.f. = 21) C. jobimena 4.98 (P < 0.01) – C. pumilus 15.16 (P < 0.001) 16.26 (P < 0.001)

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sites (main Andrafiabe Cave entrance in The most central site at which we have western cliff, the Andrafiabe cave in 3rd recorded C. jobimena is in the Parc National canyon next to Andôkotôkana River, and de l’Isalo, a sandstone massif that has been the Andrafiabe cave in the 2nd canyon). heavily eroded and down-cut by water ac- These three localities are within a radius of tion. Specimens from this area were cap- 2 km of one another, are connected by the tured during two different field trips, both at Andrafiabe cave complex, are all close to the same exact locality in the Namaza River the western face of the massif, and are with- valley. The Namaza River starts as a dead- in several hundred meters of the ecotone be- end canyon deep within the massif, with tween dry semi-deciduous forest and an- towering cliff faces about 150 m tall. The thropogenic savanna. All individuals of this site from which the bats were obtained, is species captured at Ankarana were exiting situated just after the termination of the cliff caves after sunset and in the early hours face; all individuals were netted soon after of night. The known bat fauna of the Ré- dark, over the river, and in a direction exit- serve Spéciale d’Ankarana is composed of ing the canyon. Near the capture site there 14 documented species (Goodman et al., In is gallery forest growing along the river’s press). banks and about 250 m downstream the

TABLE 4. Loadings of canonical axes 1 and 2 (C1 and C2) from canonical variates analysis for skull, body, and wing characters of C. jobimena, C. ansorgei, and C. pumilus. Eigenvalues represent the variation among species centroids explained by each axis. The data used are the same used in the discriminant function analysis (Table 3). Missing data for some specimens required the use of only select skull measurements and of seperate analyses for skull, body, and wing measures. Loadings suggest that upper toothrow, maxillary toothrow, total length, forearm length, hindfoot length, third digit first phalanx, and fourth digit second phalanx are important in distinguishing the species Character category Measure C1 C2 Skull Greatest zygomatic breadth 0.840 -0.006 Postorbital width 0.858 -0.153 Lacrimal width 0.762 0.123 Maxillary toothrow 0.947 0.016 Upper molariform row 0.700 0.400 Upper toothrow 0.961 0.046 Width M3 0.769 0.460 Eigenvalues 1.797 0.626 % variation explained 74.2 25.8 Body Total length 0.821 -0.311 Tail length 0.274 -0.525 Hindfoot length -0.083 0.844 Ear length 0.877 -0.255 Forearm length 0.911 0.398 Eigenvalues 18.708 1.636 % variation explained 92.0 8.0 Wing Third digit metacarpal 0.749 0.471 Third digit first phalange 0.938 0.064 Third digit second phalange 0.574 0.667 Fourth digit metacarpal 0.792 0.537 Fourth digit first phalange 0.853 0.223 Fourth digit second phalange 0.051 0.763 Eigenvalues 7.188 2.12 % variation explained 77.2 22.8

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natural vegetation shifts to a narrow band of little topographic variation, and we are dry deciduous forest and then anthro- unaware of any rock outcrops in the vici- pogenic savanna. A total of 12 bat species nity of the capture site. The specimen of have been recorded within this park (Good- C. jobimena reported on from this site was man et al., In press). previously identified as C. pumilus (Raso- The third site from which we have ob- lozaka, 1994). The bat fauna of the Zom- tained material of C. jobimena is the transi- bitse-Vohibasia forest is composed of 10 tional dry deciduous-spiny bush forest of species (Goodman et al., In press). the Parc National de Zombitse-Vohibasia. We suspect that with further exploration Here the forest rests on alluvium soils with of western Madagascar, particularly in

