Search

199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 667-668, DOI: 10.5281/zenodo.6887260

204. Long-nosed Luzon Forest Mouse Apomys sacobianus French: Apomys de la Sacobia / German: Langnasen-Philippinenwaldmaus / Spanish: Raton de bosque de Luzon de hocico largo Other common names: Long-nosed Luzon Apomys, Pinatubo Volcano Mouse Taxonomy. Apomys sacobianus D. H. Johnson, 1962, “Sacobia River, Clark Air Base, Pampanga Province, Luzon, Philippine Islands.” Apomys sacobianus, a member ofsubgenus Megapomys, may be most closely related to A. lubangensis. It co-occurs with A. zambalensis on the slopes of Mount Pinatubo, and was the more commonly captured of the two species. Published records of A. sacobianus from Isabela, Mountain, and Zambales Provinces of Luzon were disputed by G. G. Musser and M. D. Carleton in 2005, and are no longer included within this species. Monotypic. Distribution. Recorded only from foothills and lower slopes of Mt Pinatubo, Zambales Mts, C Luzon, Philippines; it may be more widely distributed in Zambales Mts, which remain poorly surveyed for small mammals. Descriptive notes. Head-body 135-158 mm,tail 115-159 mm, ear 21-25 mm, hindfoot 34-40 mm; weight 71-105 g. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite offeatures, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on eachside, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground; in most forest habitats across Luzon Island they are the most abundant terrestrial mammal in any given community. The Long-nosed Luzon Forest Mouse, a relatively large Megapomys, has fur on upperparts brown with grayish tones, fur on underparts and inside of limbs overall grayish white (hairs gray at bases and with short white tips), the border between upperside and underside being slightly diffuse; head has whitish-gray fur on throat, cheeks, and sides of snout, and often apatch ofdarker fur on top ofsnout. Dark fur on limbs terminates at wrist and ankle, upper surfaces offoreand hindfeet being mostly white with scattered dark hairs; hindfeet relatively long but moderately broad, undersurface heavily pigmented, all plantar pads well separated, digits relatively long and bearing sharp, curved claws. Tail is relatively long (95-100% of head-body length), unusually thick, and sharply bicolored, nearly black above for entire length and pure white or with scattered dark scales and hairs below. Cranium is similar to those of other Megapomys, but somewhat more gracile and with smaller molars than in co-occurring A. zambalensis. Habitat. Original specimen of the Long-nosed Luzon Forest Mouse was caught in lowland secondary forest, and subsequent observations of this species in sparse, regenerating vegetation confirmed that it ishighly tolerant of disturbance. The species survived the violent eruption of Mount Pinatubo in June 1991. Recorded between ¢.100 m and at least 1080 m. Food and Feeding. Probably omnivorous. Recent observations found that this species eagerly consumed both live earthworms and fried coconut baits. Breeding. Five pregnant females each had two embryos, and males have testes measuring up to 13 mm x 6 mm. Activity patterns. The Long-nosed Luzon Forest Mouse was recently reported as being active “almost exclusively at night” and very rarely climbing above ground surface (and “never more than 1 m above ground”). Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. For many years the Long-nosed Luzon Forest Mouse was known only from the holotype, which came from an area impacted by the 1991 eruption of Mount Pinatubo, butit was rediscovered in 2011-2012 and found to be abundant on the slopes of Pinatubo to at least 1080 m. Bibliography. Balete et al. (2009), Johnson (1962), Kennerley (2016h), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Heaney, Balete, Veluz et al. (2014), Musser (1982a), Musser & Carleton (1993, 2005)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 669-670, DOI: 10.5281/zenodo.6887260

