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New distributional records of the recently described and endangered shrew tenrec Microgale nasoloi (Tenrecidae: Afrosoricida) from central western Madagascar
ARTICLE IN PRESS
www.elsevier.de/mambio
SHORT COMMUNICATION
New distributional records of the recently described and endangered shrew tenrec Microgale nasoloi (Tenrecidae: Afrosoricida) from central western Madagascar Voahangy Soarimalalaa,b, Steven M. Goodmanb,c, a
De´partement de Biologie Animale, Universite´ d’Antananarivo, BP 906, Antananarivo (101), Madagascar Vahatra, BP 3972, Antananarivo (101), Madagascar c Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA b
Received 27 March 2007; accepted 6 September 2007
Keywords: Microgale nasoloi; Tenrecidae; Western Madagascar; Distribution limit; Conservation status
Recent biological explorations of the dry forests of western and southern Madagascar have revealed that this zone has much higher small mammal species diversity than previously realized. Further, many of these discoveries comprise new mammals to science, including an assortment of bats (e.g., Goodman et al. 2005, 2006a, b, 2007), rodents (e.g., Carleton et al. 2001; Goodman and Soarimalala 2005; Carleton and Goodman 2007), and shrew tenrecs (e.g., Jenkins and Goodman 1999; Goodman and Soarimalala 2004). One of these recently described taxa from the dry forests of the island is the distinctive shrew tenrec Microgale nasoloi, which was described by Jenkins and Goodman (1999) based on two specimens from the southwest. Currently, this species is considered in the IUCN Red Data Book as endangered (www.iucnredlist.org). No other records of M. nasoloi were known until 2006, when two additional specimens were taken a notable distance from the sites of the type series and in a different habitat. Herein we report on these two new specimens collected by Voahangy Soarimalala (VS) and deposited in the Universite´ d’Antananarivo, De´partement de Biologie Animale (UADBA), Antananarivo, Madagascar, while the type series are held by the Field Museum of Natural History (FMNH), Chicago, USA (Table 1). Corresponding author.
E-mail address: sgoodman@fieldmuseum.org (S.M. Goodman).
The original description of M. nasoloi was based on two animals, the holotype coming from the Province de Toliara, Foreˆt de Vohibasia, 59 km NE de Sakaraha 22127.50 S, 44150.50 E, 780 m, in transitional dry deciduous forest, and the other specimen from the Province de Toliara, Foreˆt d’Analavelona, 12.5 km NW d’Andranoheza, 22140.70 S, 44111.50 E, 1050 m, on an isolated massif with elements of eastern humid and western deciduous forest (Jenkins and Goodman 1999; Fig. 1). Although these two sites are only about 70 km from one another, the local habitats are notably different. The central portion of western Madagascar, a region known as the Menabe, has been extensively surveyed for small mammals and the only species of shrew tenrecs known from the region are Microgale brevicaudata and Microgale longicaudata/majori (Ganzhorn et al. 1996; Raharivololona 1996; Olson et al. 2004; Hilgartner 2005). However, during an inventory of two sites in March 2006, two specimens were collected, which were tentatively identified as M. nasoloi and include: (1) Province de Toliara, Fivondronana de Morondava, Commune Rurale de Bemanonga, Kirindy (CFPF), Foreˆt d’Amboropotsy, 16.3 km NE de Marofandilia, 20103.20 S, 44141.00 E, 80 m, in dry deciduous forest (VS-1151, UADBA 48052), 19 March 2006. (2) Province de Toliara, Fivondronana de Belo sur Tsiribihina, Commune Rurale de Beroboka, Foreˆt
1616-5047/$ - see front matter r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.09.002 Mamm. biol. 73 (2008) 468–471
ARTICLE IN PRESS V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
Table 1.
