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27. Red Forest Rat Nesomys rufus French: Nésomys roux / German: Rote Inselratte / Spanish: Rata de bosque roja Other common names: Eastern Red Forest Rat, Rufous Nesomys Taxonomy. Nesomys rufus Peters, 1870, “Ein getrocknetes mannliches Exemplar aus Vohima,” Madagascar. This species is monotypic. Distribution. Endemic to N & E Madagascar. Descriptive notes. Head-body 170-200 mm, tail 160-180 mm; weight 135-185 g. Dorsum of the Red Forest Rat is reddish brown, often mixed with black hair especially in middle of back and more saturated with red along flanks. Venter ranges from uniform rusty red, with some parts, particularly along midline, off-white. Ears are moderately long. Proximal part of tail has short and sparse black hair that becomes progressively slightly longer; distal 8-20 mm oftail is white. Legs, feet, and toes are dark brown. Habitat. Eastern humid lowland and montane forest at elevations of 650-2000 m. The Red Forest Ratis distinctly uncommon above elevations of 1900 m, and occurs sympatrically with Audebert’s Forest Rat (N. audeberti) at certain locations. Food and Feeding. The Red Forest Rat eats seeds of different forest trees, including Cryptocarya (Lauraceae), Canarium (Burseraceae), and Sloanea (Elaecocarpaceae), and caches fatrich seeds such as Canarium. It has been observed in areas of slash-and-burn agriculture within short distances of relatively intact native forest. Breeding. Reproductive season of the Red Forest Rat is from mid-Octoberto late December. Females have three pairs of mammae and are known to have litters of up to four young. Activity patterns. The Red Forest Rat is diurnal and terrestrial, tending to be crepuscular. It lives in complex underground burrow systems, with numerous holes and often placed in root complexes. Burrows have sections with freshly clipped plant material and sleeping chambers with finer plant material. It is preyed on by snakes (boas and the genus Pseudoxyrhopus) and carnivores such as the Ring-tailed Vontsira (Galidia elegans) and the Fosa (Cryproproctaferox). Movements, Home range and Social organization. Home range of the Red Forest Rat has been calculated to average 0-5 ha. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Red Forest Rat is strictly forest-dwelling and is known from widely separated localities across much of the eastern humid forests of Madagascar. It occurs in lowland formations that have been drastically reduced in their extent due to human pressure, and the mediumand long-term future in that habitat is uncertain. Bibliography. Cadle (1999), Carleton & Schmidt (1990), Carleton, Smeenk et al. (2014), Goodman & Carleton (1996), Goodman etal. (2013), Ryan (2003), Ryan et al. (1993), Soarimalala & Goodman (2011), Soarimalala etal. (2001)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Nesomyidae, pp. 156-203 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 192, DOI: 10.5281/zenodo.6600357

25. Audebert’s Forest Rat Nesomys audeberti French: Nésomys d’Audebert / German: Audebert-Inselratte / Spanish: Rata de bosque de Audebert Other common names: Lowland Red Forest Rat, White-bellied Nesomys Taxonomy. Hallomys audeberti Jentink, 1879, “ Maisine and Savary, N. E. Madagascar.” Restricted by M. D. Carleton and colleagues in 2014 to “ Toamasina Province (former), Analanjirofo Région, west of Antongil Bay toward the western frontier of Fivondronana Mananara Avaratra, ‘ Savary.” This species is monotypic. Distribution. Endemic to E Madagascar. Descriptive notes. Head-body 195-203 mm, tail 169-173 mm; weight 193-239 g. Dorsum of Audebert’s Forest Ratis reddish brown, often mixed with black hair especially in middle of back, and venteris largely rusty-red, with central portion and throat white. Snout is elongated, and ears are moderately long. Base oftail has short black hair that becomes progressively longer and denser toward tip, which can be white. Legs, feet, and toes have dark brown fur. Habitat. Eastern humid lowland and montane forest from nearsea level to elevations of ¢.1050 m. Audebert’s Forest Rat has a patchy distribution and at some localities occurs in sympatry with the Red Forest Rat (N. rufus). Food and Feeding. Audebert’s Forest Rat is presumed to be largely granivorous and perhaps on occasion frugivorous or insectivorous. Breeding. Breeding season of Audebert’s Forest Rat coincides with the end of the dry season; reproductive activity begins in July-August, and young are born c.6 weekslater. Females have three pairs of mammae, and maximum recorded litter size is two young. Activity patterns. Audebert’s Forest Rat is diurnal and terrestrial, with a tendency to be crepuscular. It lives in underground in complex burrows with numerous holes and often placed in root complexes. These burrows have sections with freshly clipped plant material and sleeping chambers with finer plant material. Movements, Home range and Social organization. Average home range of Audebert’s Forest Rat is c.1-5 ha, almost three times larger than that of the Red Forest Rat, with some sexual differences (females averaging 0-5 ha and males 1-4 ha). Home ranges do not appear to be defended and can be occupied by a male and numerous females. An Audebert’s Forest Rat can travel a ground distance of ¢.500 m/day. Status and Conservation. Classified as Least Concern on The IUCN Red List. Audebert’s Forest Rat is strictly forest-dwelling and is known from widely separated localities across much of the eastern humid forests of Madagascar. It frequents lowland formations that have been drastically reduced in their extent due to human pressure, and its mediumand long-term future is uncertain. Bibliography. Carleton, Smeenk et al. (2014), Goodman, Ganzhorn & Rakotondravony (2003), Goodman, Soarimalala et al. (2013), Rakotomalala et al. (2007), Ryan (2003), Ryan et al. (1993), Soarimalala & Goodman (2011)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Nesomyidae, pp. 156-203 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 191, DOI: 10.5281/zenodo.6600357