195. Mindanao Mossy Forest Mouse Apomys hylocoetes French: Apomys des mousses / German: Nebelwald-Philippinenwaldmaus / Spanish: Ratén de bosque de Mindanao Other common names: Mindanao Apomys, Mount Apo Forest Mouse Taxonomy. Apomys hylocoetes Mearns, 1905, “Mount Apo at 6,000 feet altitude [= 1829 m], southern Mindanao, Philippine Islands.” Original spelling hylocoetes is a latinized Greek word and cannot be modified, so widely used specific epithet hylocetes has been changed. Apomys hylocoetes is type species of genus and subgenus. Molecular evidence suggests special affinity with A. insignis from Mindanao, A. caminguensis from (Camiguan, and an undescribed species from Bohol, Leyte, Samar, and Biliran. Specimens from 2286 m on Mount Apo have been described as a furtherspecies, A. petraeus, but differences are slight and very likely due to age and elevational differences; A. hylocoeles 1s sympatric with A. insignis at 1829 m on Mount Apo. Mitochondrial sequences do not distinguish A. hylocoetes from sympatric A. insignis from Mount Kitanglad Range, but this may be due to introgression or mitochondrial capture as the two species differ markedly in karyotype. Currently regarded as monotypic but in need of further assessment. Distribution. Mindanao I, S Philippines, where recorded from several localities in Kitanglad Range at 1900-2800 m and from Mt Apo at 1829-2286 m;it probably occurs in the Kamangkil-Lumut Ranges that extend into L.anao del Sur Province. Descriptive notes. Head-body 111-115 mm, tail 120-150 mm, ear 18-22 mm, hindfoot 29-34 mm; weight 33-45 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes,relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed species typically most active in canopy. The Mindanao Mossy Forest Apomys has fur on upperparts, flanks and outer surfaces of limbs dense and very soft, without spines or stiff hairs, dark brown without “saltand-pepper speckling”; fur on underparts and inside of limbs is “deep buffy gray”; colors of upperside and underside merge gradually on flanks; ears relatively longer than in sympatric A. insignis, darkly pigmented, and with short hairs on outer surface; eyes without any conspicuous emargination. Dark fur of limbs extends onto upper surfaces of foreand hindfeet, but digits are unpigmented and covered with shorter white hairs; hindfeet relatively shorter and wider than in other members of subgenus Apomys, undersurface lightly pigmented, plantar pads small and well separated, hallucal pad short and positioned farther back than in other species of subgenus Apomys, digits moderately long and slender, with delicate and sharply pointed claws. Tail long (averaging c.130% of head-body length), clearly bicolored, brown above and pale gray below for nearly entire length, terminating in short white tip;tail scales conspicuous but relatively small, in 14-15 rows per cm near tail base, and usually with three short hairs per scale, hair length increasing toward tail tip but without terminal brush. Cranium is very similar to that of other members of subgenus Apomys, but braincase rounder and more inflated than in sympatric A. insignis. Chromosomal complement has 2n = 48 and FN = 56, autosomes include one pair of small submetacentric, three pairs of small subtelocentric, and 20 pairs of telocentric chromosomes graded large to small. Sex chromosomes not obvious, hence X and Y chromosomes presumably both telocentric. Habitat. Forest inhabited by Mindanao Mossy Forest Mice is evergreen tropical rainforest of the kinds categorized by Philippine ecologists as “montane forest” and “mossy forest,” at elevations of ¢.1800-2800 m. The species occurs in areas of natural disturbance such as regeneration after landslides. Original captures of this species were made in primary forest and at higher elevation, among rocks above forest zone. Food and Feeding. The Mindanao Mossy Forest Mouse is thought to be omnivorous. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List (as A. hylocetes). This species appears to be most common in primary mossy forest, where it is the most abundant murid species. It is less abundant at lower elevations in primary montane forest. Populations are present in three National Parks (Mount Kitanglad, Mount Apo, and Mount Kalatungan Range). Bibliography. Heaney (2016b), Heaney, Balete et al. (1998), Heaney, Tabaranza, Rickart et al. (2006), Mearns (1905), Musser (1982a), Musser & Carleton (2005), Rickart & Heaney (2002)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 666, DOI: 10.5281/zenodo.6887260