A 3 2

1

0

C2 -1

-2

-3

-4 -4-4 -3 -2 -1 -1 0 1 2 2 3 B 6 5 4 3 2

C2 1 0 -1 -2 -3 -12-12 -10 -8 -6 -4 -4 -22 0 2 4 6 6 C 4 3

2

1

0 C2 -1

-2

-3

-4 -10-8-6-4-20-10-8-6-4-20 2 4 C1

FIG. 5. Canonical variates analysis axes scores for skull (A), body (B), and wing (C) measurements for specimens of Chaerephon ansorgei (Î), C. jobimena sp. nov. ( ), and C. pumilus (F). Missing values for some specimens required the separate analysis of skull, body, and wing measurements and the use of only select skull measures. Axes are scaled relative to their eigenvalues. Loading contribution of variables to each axis are shown in Table 4

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zones with water-cut rock outcrops, re- Ambovombe. In portions of the east speci- searchers will find C. jobimena at other mens of M. jugularis came from the village sites. All three of the currently known local- of Andapa, within 100 m of another colony ities for this species are within protected of C. pumilus (FMNH and UADBA); in areas. the train station of Andrambovato, along the eastern escarpment and in a zone RECORDS OF OTHER MOLOSSIDAE ON MADA- largely surrounded by forest (FMNH and GASCAR UADBA); and in a building at Manantenina (USNM). In November 2003 in the Parc Over the course of the past decade SMG National de Midongy du Sud a small colony and others have visited a number of sites to of less than 10 individuals of M. jugularis inventory bats and have captured numerous was found inhabiting a rock fissure along a molossids. We present here, by species, boulder-strewn river wash. This locality is records that add information to the distribu- within eastern humid forest, at least 4 km tion and natural history of this group, as from the forest edge, and bears evidence of summarized by Peterson et al. (1995) and the type of roosting site that this species nat- Eger and Mitchell (2003). We also present urally occupies in this habitat type. information on Malagasy specimens held in During a late January 2001 visit to natural history museums not associated with Cathedral Cave (an extension of the Andra- our recent inventories on the island. We fiabe Cave system) in Ankarana, we found present coordinates, elevation, and habitat M. jugularis at the end of its breeding sea- types of these different sites (Fig. 1, Appen- son. Near the entrance of the cave was a dix). Unless otherwise stated, all records are nursery colony with an estimated 1,000 represented by voucher specimens held in young nearly capable of flying but still with the Field Museum of Natural History and/or subadult dentitions, as milk teeth were pres- The Université d’Antananarivo, Départe- ent. Close to 100 dead and dying young ment de Biologie Animale. were found on the cave floor in a zone with Mormopterus jugularis. Peterson et al. sticky mud. Several of these individuals had (1995) reported this endemic and largely large and partially hardened mud balls at- commensal species from a range of sites tached to their feet. These balls presumably from eastern, central, southwestern, and formed after the bats dropped to the cave southern Madagascar, but not from western floor, crawled along the ground, and were Madagascar. We have captured it at several unable to reach a vertical surface before sites in areas of dry habitat, most in the they became too laden with mud. western portion of the island (listed north to Tadarida f. fulminans. The holotype south): Andrafiabe/Cathedral Cave (Anka- (BMNH 82.3.1.34) of the nominate form, rana), Ankijabe (), Ambilobe (with which is considered endemic to Madagas- Mops leucostigma), several sites in Isalo car, was collected in the vicinity of Fiana- (Namaza River, Canyon des Singes, along rantsoa by Rev. W. D. Cowan (Thomas, Menamaty River), Efoetse (Tsimanam- 1903) and there had been no subsequent petsotsa), and Andohahela (parcel II). The record of it on Madagascar (Peterson et al., vast majority of these individuals were 1995) until 1999. In mid-April of that year, found at roost sites in caves or buildings. SMG captured two individuals referable to There are also specimens (USNM) from this species along the Namaza River, Parc the southern portion of the island captured National de l’Isalo. One of these individu- at Faux Cap, Ankapoky, Tsihombe, and als, which was saved as a voucher specimen

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(FMNH 166074), was compared to the ) ) 6 holotype of T. f. fulminans in the BMNH, 6 and found to be similar to one another in general shape and cranial and dental meas- urements (Table 5). The Madagascar specimens were com- ) (13.4–14.1, ) (12.7–13.5, 6 pared to a small series of T. fulminans col- 5 lected in Kenya and Tanzania and Mo-

zambique, the former being notably larger 23.0 13.7 in external measurements than the east- ern African specimens (Table 5). Given the

general similarity in cranial measurements ) (21.9–23.2, 5 among these specimens, we assume the variation in external characters can be at- tributed to different techniques the field collectors used to measure the specimens. ) (23–31,