469
Measurements of the four known specimens of Microgale nasoloi
Measurements
UADBA 48052
UADBA 48053
FMNH 156187 (holotype)
FMNH 161575
Sex Total length Head and body length Tail length Hind foot length Ear length Weight Condylo-incisive length Upper toothrow length Rostral breadth Interorbital breadth Braincase breadth Braincase height
~ 130 80 52 11 15 7.3 22.1 9.9 3.0 4.1 9.1 5.0
~ 127 78 50 12 15 8.2 22.4 10.1 3.0 4.2 9.0 5.0
~ – 81 53 13 16 14.0 23.2 10.2 3.1 4.3 9.2 4.9
# – 70 62 14 16 5.9 – – – – – –
All measurements in mm with the exception of weight, which is in gm. The measurements for the two FMNH specimens are taken from Jenkins and Goodman (1999).
Fig. 1. Map of western and southwestern Madagascar showing the four sites Microgale nasoloi has been found and other localities mentioned in the text. The inset map of Madagascar shows the zone delineated in the regional map to the left.
de Lambokely, 11 km NNE de Beroboka, 19152.20 S, 44138.70 E, 85 m, in dry deciduous forest (VS-1166, UADBA 48053), 29 March 2006.
These two specimens were subsequently compared to the different medium-sized Microgale spp. known from Madagascar (Jenkins 2003; Goodman and Soarimalala
ARTICLE IN PRESS 470
V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
2004) and based on various aspects of pelage, external measurements, and cranial and dental morphology are referable to M. nasoloi. All of the salient characters noted in the diagnosis of M. nasoloi by Jenkins and Goodman (1999) are discernable in the new material from the Menabe region. On the basis of tooth eruption patterns UADBA 48053 is at stage 3 (MacPhee 1987), showed no sign of current or past reproduction, and had notable subcutaneous fat deposits. In contrast, UADBA 48052 is younger in age and based on tooth ontogeny patterns fits MacPhee’s (1987) stage 1/2. This latter individual showed no sign of reproductive activity and was presumably nuliparous. At each of the two sites inventoried in 2006, three pitfall lines with plastic drift fences were installed in the forest (see Raxworthy and Nussbaum (1994) for a description of the technique). Each line comprised 11 buckets, 15 l in capacity, and placed at 10 m intervals. At each site, the pit-falls were installed for 6 consecutive nights, comprising an accumulated 198 pit-fall trap days. In the Foreˆt d’Amboropotsy, other than the single specimen of M. nasoloi, the other small mammals captured in these devices included Geogale aurita (n ¼ 1), M. brevicaudata (n ¼ 4) and Setifer setosus (n ¼ 1), while in the Foreˆt de Lambokely associated species included Suncus madagascariensis (n ¼ 1) and S. setosus (n ¼ 1). Hence, even though the surveys took place during the rainy season, when shrew tenrecs are presumed to be active, small mammal pit-fall capture rates were notably low, with seven animals (3.5% capture success) at Foreˆt d’Amboropotsy and three animals (1.5% capture success) in the Foreˆt de Lambokely. In February and March 2007, three different sites in the Kirindy-Mitea National Park were surveyed and 198 pit-fall trap days were accrued at each site and no individual of M. nasoloi was captured. Further, no sign of this species was found during extensive small mammal surveys in the zone north of the Tsiribihina River (V. Soarimalala and S. M. Goodman, unpublished data). The Foreˆt d’Amboropotsy is within the Kirindy (CFPF) forest and close to the research station, which has been an area of intensive biological research for several decades (Sorg et al. 2003), including inventory programs on small mammals (e.g., Ganzhorn et al. 1996; Raharivololona 1996). The discovery of M. nasoloi in this forest block, a species that is either rare or difficult to capture, indicates the importance of continued research with varied techniques and during different seasons to reach a more complete estimation of local small mammal diversity. Further, given the low pit-fall capture rates in the dry forests of Madagascar, it is clear that extensive survey work is needed to approach complete knowledge of the local small mammal fauna. The forest habitat where M. nasoloi was captured in the Menabe region is a seasonally dry deciduous habitat
and distinctly more mesic than the Zombitse-Vohibasia zone near Sakaraha. The Analavelona Massif site is rather particular, as it is notable moister than surrounding habitats associated with the summital zone receiving orographic precipitation. Based on these new records from the Menabe region, the habitat breadth and distribution of M. nasoloi is greatly expanded. The triangular-shaped polygon encompassing the four known localities for this taxon comprises an area of about 10,260 km2. However, whether it occurs across this zone is unclear and given its very recent discovery in the Kirindy (CFPF) area, much more detailed inventories will be needed to properly establish this point. M. nasoloi has been classified by the IUCN (www. iucnredlist.org) as endangered (EN B1ab(iii)), at least in part associated with its previously very limited geographic distribution and limited number of records. The new distributional data presented here clearly indicate that this conservation status needs to be re-evaluated. Further, given the remarkable level of new mammal species discoveries in the dry forests of Madagascar, the notable levels of deforestation of this region (Smith 1997), and high levels of habitat heterogeneity and species turnover, it is clear that this zone of the island needs to be given one of the highest priorities for new conservation actions.