192. Camiguin Forest Mouse Apomys camiguinensis French: Apomys de Caminguin / German: Camiguin-Philippinenwaldmaus / Spanish: Ratén de bosque de Camiguin Other common names: Camiguin Apomys Taxonomy. Apomys camiguinensis Heaney & Tabaranza, 2006, “Barangay Kital-is, Sagay Municipality, 2 km N, 62 kw W Mahinog, 1000 m elevation, Camiguin Province, Camiguin Island. 9°9.5°N, 124°43.5’E.” A member of subgenus Apomys, A. camiguinensis was newly collected in 1994 and, prior to formal description, was referred to as“ Apomys sp. D.” Molecular evidence suggests possible special affinity with A. Aylocoetes and A. insignis from Mindanao Island, and with an undescribed species of Apomys from Leyte and Biliran. Monotypic. Distribution. Camiguin I, S Philippines. Descriptive notes. Head-body 106 mm, tail 140-160 mm, ear 18-20 mm, hindfoot 31-34 mm; weight 33-48-5 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond,relatively large and thinly furred ears, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed species active primarily in canopy. The Camiguin Forest Mouse is a medium-sized species of subgenus Apomys. Fur on upperparts, flanks, and outer surfaces of limbs is dense and very soft, without spines or stiff hairs, rich brownish russet with small amount of “salt-andpepper” speckling; fur on underparts and inside of limbs is usually nearly white with a buffy or pale russet wash, sometimes with pure white blaze on chest; colors of upperside and lower parts merge gradually; ears are darkly pigmented and with short hairs on outer surface; eyes ringed with thinly furred pale skin. Dark fur of limbs extends onto upper surfaces of foreand hindfeet, but digits are unpigmented and covered with shorter white hairs; hindfeet relatively elongate but somewhat broader than those of some other members of subgenus Apomys, undersurface lightly pigmented, plantar pads small and well separated, hallucal pad short but well forward, digits moderately long. Tail is relatively long (averaging slightly longer than head-body length), sharply bicolored, dark brown above and white below for entire length, tail scales conspicuous and relatively large, in 12-125 rows per cm near tail base, and usually with three short hairs per scale, hairiness increasing toward tail tip but without terminal brush. Cranium very similar to those of other members of subgenus Apomys, but braincase and orbit unusually long, and rostrum long but robust. Habitat. Known only from evergreen tropical rainforest on Mount Timpoong,at elevations of ¢.1000-1400 m. It was common in 1994-1995 in disturbed lowland rainforest at 1000 m and in undisturbed montane forest at 1275 m, and less so in primary mossy forest at 1200-1400 m. It is likely to occur on other forested peaks on Camiguin Island, but is probably absent from extensively deforested areas below 1000 m. Food and Feeding. No information. Breeding. A pregnant female captured in May 1994 had a single embryo, while two other adult females captured in May 1994 and March 1995 each had two placental scars. Adult females with enlarged teats weighed 37-5-42 g, whereas nulliparous females weighed up to 35 g. Adult males with scrotal testes weighed 36-5-48 g, whereas immature males with abdominal testes weighed 34 g or less. Activity patterns. All individuals captured in 1994-1995 were trapped on the ground among roots or beneath fallen and rotten logs, and at night. The Camiguin Forest Mouse is likely to be scansorial or arboreal, as are other species of subgenus Apomys. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Vulnerable on The IUCN Red List. The Camiguin Forest Mouse was locally abundant in 1994-1995 in both primary and disturbed habitats but its current status is unknown. Its known population is within Timboong Hibok-Hibok Natural Monument. Bibliography. Heaney (2017), Heaney & Tabaranza (1997 2006a, 2006b), Heaney et al. (2011), Heaney, Tabaranza, Balete & Rigertas (2006), Justiniano et al. (2015), Musser & Carleton (2005).