For the most part the cranial variables for 3 the two Malagasy specimens tend to be to- )

wards the upper limit or slightly larger than n the Kenya-Tanzania material. In contrast the specimens from Mozambique tend to be ) (54–60, ) – – (22.6–23.6, 7

larger than the Madagascar material. The 7 white ventral pelage coloration of the Kenyan material is often restricted to a rel- atively narrow portion of the mid-venter area and is similar to that in the Isalo spec- ) (19.5–24, ) (21–24, 7 imen. In contrast, specimens referable to T. 7 ± SD (minimum–maximum,

f. mastersoni from Mozambique have much 0 more extensive white on the venter, almost reaching the lower flanks in a few speci-

mens. Africa from Madagascar (including holotype for which cranial measurements are only available), eastern ) (10–14, ) (11–13, 7 The site at which we captured the Isalo 7 specimens of T. fulminans was over a river at the entrance of a deep sandstone canyon. The habitat type fits that of African popula- ) (46–58, ) (47–58, 6 tions of this species, which authors have 7 noted to occur in granite regions with high fulminans Tadarida ––––––

and steep-sided inselbergs, and to be dis- Total Tail Hindfoot Ear Forearm Body Greatest Greatest length length length length length mass length skull zygomatic breadth tinctly rupicolous (Cotterill, 2001). Chaerephon leucogaster. Peterson et al. ) (131–149, (1995) listed a few records of this species ) (131–144, XX from the western coastal area of the island, XX to which we can add the following sites and 4 and 2 (listed north to south): Ambilobe (with M. Sample 5. Measurements of adult YY YY jugularis), Katsepy (with M. leucostigma), from Isalo (BMNH 82.3.1.34) X (FMNH 166074)(3 152 61 11 21 61 27.5 23.4 13.4 (5 Antsalova (with M. leucostigma), Kirindy ABLE Holotype f. fulminans T. f. fulminans T. TanzaniaKenya and 138.8 ± 5.98 52.3 ± 4.31 12.0 ± 1.73 22.2 ± 1.63 57.3 ± 3.06 27.2 ± 2.95 22.5 ± 0.47 13.1 ± 0.27 Mozambique 136.7 ± 4.68 51.9 ± 3.34 12.0 ± 0.82 22.6 ± 0.98 – – 23.2 ± 0.36 13.8 ± 0.25 T (Tanzania and Kenya), and Mozambique. Statistics presented as and Kenya), Mozambique. Statistics (Tanzania

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(CFPF), Belo-sur-Mer, Betakilotra (Kirin- ) ) 6

6 dy-Mitea), Morombe, Ankiloaka (with M. leucostigma), and Ranobe (with M. leuco- stigma). The vast majority of these records are individuals captured at roost sites in buildings. There are few records of this spe- ) (9.1–9.8, ) (9.7–10.3, cies in the wild. 7 6 On 16 November 2002 a pregnant fe- male C. leucogaster was obtained near the village of Betakilotra. The was found roosting under the exfoliating bark of ) (5.9–6.2, ) (6.4–6.6, 7 6 a standing dead tree, about 5 m off the ground, and in a sandy valley with agricul- tural fields and stands of massive baobab trees. Three other members of this species flew off from the site. ) (7.8–8.5, ) (8.4–9.1, 6 6 Chaerephon pumilus. Peterson et al. (1995) noted records from central portion of the eastern escarpment. We captured a single individual of this species in the vil- lage of Andapa in the northern portion of ) (8.3–8.8, ) (8.9–9.4, 6 6 the eastern humid forest. In the same village was a roost of M. jugularis. This record is the westernmost we are aware of for C. pu- milus on the island. Mops leucostigma. The only record of ) (9.0–10.5, ) (9.8–10.4, 6 6 this species from the drier portions of Madagascar, presented by Peterson et al. (1995), was from Ampijoroa, inland from Mahajanga. We have collected it at the fol- ) (7.7–8.3, ) (8.1–9.0, 6 6 lowing sites (listed north to south): Ambi- lobe (near colonies of M. jugularis and C. leucogaster), Katsepy (with C. leucogas- ter), Antsalova (with C. leucogaster), Kirindy (CFPF), Ankiloaka (with C. leu- ) (12.6–13.5, ) (13.1–13.7, cogaster), Tsifota, Ranobe, and Saint Augu- 7 6 stin. All of these records are associated with human-built structures, except at Kirindy width breadth width length length toothrow row toothrow Postorbital Mastoid Lacrimal Post-palatal Palatal Maxillary Molariform Upper