26. Lamberton’s Forest Rat Nesomys lambertoni French: Nésomys de Lamberton / German: Lamberton-Inselratte / Spanish: Rata de bosque de Lamberton Other common names: Western Nesomys, Western Red Forest Rat Taxonomy. Nesomys lambertoni G. Grandidier, 1928, type locality not given. Identified by M. D. Carleton and colleagues in 2014 as “Mahajanga Province (former), Melaky Région, Antsalova District, tsingy habitat at the western margin of the Bemaraha Massif and east of Antsalova.” This species is monotypic. Distribution. Endemic to CW Madagascar (Tsingy de Bemaraha area). Descriptive notes. Head-body 189-195 mm, tail 183-191 mm (two individuals); weight 225-243 g. Lamberton’s Forest Rat is distinctly large and appears squirrel-like from a distance. Dorsum is dark reddish brown, mixed with black hair especially in middle of back, and it becomes more saturated with red along flanks. Venteris light brown. It has an elongated snout and distinctly long ears. Tail is completely covered with long, thick dark brown hair. Legs are dark brown, tending to be black. Habitat. Dry deciduous forest, specifically karst areas forming “tsingy” (limestone pinnacle) habitat, at elevations of ¢.100 m. Lamberton’s Forest Rat makes its dens in small caves and rock shelters in limestone. Food and Feeding. LLamberton’s Forest Rat is presumed to be largely granivorous and frugivorous. Breeding. Female Lamberton’s Forest Rats have three pairs of mammae, and based on limited information, litter size is one young. Activity patterns. Lamberton’s Forest Rat is diurnal and terrestrial, tending to be crepuscular. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Endangered on The IUCN Red List. Lamberton’s Forest Ratis strictly forest-dwelling and is known from a restricted zone of dry deciduous forest that grow on limestone substrate. Its complete known distribution occurs in Tsingy de Bemaraha UNESCO World Heritage Site, which is subjected to fire and other anthropogenic pressures. Bibliography. Carleton, Smeenk et al. (2014), Goodman & Schutz (2003), Goodman, Ganzhorn & Rakotondravony (2003), Goodman, Soarimalala et al. (2013), Ryan (2003), Soarimalala & Goodman (2011)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Nesomyidae, pp. 156-203 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 191-192, DOI: 10.5281/zenodo.6600357

International audience; Since the mid-19th century, the Malagasy subfossil macrofauna has been the object of numerous studies and publications, contrary to the microfauna. New fieldwork, initiated in 2001, in the North West of Madagascar (Province of Mahajanga) led to the discovery of sites rich in fossiliferous breccias, containing micromammals. In this article, we describe two new subfossil species of rodent: Brachytarsomys mahajambaensis, the smallest within the genus and Nesomys narindaensis, the largest within the genus. Most of the extant species of these two genera live in the tropical rainforests of the eastern and north-eastern areas of Madagascar, far away from the north-western part of the island where a dry deciduous forest occurs. The presence of the two taxa in the subfossil record in the Northwest of the country suggests the occurrence of wetter conditions in the past in this region

Nesomys rufus Peters, 1870. Sitb. Ges. Naturf. Fr. Berlin, p. 55. TYPE LOCALITY: Madagascar, Antsiranana Prov., Vohima. DISTRIBUTION: Broadly distributed in N and E forest (audeberti and rufus), and one locality in WC Madagascar (lambertoni). SYNONYMS: audeberti, lambertoni. COMMENTS: Ellerman (1941, 1949a) listed audeberti and lambertoni as species as described, but Petter (1972c, 1975a) arranged them as subspecies of N. rufus. Carleton and Schmidt (1990) indicated that each will stand as a species upon revision., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 679, DOI: 10.5281/zenodo.7353098, {"references":["Ellerman, J. R. 1941. The families and genera of living rodents. Vol. II. Family Muridae. British Museum (Natural History), London, 690 pp.","Ellerman, J. R. 1949 a. The families and genera of living rodents. Vol. III, Appendix II [Notes on the rodents from Madagascar in the British Museum, and on a collection from the island obtained by Mr. C. S. Webb]. British Museum (Natural History), London, 210 pp.","Petter, F. 1972 c. The rodents of Madagascar: The seven genera of Malagasy rodents. Pp. 661 - 666, in Biogeography and ecology in Madagascar (R. Battistini and G. Richard-Vincard, eds.). Monographiae Biologicae, 21: 1 - 765.","Petter, F. 1975 a. Family Cricetidae: Subfamily Nesomyinae, Part 6.2. Pp. 1 - 4, in The mammals of Africa: An identification manual (J. Meester and H. W. Setzer, eds.) [issued 10 Dec 1975]. Smithsonian Institution Press, Washington, D. C., not continuously paginated.","Carleton, M. D., and D. F. Schmidt. 1990. Systematic studies of Madagascar's endemic rodents (Muroidea: Nesomyinae): An annotated gazetteer of collecting localities of known forms. American Museum Novitates, 2987: 1 - 36."]}