Acknowledgements We are grateful to the Malagasy authorities for permits to conduct this research, which was supported by a grant from the John D. and Catherine T. MacArthur Foundation. Lucienne Wilme´ kindly prepared Figure 1.
References Carleton, M., Goodman, S.M, 2007. A new species of the Eliurus majori complex (Rodentia: Muroidea: Nesomyidae) from south-central Madagascar, with remarks on emergent species groupings in the genus Eliurus. Am. Mus. Novit. 3547, 1–21. Carleton, M., Goodman, S.M., Rakotondravony, D., 2001. A new species of tufted-tailed rat, genus Eliurus (Muridae: Nesomyinae, from western Madagascar, with notes on the distribution of E. myoxinus. Proc. Biol. Soc. Wash. 114, 972–987. Ganzhorn, J.U., Sommer, S., Abraham, J.-P., Ade, M., Raharivololona, B.M., Rakotovao, E.R., Rakotondrasoa, C., Randriamarosoa, R., 1996. Mammals of the Kirindy forest with special emphasis on Hypogeomys antimena and the effects of logging on the small mammal fauna. In: Ganzhorn, J.U., Sorg, J.-P. (Eds.), Ecology and Economy of a Tropical Dry Forest in Madagascar. Primate Report 46-1, pp. 215–232.
ARTICLE IN PRESS V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
Goodman, S.M., Soarimalala, V., 2004. A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Foreˆt des Mikea of southwestern Madagascar. Proc. Biol. Soc. Wash. 117, 251–265. Goodman, S.M., Soarimalala, V., 2005. A new species of Macrotarsomys (Rodentia: Muridae: Nesomyinae) from the Foreˆt des Mikea of southwestern Madagascar. Proc. Biol. Soc. Wash. 118, 450–464. Goodman, S.M., Jenkins, R.K.B., Ratrimomanarivo, F.H., 2005. A review of the genus Scotophilus (Chiroptera: Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27, 867–882. Goodman, S.M., Ratrimomanarivo, F.H., Randrianandrianina, F.H., 2006a. A new species of Scotophilus (Chiroptera: Vespertilionidae) from western Madagascar. Acta Chiropter 8, 21–37. Goodman, S.M., Cardiff, S.G., Ranivo, J., Russell, A.L., Yoder, A.D., 2006b. A new species of Emballonura (Emballonuridae: Chiroptera) from the dry regions of Madagascar. Am. Mus. Novit. 3538, 1–24. Goodman, S.M., Rakotondraparany, F., Kofoky, A., 2007. The description of a new species of Myzopoda (Myzopodidae: Chiroptera) from western Madagascar. Mamm. Biol. 72, 65–81. Hilgartner, R.D., 2005. Some ecological and behavioural notes on the shrew tenrec Microgale cf. longicaudata in the dry deciduous forest of western Madagascar. Afrotherian Conserv. 3, 3–5. Jenkins, P.D., 2003. Microgale, shrew tenrecs. In: Goodman, S.M., Benstead, J.P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, Chicago, pp. 1273–1278.