(5.0–5.2, (CFPF). At this locality, M. leucostigma was obtained flying over a small waterhole along the Kirindy River in a large area of dry deciduous forest. Mops midas miarensis. Previously there

Sample were few records of this Malagasy endemic 5. Extended subspecies (Peterson et al., 1995). In recent (BMNH 82.3.1.34)from Isalo 4.9(FMNH 166074)Tanzania 4.6 13.0 (5.0–5.7, 12.9 8.5 8.4 9.6 10.1 9.3 9.8 8.7 8.8 6.4 6.4 10.0 10.1 ABLE years it has been netted in the Parc National Holotype f. fulminans T. f. fulminans T. Kenya and 5.2 ± 0.23 13.0 ± 0.33 7.9 ± 0.23 9.7 ± 0.53 8.6 ± 0.23 8.2 ± 0.23 6.0 ± 0.11 9.4 ± 0.28 Mozambique 5.1 ± 0.06 13.4 ± 0.26 8.7 ± 0.35 10.1 ± 0.22 9.3 ± 0.20 8.9 ± 0.26 6.5 ± 0.01 10.1 ± 0.22 T

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de Zombitse-Vohibasia (Rasolozaka, 1994), Environmental Exploration) through Hannah Thomas identified from Barn Owl (Tyto alba) pellets and Annette Olsson. Field research associated with collected near the Réserve Spéciale de Beza this paper has been generously supported by WWF- Madagascar, National Geographic Society (6637-99 Mahafaly (Goodman et al., 1993), found and 7402-03), and the Volkswagen Foundation. For roosting in the attic of a house in Ambo- valuable comments on a previous version of this pa- vombe (USNM), and netted near Ranobe per we are grateful to two anonymous reviewers. (FMNH) just outside of a building housing a colony of C. leucogaster and M. leucosti- LITERATURE CITED gma (H. Thomas, pers. comm.). The locali- ty reported for a specimen of this species in ALLEN, J. A. 1917. The American Museum Congo ex- pedition collection of bats. Part I. Systematic list. southwestern Madagascar at the site of Bulletin of the American Museum of Natural ‘Tamabuc’ (Peterson et al., 1995) is almost History, 37: 405–496. certainly a cacography for ‘Tamatave’ BATES, P. J. J., and D. L. HARRISON. 1997. Bats of the [=Toamasina] of eastern Madagascar [spec- Indian subcontinent. Harrison Zoological Muse- imen in question is BMNH 99.11.3.6]. um, Kent, England, xvi + 258 pp. BONACCORSO, F. J. 1998. Bats of Papua New Guinea. Otomops madagascariensis. This en- Conservation International, Washington, D.C., demic species was previously known from 489 pp. two sites on Madagascar: Namoroka, where BOUCHARD, S. 1998. Chaerephon pumilus. - the holotype was collected, and the Sarodra- ian Species, 574: 1–6. no area south of Toliara. We have captured CARDIFF, S. G., and J. BEFOUROUACK. 