193. Northern Luzon Forest Mouse Apomys datae French: Apomys du Data / German: Mount-Data-Philippinenwaldmaus / Spanish: Ratén de bosque de Luzon septentrional Other common names: Cordillera Forest Mouse, Luzon Montane Forest Mouse, Northern Luzon Apomys Taxonomy. Mus datae A. B. Meyer, 1899, “Berge Data, Lepanto, Nord Luzon [Is- land], von 8000 Fuss [= Mount Data, Mountain Province, Luzon Island, Philip- pines, 2438 m].” Apomys datae was included variously in Mus, Epimys, and Rattus before J. R. Ellerman cor- rectly affiliated datae with hylocoetes, placing them in subgenus Apomys of Rattus. D. H. Johnson in 1962 was the first to use the combination Apomys datae. G. S. Miller described Apomys majorin 1910 from a single specimen and without comparing it with Meyer's taxon; its similarity to A. dataewas noted by G. G. Musser in 1982, a conclusion upheld by all subsequent analyses. A. dataeis not fully differentiated from A. abrae on published mitochondrial sequences, but this is interpreted as evidence of introgression following occasional hybridization on Mount Data. Genetic and morphometric analyses demonstrate phylogeographic structure among the various isolated populations of A. datae. This speciesis sympatric with A. abrae over a relatively narrow elevational band between 1500 m and 1650 m, but the two species are generally segregated locally by habitat, A. datae favoring wetter, more closed situations. Currently regarded as monotypic but in need of further assessment. Distribution. Recorded from high elevations in the S two-thirds of the Cordillera Central, NW Luzon I, Philippines. Previous accounts of A. datae in the Sierra Madre and in the lowlands of Luzon refer to other species of Apomys that remained unrecognized until 2011. Descriptive notes. Head—body 125-155 mm, tail 121-139 mm, ear 18-23 mm, hindfoot 33-39 mm; weight 63-105 g. Males average slightly more than females for all measurements, including weight (87-8 g vs. 77-2 g). Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species that are active primarily on ground; in most forest habitats across Luzon, they are the most abundant terrestrial mammal in any local community. The Northern Luzon Forest Mouse is a medium-sized Megapomys. Fur on upperparts is dark brown with slight rustyorange tint and fine black flecking, the latter produced by short black guard hairs, body hairs with dark gray bases; fur on underparts and on inside of limbs is dark gray at base with white or buff tipping; upperside and venter colors with sharply defined boundary; skin oflips, lower snout, and base of ears darkly pigmented; dark fur of limbs extends part way onto upper surfaces of foreand hindfeet, but digits unpigmented and with short white hairs, sometimeslightly peppered with black; hindfeet are relatively elongate and narrow, undersurface with dark gray pigment for proximal half, digits relatively short and stocky, plantar pads moderately large but well separated. Tail is relatively short (on average 89-95% of head-body length in various populations), dark above and white below for entire length, with dark tail hairs on undersurface. Cranium is larger and more robust than that of most Megapomys, with long, stout rostrum but relatively small molars. Chromosomal complement has 2n = 44 and FN 2 54, detail not well documented but probably most similar to arrangement in A. abrae. A karyotype originally reported by E. A. Rickart and L.. R. Heaney in 2002 as an exemplar of A. dataeis now attributed to A. abrae. Habitat. Abundant in evergreen tropical montane forests, including both montane and “mossy” types distinguished in Philippine ecological literature. It occurs also in related secondary forest communities, as well as in shrubby habitats adjacent to agricultural fields, but is generally absent from the agricultural landscape and from pine forest (except when broadleaf forest is regenerating below pine canopy). Elevational limits are 1500-2800 m. Food and Feeding. Diet reported consists of earthworms,insects, and seeds. Breeding. Heaney and coworkers reported in 2016 that “twenty-one pregnant females had an average of 1:66 embryos (range =1-3).” The type specimen contained “two small foetuses.” Activity patterns. The Northern Luzon Forest Mouse is reported as being nocturnal and foraging on ground. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. This species is locally abundant in both primary and disturbed habitats, and is recorded from various protected areas, including Mount Pulag National Park. Bibliography. Danielsen et al. (1994), Ellerman (1941), Heaney (2016a), Heaney, Balete, Dolar et al. (1998), Heaney, Balete, Gee et al. (2005), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Heaney, Balete, Rickart, Veluz & Sarmiento (2004), Heaney, Balete, Rosell-Ambal et al. (2013), Johnson (1962), Justiniano et al. (2015), Mallari & Jensen (1993), Miller (1910), Musser (1982a), Musser & Carleton (1993, 2005), Rabor (1955), Rickart & Heaney (2002), Rickart, Balete, Rowe & Heaney (2011), Rickart, Balete & Heaney (2007), Rickart, Heaney et al. (2011), Sanborn (1952a), Stuart et al. (2007)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 665