471
Jenkins, P.D., Goodman, S.M., 1999. A new species of Microgale (Lipotyphla, Tenrecidae) from isolated forest in southwestern Madagascar. Bull. Nat. Hist. Mus., London 65, 155–164. MacPhee, R.D.E., 1987. The shrew tenrecs of Madagascar: systematic revision and holocene distribution of Microgale (Tenrecidae, Insectivora). Am. Mus. Novit. 2889, 1–45. Olson, L., Goodman, S.M., Yoder, A.D., 2004. Reciprocal illumination of cryptic species: morphological and molecular support for several undescribed species of shrew tenrecs (Mammalia: Tenrecidae; Microgale). Biol. J. Linn. Soc. 83, 1–22. Raharivololona, B.M., 1996. Impact de l’exploitation se´lective de la foreˆt et la variation saisonnie`re sur la composition de la population de rongeurs et insectivores dans une foreˆt se`che de l’Ouest de Madagascar. Me´moire DEA. Universite´ d’Antananarivo, Antananarivo. Raxworthy, C.J., Nussbaum, R.A., 1994. A rainforest survey of amphibians, reptiles and small mammals at Montagne d’Ambre, Madagascar. Biol. Conserv. 69, 65–73. Smith, A.P., 1997. Deforestation, fragmentation, and reserve design in western Madagascar. In: Laurance, W.F., Bierregaard, Jr., R.O.W. (Eds.), Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities. University of Chicago Press, Chicago, pp. 415–441. Sorg, J.-P., Ganzhorn, J.U., Kappeler, P.M., 2003. Forestry and research in the Kirindy forest/centre de formation professionnelle Forestie`re. In: Goodman, S.M., Benstead, J.P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, Chicago, pp. 1512–1519.
www.elsevier.de/mambio
SHORT COMMUNICATION
New distributional records of the recently described and endangered shrew tenrec Microgale nasoloi (Tenrecidae: Afrosoricida) from central western Madagascar Voahangy Soarimalalaa,b, Steven M. Goodmanb,c, a
De´partement de Biologie Animale, Universite´ d’Antananarivo, BP 906, Antananarivo (101), Madagascar Vahatra, BP 3972, Antananarivo (101), Madagascar c Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, USA b
Received 27 March 2007; accepted 6 September 2007
Keywords: Microgale nasoloi; Tenrecidae; Western Madagascar; Distribution limit; Conservation status
Recent biological explorations of the dry forests of western and southern Madagascar have revealed that this zone has much higher small mammal species diversity than previously realized. Further, many of these discoveries comprise new mammals to science, including an assortment of bats (e.g., Goodman et al. 2005, 2006a, b, 2007), rodents (e.g., Carleton et al. 2001; Goodman and Soarimalala 2005; Carleton and Goodman 2007), and shrew tenrecs (e.g., Jenkins and Goodman 1999; Goodman and Soarimalala 2004). One of these recently described taxa from the dry forests of the island is the distinctive shrew tenrec Microgale nasoloi, which was described by Jenkins and Goodman (1999) based on two specimens from the southwest. Currently, this species is considered in the IUCN Red Data Book as endangered (www.iucnredlist.org). No other records of M. nasoloi were known until 2006, when two additional specimens were taken a notable distance from the sites of the type series and in a different habitat. Herein we report on these two new specimens collected by Voahangy Soarimalala (VS) and deposited in the Universite´ d’Antananarivo, De´partement de Biologie Animale (UADBA), Antananarivo, Madagascar, while the type series are held by the Field Museum of Natural History (FMNH), Chicago, USA (Table 1). Corresponding author.
E-mail address: sgoodman@fieldmuseum.org (S.M. Goodman).