2003. The An- this species in the Réserve Spéciale d’Anka- karana Special Reserve. Pp. 1501–1507, in The natural history of Madagascar (S. M. GOODMAN rana (several different caves), Parc National and J. P. BENSTEAD, eds.) The University of de Bemaraha (Grotte d’Anjohimbazimba), Chicago Press, Chicago, xxi + 1709 pp. Parc National de l’Isalo (Namaza River), COTTERILL, F. P. D. 2001. Further notes on large Afro- and near Sarodrano (Grotte d’Ambanilia tropical free-tailed bats of the genus Tadarida and Grotte de Bisihiko). (Molossidae: Mammalia). Arnoldia Zimbabwe, 10 (18): 199–210. EGER, J. L., and L. MITCHELL. 2003. Chiroptera, bats. ACKNOWLEDGEMENTS Pp. 1287–1298, in The natural history of Mada- gascar (S. M. GOODMAN and J. P. BENSTEAD, eds.) We are grateful to the Direction des Eaux et The University of Chicago Press, Chicago, xxi + Forêts and Association Nationale pour la Gestion des 1709 pp. Aires Protégées for issuing permits to conduct faunal EGER, J. L., and R. L. PETERSON. 1979. Distribution surveys. For access to specimen material under their and systematic relationships of Tadarida bivittata care we are indebted to Michael Carleton, National and Tadarida ansorgei (Chiroptera: Molossidae). Museum of Natural History, Washington, D.C.; Canadian Journal of Zoology, 57: 1887–1895. Christiane Denys, Muséum National d’Histoire FENTON, M. B., and J. L. EGER. 2002. Chaerephon Naturelle, Paris; Paulina Jenkins, The Natural Histo- chapini. Mammalian Species, 696: 1–2. ry Museum, London; Judith L. Eger, Royal On- FREEMAN, P. W. 1981. A multivariate study of the tario Museum; Daniel Rakotondravony, Université family Molossidae (Mammalia: Chiroptera): d’Antananarivo, Antananarivo; and Paula Holahan, Morphology, ecology, evolution. Fieldiana: Zool- University of Wisconsin Zoological Museum, Mad- ogy (N.S.), 7: 1–173. ison. James M. Hutcheon kindly made specimens GOODMAN, S. M., O. LANGRAND, and C. J. RAX- available for study. Lucienne Wilmé thoughtfully pre- WORTHY. 1993. The food habits of the Barn Owl pared Fig. 1. We are grateful to Harald Schütz and Tyto alba at three sites on Madagascar. Ostrich, John Weinstein for photographs, Rebecca Kramer for 64: 160–71. the line drawing, and Tom Gnoske for some literature. GOODMAN, S. M., D. ANDRIAFIDISON, R. ANDRIA- Vola Razakarivony and Achille Raselimanana took NAIVOARIVELO, S. G. CARDIFF, E. IFTICENE, R. K. part in several field trips and we are grateful for their B. JENKINS, A. KOFOKY, T. MBONOAHY, D. RAKO- aid. Unpublished information and specimens were TONDRAVONY, J. RANIVO, F. RATRIMOMANARIVO, made available by Frontier-Madagascar (Society for J. RAZAFIMANAHAKA, and P. A. RACEY. In press.