Nesomys Peters, 1870. Sitzb. Ges. Naturf. Fr. Berlin, p. 54. TYPE SPECIES: Nesomys rufus Peters, 1870. SYNONYMS: Hallomys. COMMENTS: Major (1897) synonymized Jentink's (1879) Hallomys (type species = H. audeberti) under Nesomys and allocated the genus to the Cricetinae., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 679, DOI: 10.5281/zenodo.7353098, {"references":["Major, C. I. F. 1897. On the Malagasy rodent genus Brachyuromys; and on the mutual relations of some groups of the Muridae (Hesperomyinae, Microtinae, Murinae, and \" Spalacidae \") with each other and with the Malagasy Nesomyinae. Proceedings of the Zoological Society of London, 1897: 695 - 720."]}

Nesomys rufus Peters, 1870. Sitzb. Ges. Naturf. Fr. Berlin, p. 55. TYPE LOCALITY: Madagascar, Vohima. DISTRIBUTION: Madagascar. COMMENT: Includes audeberti and lambertoni; see Petter, 1975, Part 6.2:3. ISIS NUMBER: 5301410008066003001 as N. rufus. 5301410008066001001 as N. audeberti. 5301410008066002001 as N. lambertoni., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 3), pp. 392-476 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 432, DOI: 10.5281/zenodo.7353031

Nesomys Peters, 1870. Sitzb. Ges. Naturf. Fr. Berlin, p. 54. REVIEWED BY: R. D. Owen (RDON). COMMENT: Subfamily Nesomyinae; see comment under Cricetidae. ISIS NUMBER: 5301410008066000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 3), pp. 392-476 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 432, DOI: 10.5281/zenodo.7353031

Three previously described members of the Madagascar Haemaphysalis (Rhipistoma) elongata group have a strong ventral spine on palpal segment 3 and exceptionally long and numerous spines on the coxae and trochanters. These species parasitize spiny insectivores (Family Tenrecidae). In H. (R.) anoplos sp. n., a parasite of the soft-furred Nesomys rufus Peters (Rodentia, Nesomyinae), all these spines have become obsolete. H. (R.) simplex Neumann, the other member of this group and also a spiny insectivore parasite, has only moderately developed palpal and coxal spurs. H. (R.) anoplos sp. n. is postulated to represent an extreme reversal to the "simplex type" of spur structure and to reflect a functional adaptation to the fine fur of the host. H. (Dermaphysalis) nesomys Hoogstraal, Uilenberg, and Klein, another parasite of Nesomys rufus, has similarly reduced spurs and has diverged so markedly in other structural characteristics from its Rhipistoma or Rhipistoma-like ancestors that a new subgeneric taxon was created for it. Haemaphysalis (Rhipistoma) anoplos sp. n., described below, appears to represent an extreme reversal from the uniquely strongly spined members of the H. (R.) elongata group (elongata Neumann, subelongata Hoogstraal, and tiptoni Hoogstraal) to the more primitively spined structural type seen in H. (R.) simplex Neumann. The four previously described members of this group parasitize spiny insectivores (Family Tenrecidae); the newly described species parasitizes the soft-furred Nesomys rufus Peters (Rodentia, Nesomyinae), from which the structurally remarkable H. (Dermaphysalis) nesomys was recently described by the present writers (1966). Received for publication 21 April 1967. *From Research Project MR005.09-1402.3, Bureau of Medicine and Surgery, Department of the Navy, Washington, D.C. The opinions and assertions contained herein are the private ones of the authors and are not to be construed as official or as reflecting the views of the Department of the Navy or of the naval service at large. The illustrations in this report were prepared under the auspices of Agreement 352505 between the National Institutes of Health and NAMRU-3. t Head, Department of Medical Zoology, United States Naval Medical Research Unit Number Three, Cairo, Egypt, U. A. R. t Chef du Service d'Entomologie et Protozoologie, Laboratoire Central de l'Elevage, Institut d'Elevage et de Medecine Veterinaire des Pays Tropicaux, Tananarive, Madagascar. ? Entomologiste Medical, O.R.S.T.O.M., Institut Pasteur, Phnom-Penh, Cambodia. Haemaphysalis (Rhipistoma) anoplos sp. n. The Madagascar Spurless Rodent-Haemaphysalid (Figs. 1-10)