188. Luzon Cordillera Forest Mouse Apomys abrae French: Apomys d’Abra / German: Cordillera-Philippinenwaldmaus / Spanish: Raton de bosque de Luzon Other common names: Cordillera Pine Forest Mouse, Luzon Cordillera Apomys Taxonomy. Rattus (Apomys) abrae Sanborn, 1952, “Massisiat, altitude 3500 feet [= 1067 m |], Abra Province, Luzon Island, Philippine Islands.” Apomys abrae is a member of subgenus Megapomys, erected by L. R. Heaney and cow-orkers in 2011. Originally described as a species of subgenus Apomys within Rattus, reflecting J. R. Ellerman’s mistaken view that many Murinae of generalized body form and cranio-dental morphology were closely related to core species of genus Rattus. The combination Apomys abrae was first used by D. H. Johnson in 1962. A. abrae is not fully differentiated from A. datae in published mitochondrial sequences, but this is interpreted as evidence of introgression following occasional hybridization on Mount Data. Genetic and morphometric analyses failed to identify any strong phylogeographic structure within the range of A. abrae. Currently regarded as monotypic but in need of further assessment. Distribution. Cordillera Central of N Luzon I, N Philippines. Descriptive notes. Head-body 121-141 mm, tail 120-144 mm,ear 21-24 mm, hindfoot 33-39 mm; weight 50-79 g. Published mensural data demonstrate weak male-biased sexual dimorphism in external and cranio-dental measures for this species. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; they are distinguished from other Philippine murines by a distinctive suite offeatures, including long, narrow hindfeet with sharp, recurved claws on all digits, thinly furred tail with weakly overlapping scales and three hairs per tail scale, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both 1nguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species that are active primarily on ground; in most forest habitats across Luzon, they are the most abundantterrestrial mammalin any local community. The Luzon Cordillera Forest Mouse is a medium-sized Megapomys. Fur on upperparts, flanks, and outer surfaces of foreand hindlimbsis rich dark brown with slight rusty tint and black flecking, body hairs dark gray basally, guard hairs black and narrowly projecting through fur; fur on underparts and inside of foreand hindlimbs dark gray at bases, with white tips; upperside and lower parts with sharply defined boundary. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and covered with short white hairs, sometimes lightly peppered with black; hindfeet relatively elongate and narrow, undersurface with pale gray pigment for proximal half, plantar pads small and well separated, digits relatively long. Tail is relatively long (averaging 97-101% of head-body length), dark above and white below for entire length,tail hairs dark on upper surface but all white on underside. Cranium is smaller than that of most Megapomys, with moderately short and slender rostrum and relatively large molars. Chromosomal complement has 2n = 44 and FN = 54, autosomes including two pairs large submetacentric, two pairs small subtelocentric, one pair very small metacentric, remainder telocentric; X and Y chromosomes both telocentric, X large, Y very small. This karyotype originally reported by E. A. Rickart and Heaney in 2002 as belonging to the Northern Luzon Forest Mouse (A. datae). Habitat. [Luzon Cordillera Forest Mice have been recorded in dense evergreen montane rainforest, mixed grass and shrubs beneath pines, and in pine forest with broadleaf understory, at elevations of 925-2200 m. In southern part of Central Cordillera, where it co-occurs with the Northern Luzon Forest Mouse between 900 m and 1650 m, the Luzon Cordillera Forest Mouse is generally found at lower elevations and in relatively drier and more open habitats. Food and Feeding. According to Heaney and coworkers in 2016, Luzon Cordillera Forest Mice are “omnivorous, consuming seeds, insects, and earthworms.” Breeding. Heaney and coworkers reported in 2016 that “four pregnant females had an average oftwo embryos (range = 1-3).” Activity patterns. Luzon Cordillera Forest Mice are nocturnal and have been reported as foraging on surface of ground. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Locally abundant in both primary and disturbed habitats. Bibliography. Heaney, Balete, Dolar et al. (1998), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Johnson (1962), Justiniano et al. (2015), Kennerley (2016e), Rabor (1955), Rickart & Heaney (2002), Rickart, Balete & Heaney (2007), Rickart, Balete, Rowe & Heaney (2011), Rickart, Heaney et al. (2011), Sanborn (1952a)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 663, DOI: 10.5281/zenodo.6887260