The original description of M. nasoloi was based on two animals, the holotype coming from the Province de Toliara, Foreˆt de Vohibasia, 59 km NE de Sakaraha 22127.50 S, 44150.50 E, 780 m, in transitional dry deciduous forest, and the other specimen from the Province de Toliara, Foreˆt d’Analavelona, 12.5 km NW d’Andranoheza, 22140.70 S, 44111.50 E, 1050 m, on an isolated massif with elements of eastern humid and western deciduous forest (Jenkins and Goodman 1999; Fig. 1). Although these two sites are only about 70 km from one another, the local habitats are notably different. The central portion of western Madagascar, a region known as the Menabe, has been extensively surveyed for small mammals and the only species of shrew tenrecs known from the region are Microgale brevicaudata and Microgale longicaudata/majori (Ganzhorn et al. 1996; Raharivololona 1996; Olson et al. 2004; Hilgartner 2005). However, during an inventory of two sites in March 2006, two specimens were collected, which were tentatively identified as M. nasoloi and include: (1) Province de Toliara, Fivondronana de Morondava, Commune Rurale de Bemanonga, Kirindy (CFPF), Foreˆt d’Amboropotsy, 16.3 km NE de Marofandilia, 20103.20 S, 44141.00 E, 80 m, in dry deciduous forest (VS-1151, UADBA 48052), 19 March 2006. (2) Province de Toliara, Fivondronana de Belo sur Tsiribihina, Commune Rurale de Beroboka, Foreˆt
1616-5047/$ - see front matter r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.09.002 Mamm. biol. 73 (2008) 468–471
ARTICLE IN PRESS V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
Table 1.
469
Measurements of the four known specimens of Microgale nasoloi
Measurements
UADBA 48052
UADBA 48053
FMNH 156187 (holotype)
FMNH 161575
Sex Total length Head and body length Tail length Hind foot length Ear length Weight Condylo-incisive length Upper toothrow length Rostral breadth Interorbital breadth Braincase breadth Braincase height
~ 130 80 52 11 15 7.3 22.1 9.9 3.0 4.1 9.1 5.0
~ 127 78 50 12 15 8.2 22.4 10.1 3.0 4.2 9.0 5.0
~ – 81 53 13 16 14.0 23.2 10.2 3.1 4.3 9.2 4.9
# – 70 62 14 16 5.9 – – – – – –
All measurements in mm with the exception of weight, which is in gm. The measurements for the two FMNH specimens are taken from Jenkins and Goodman (1999).
Fig. 1. Map of western and southwestern Madagascar showing the four sites Microgale nasoloi has been found and other localities mentioned in the text. The inset map of Madagascar shows the zone delineated in the regional map to the left.
de Lambokely, 11 km NNE de Beroboka, 19152.20 S, 44138.70 E, 85 m, in dry deciduous forest (VS-1166, UADBA 48053), 29 March 2006.
These two specimens were subsequently compared to the different medium-sized Microgale spp. known from Madagascar (Jenkins 2003; Goodman and Soarimalala
ARTICLE IN PRESS 470
V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
2004) and based on various aspects of pelage, external measurements, and cranial and dental morphology are referable to M. nasoloi. All of the salient characters noted in the diagnosis of M. nasoloi by Jenkins and Goodman (1999) are discernable in the new material from the Menabe region. On the basis of tooth eruption patterns UADBA 48053 is at stage 3 (MacPhee 1987), showed no sign of current or past reproduction, and had notable subcutaneous fat deposits. In contrast, UADBA 48052 is younger in age and based on tooth ontogeny patterns fits MacPhee’s (1987) stage 1/2. This latter individual showed no sign of reproductive activity and was presumably nuliparous. At each of the two sites inventoried in 2006, three pitfall lines with plastic drift fences were installed in the forest (see Raxworthy and Nussbaum (1994) for a description of the technique). Each line comprised 11 buckets, 15 l in capacity, and placed at 10 m intervals. At each site, the pit-falls were installed for 6 consecutive nights, comprising an accumulated 198 pit-fall trap days. In the Foreˆt d’Amboropotsy, other than the single specimen of M. nasoloi, the other small mammals captured in these devices included Geogale aurita (n ¼ 1), M. brevicaudata (n ¼ 4) and Setifer setosus (n ¼ 