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The distribution and conservation of bats in eds.). Recherches pour le Développement, Série the dry regions of Madagascar. Animal Conser- Sciences Biologiques, Spécial, Centre d’Infor- vation. mation et de Documentation Scientifique et HARRISON, D. L. 1975. A new species of African free- Technique, Antananarivo, 106 pp. tailed bat (Chiroptera: Molossidae) obtained by RAUTENBACH, I. L., D. A. SCHLITTER, and L. E. O. the Zaire River Expedition. Mammalia, 39: BRAACK. 1984. New distributional records of 313–318. bats for the Republic of South Africa, with spe- HAYMAN, R. W., and J. E. HILL. 1971. The cial reference to the Kruger National Park. of Africa. An identification manual. Part 2. Order Koedoe, 27: 131–135. Chiroptera. Smithsonian Institution Press, Wash- SAS. 1990. SAS User’s Guide, version 8. SAS Insti- ington, D.C., 1–73 pp. tute, Cary, North Carolina. HILL, J. E. 1974. New records of bats from southeast- SIMMONS, N. B. In press. Order Chiroptera. In Mam- ern Asia, with taxonomic notes. Bulletin of the mal species of the World: a taxonomic and geo- British Museum of Natural History (Zoology), graphic reference, 3rd edition (D. E. WILSON and 27: 127–138. D. M. REEDER, eds.). Smithsonian Institution KINGDON, J. 1974. East African mammals. An atlas of Press, Washington, D.C. evolution in Africa. Vol. II, Part A (insectivores TAYLOR, P. J. 1999. Problems with the identification and bats). Academic Press, London, xi + 341 pp. of southern African Chaerephon (Molossidae), KOOPMAN, K. F. 1994. Chiroptera: systematics. Pp. and the possibility of a cryptic species from 1–217, in Handbuch der Zoologie, Vol. VIII, South Africa and Swaziland. Acta Chiropterolo- Mammalia, Part 60 (J. NIETHAMMER, H. SCHLIE- gica, 1: 191–200. MANN, and D. STARCK, eds.). Walter de Gruyter, TAYLOR, P. J. 2000. Bats of southern Africa. Berlin, vii + 217 pp. University of Natal Press, Pietermaritzburg, vii + PETERSON, R. L., J. L. EGER, and L. MITCHELL. 1995. 206 pp. Chiroptères. Vol. 84. Faune de Madagascar. THOMAS, O. 1903. Three new species of Nyctinomus. Muséum National d’Histoire Naturelle, Paris, The Annals and Magazine of Natural History, 12 204 pp. (7): 501–505. RASOLOZAKA, I. H. 1994. Les micro-chiroptères. WILLIS, C. K. R., J. M. PSYLLAKIS, and D. J. H. SLEEP. Pp. 64–67, in Inventaire biologique, Forêt de 2002. Chaerephon nigeriae. Mammalian Species, Zombitse (S. M. GOODMAN and O. LANGRAND, 710: 1–3.

Received 27 October 2003, accepted 14 April 2004

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APPENDIX

Latitudinal and longitudinal coordinates and elevation of collection sites mentioned in text South East Site Elevation (m) latitude longitude Ambilobe 13°11.5’ 49°03.5’ 25 Ambovombe 25°10’ 46°05’ 135 Ampijoroa 16°18.868’ 46°48.620’ 100 Andohahela, Parcel II 24°49.0’ 46°36.6’ 120 Andrambovato 21°31’ 47°25’ 800 Andapa 14°39’ 49°38.5’ 500 Ankapoky 24°59’ 46°30’ 100 Ankijabe 13°15.6’ 49°38.1’ 125 Ankiloaka 22°46.548’ 43°36.889’ 90 Antsalova 18°39’ 44°37’ 150 Belo-sur-Mer 20°44.139’ 44°00.266’ 10 Betakilotra 20°53.2’ 44°04.8’ 35 Beza Mahafaly 23°40’ 44°37’ 150 Canyon des Singes (Isalo) 22°29.2’ 45°22.7’ 700 Cathedral Cave (Ankarana) 12°56.0’ 49°03.5’ ± 60 Efoetse 24°04’ 43°42’ 0 Faux Cap 25°35.5’ 45°32’ 0 Grotte d’Ambanila (Sarodrano) 23°32.397’ 43°44.763’ 0 Grotte d’Anjohimbazimba (Bemaraha) 18°41.7’ 44°43.0’ 110 Grotte de Bisihiko 23°32.933’ 43°46.044’ 10 Katsepy 15°45.808’ 46°14.695’ 5 Kirindy (CFPF) 20°04.578’ 44°40.492’ 30 Manantenina 24°18’ 47°20’ 30 Morombe 21°44.417’ 43°22.333’ 15 Menamaty River (near Isalo) 22°29.133’ 45°23.502’ 700 Namaza River (Isalo) 22°32.4’ 45°22.8 800 Namoroka 16°28’ 45°21’ 200 Ranobe 23°02.4’ 43°36.6’E 10 Saint Augustin 23°32.933’ 43°46.044’ 10 Tsifota 22°49.445’ 43°21.913 5 Zombitse-Vohibasia 22°50’ 44°42’ 800

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