1), while in the Foreˆt de Lambokely associated species included Suncus madagascariensis (n ¼ 1) and S. setosus (n ¼ 1). Hence, even though the surveys took place during the rainy season, when shrew tenrecs are presumed to be active, small mammal pit-fall capture rates were notably low, with seven animals (3.5% capture success) at Foreˆt d’Amboropotsy and three animals (1.5% capture success) in the Foreˆt de Lambokely. In February and March 2007, three different sites in the Kirindy-Mitea National Park were surveyed and 198 pit-fall trap days were accrued at each site and no individual of M. nasoloi was captured. Further, no sign of this species was found during extensive small mammal surveys in the zone north of the Tsiribihina River (V. Soarimalala and S. M. Goodman, unpublished data). The Foreˆt d’Amboropotsy is within the Kirindy (CFPF) forest and close to the research station, which has been an area of intensive biological research for several decades (Sorg et al. 2003), including inventory programs on small mammals (e.g., Ganzhorn et al. 1996; Raharivololona 1996). The discovery of M. nasoloi in this forest block, a species that is either rare or difficult to capture, indicates the importance of continued research with varied techniques and during different seasons to reach a more complete estimation of local small mammal diversity. Further, given the low pit-fall capture rates in the dry forests of Madagascar, it is clear that extensive survey work is needed to approach complete knowledge of the local small mammal fauna. The forest habitat where M. nasoloi was captured in the Menabe region is a seasonally dry deciduous habitat
and distinctly more mesic than the Zombitse-Vohibasia zone near Sakaraha. The Analavelona Massif site is rather particular, as it is notable moister than surrounding habitats associated with the summital zone receiving orographic precipitation. Based on these new records from the Menabe region, the habitat breadth and distribution of M. nasoloi is greatly expanded. The triangular-shaped polygon encompassing the four known localities for this taxon comprises an area of about 10,260 km2. However, whether it occurs across this zone is unclear and given its very recent discovery in the Kirindy (CFPF) area, much more detailed inventories will be needed to properly establish this point. M. nasoloi has been classified by the IUCN (www. iucnredlist.org) as endangered (EN B1ab(iii)), at least in part associated with its previously very limited geographic distribution and limited number of records. The new distributional data presented here clearly indicate that this conservation status needs to be re-evaluated. Further, given the remarkable level of new mammal species discoveries in the dry forests of Madagascar, the notable levels of deforestation of this region (Smith 1997), and high levels of habitat heterogeneity and species turnover, it is clear that this zone of the island needs to be given one of the highest priorities for new conservation actions.
Acknowledgements We are grateful to the Malagasy authorities for permits to conduct this research, which was supported by a grant from the John D. and Catherine T. MacArthur Foundation. Lucienne Wilme´ kindly prepared Figure 1.
References Carleton, M., Goodman, S.M, 2007. A new species of the Eliurus majori complex (Rodentia: Muroidea: Nesomyidae) from south-central Madagascar, with remarks on emergent species groupings in the genus Eliurus. Am. Mus. Novit. 3547, 1–21. Carleton, M., Goodman, S.M., Rakotondravony, D., 2001. A new species of tufted-tailed rat, genus Eliurus (Muridae: Nesomyinae, from western Madagascar, with notes on the distribution of E. myoxinus. Proc. Biol. Soc. Wash. 114, 972–987. Ganzhorn, J.U., Sommer, S., Abraham, J.-P., Ade, M., Raharivololona, B.M., Rakotovao, E.R., Rakotondrasoa, C., Randriamarosoa, R., 1996. Mammals of the Kirindy forest with special emphasis on Hypogeomys antimena and the effects of logging on the small mammal fauna. In: Ganzhorn, J.U., Sorg, J.-P. (Eds.), Ecology and Economy of a Tropical Dry Forest in Madagascar. Primate Report 46-1, pp. 215–232.
ARTICLE IN PRESS V. Soarimalala, S.M. Goodman / Mamm. biol. 73 (2008) 468–471
Goodman, S.M., Soarimalala, V., 2004. A new species of Microgale (Lipotyphla: Tenrecidae: Oryzorictinae) from the Foreˆt des Mikea of southwestern Madagascar. Proc. Biol. Soc. Wash. 117, 251–265. Goodman, S.M., Soarimalala, V., 2005. A new species of Macrotarsomys (Rodentia: Muridae: Nesomyinae) from the Foreˆt des Mikea of southwestern Madagascar. Proc. Biol. Soc. Wash. 118, 450–464. Goodman, S.M., Jenkins, R.K.B., Ratrimomanarivo, F.H., 2005. A review of the genus Scotophilus (Chiroptera: Vespertilionidae) on Madagascar, with the description of a new species. Zoosystema 27, 867–882. Goodman, S.M., Ratrimomanarivo, F.H., Randrianandrianina, F.H., 2006a. A new species of Scotophilus (Chiroptera: Vespertilionidae) from western Madagascar. Acta Chiropter 8, 21–37. Goodman, S.M., Cardiff, S.G., Ranivo, J., Russell, A.L., Yoder, A.D., 2006b. A new species of Emballonura (Emballonuridae: Chiroptera) from the dry regions of Madagascar. Am. Mus. Novit. 3538, 1–24. Goodman, S.M., Rakotondraparany, F., Kofoky, A., 2007. The description of a new species of Myzopoda (Myzopodidae: Chiroptera) from western Madagascar. Mamm. Biol. 72, 65–81. Hilgartner, R.D., 2005. Some ecological and behavioural notes on the shrew tenrec Microgale cf. longicaudata in the dry deciduous forest of western Madagascar. Afrotherian Conserv. 3, 3–5. Jenkins, P.D., 2003. Microgale, shrew tenrecs. In: Goodman, S.M., Benstead, J.P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, Chicago, pp. 1273–1278.
471
Jenkins, P.D., Goodman, S.M., 1999. A new species of Microgale (Lipotyphla, Tenrecidae) from isolated forest in southwestern Madagascar. Bull. Nat. Hist. Mus., London 65, 155–164. MacPhee, R.D.E., 1987. The shrew tenrecs of Madagascar: systematic revision and holocene distribution of Microgale (Tenrecidae, Insectivora). Am. Mus. Novit. 2889, 1–45. Olson, L., Goodman, S.M., Yoder, A.D., 2004. Reciprocal illumination of cryptic species: morphological and molecular support for several undescribed species of shrew tenrecs (Mammalia: Tenrecidae; Microgale). Biol. J. Linn. Soc. 83, 1–22. Raharivololona, B.M., 1996. Impact de l’exploitation se´lective de la foreˆt et la variation saisonnie`re sur la composition de la population de rongeurs et insectivores dans une foreˆt se`che de l’Ouest de Madagascar. Me´moire DEA. Universite´ d’Antananarivo, Antananarivo. Raxworthy, C.J., Nussbaum, R.A., 1994. A rainforest survey of amphibians, reptiles and small mammals at Montagne d’Ambre, Madagascar. Biol. Conserv. 69, 65–73. Smith, A.P., 1997. Deforestation, fragmentation, and reserve design in western Madagascar. In: Laurance, W.F., Bierregaard, Jr., R.O.W. (Eds.), Tropical Forest Remnants: Ecology, Management, and Conservation of Fragmented Communities. University of Chicago Press, Chicago, pp. 415–441. Sorg, J.-P., Ganzhorn, J.U., Kappeler, P.M., 2003. Forestry and research in the Kirindy forest/centre de formation professionnelle Forestie`re. In: Goodman, S.M., Benstead, J.P. (Eds.), The Natural History of Madagascar. The University of Chicago Press, Chicago, pp. 1512–1519.