196. Mindanao Montane Forest Mouse Apomys insignis French: Apomys remarquable / German: Mindanao-Philippinenwaldmaus / Spanish: Ratén de bosque de montana de Mindanano Other common names: Great Apo Mouse, Mindanao Montane Forest Apomys Taxonomy. Apomys insignis Mearns, 1905, “Mount Apo at 6,000 feet altitude [= 1829 m], southern Mindanao, Philippine Islands.” Apomys insignis 1s a member of subgenus Apomys. Form A. bardus, from Mount Bliss in the Grand Malindang Mountains of eastern Mindanao, has usually been treated as a subspecies of insignis, but more recently as a synonym. Mitochondrial sequences do not distinguish A. insignis from sympatric A. hylocoetes from Mount Kitanglad Range, but this may be due to introgression or mitochondrial capture as the two species differ markedly in karyotype; the two are sympatric also at 1829 m on Mount Apo, Mindanao. Currently regarded as monotypic but in need of further assessment. Distribution. Philippines, where it isrestricted to islands of Mindanao and Dinagat. Descriptive notes. Head-body 71-123 mm,tail 134-176 mm, ear 18-21 mm, hindfoot 31-35 mm; weight 33-52 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long narrow hindfeet, thinly furred tail, unreduced eyes, relative large and thinly furred ears,vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed species typically most active in canopy. The Mindanao Montane Forest Mouse has fur on upperparts, flanks and outer surfaces of limbs dense and very soft, without spines or stiff hairs, dark, rich brown with “burnished highlights”; fur on underparts and inside of limbs is variably rich dark buff suffused with dark gray or grayish pale buff, many with pure white patches on midline of chest and abdomen; colors of upperside and underside merge gradually on flanks; ears relatively shorter than in sympatric A. hylocoetes, darkly pigmented, and with short hairs on outer surface; eyes without any conspicuous emargination. Dark fur of limbs extends onto upper surfaces of foreand hindfeet, but digits are unpigmented and covered with shorter white hairs; hindfeet relatively longer and narrower than in other members of subgenus, undersurface lightly pigmented, plantar pads small and well separated, hallucal pad short and positioned farther forward than in A. hylocoetes, digits moderately long and slender, with sharply pointed claws. Tail is relatively long (averaging 130-140% of head-body length), weakly to strongly bicolored to tip or with short white tip, brown above and variably brown, gray or white below, often mottled; tail scales conspicuous, in 13-14 rows per cm neartail base, and usually with three short hairs per scale, hair length increasing toward tail tip but without terminal brush. Cranium is very similar to that of other members of subgenus Apomys, but braincase rounder and more inflated than in sympatric A. insignis. Chromosomal complement has 2n = 36 and FN = 36, and autosomes include five pairs of small to large metacentric, three pairs of submetacentric, and ten pairs of subtelocentric chromosomes graded medium to small. Sex chromosomes not obvious, hence X and Y chromosomes presumably both subtelocentric. Habitat. Restricted to evergreen tropical forest, including both lowland and montane rainforests, and extending narrowly into mossy forest. The Mindanao Montane Forest Mouse appears to be most common in primary montane forest but occursalso in areas of regeneration after natural and low-intensity anthropogenic disturbance. Its less often encountered in lowland contexts where deforestation has destroyedmuch ofits original habitat. One specimen was captured in a coffee plantation at 899 m on Mount McKinley. Food and Feeding. No information. Breeding. No information. Activity patterns. Many captures of Mindanao Montane Forest Mice have been made on the ground among tree roots, under logs and among boulders. The species’ relatively long tail also points to a significant degree of arboreal activity. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Populations are present in three National Parks (Mount Kitanglad, Mount Apo, and Mount Kalatungan Range). The Mindanao Montane Forest Mouse is widespread on Mindanao, from near sea level to high elevations, but lowland populations are fragmented owing to forest loss. The population on Dinagat remains poorly documented. Bibliography. Heaney, Balete et al. (1998), Heaney, Tabaranza & Balete (2016), Heaney, Tabaranza, Rickart et al. (2006), Hoogstraal (1951), Justiniano et al. (2015), Mearns (1905), Musser (1982c), Musser & Carleton (2005), Rickart & Heaney (2002), Sanborn (1952a)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 666-667, DOI: 10.5281/zenodo.6887260

190. Banahaw Forest Mouse Apomys banahao French: Apomys du Banahaw / German: Banahaw-Philippinenwaldmaus / Spanish: Raton de bosque de Banahaw Other common names: Banahaw Apomys Taxonomy. Apomys banahao Heaney et al., 2011, “Philippines: Luzon Island: Quezon Province: Mt. Banahaw: Barangay Lalo, 1465 m; 14-06635°N, 121-50855°E..” Apomys banahao was newly collected in 2004. It belongs to subgenus Megapomys and is most closely related to A. brownorum. It is sympatric at mid-elevations on Mount Banahaw with A. magnus of subgenus Megapomys, and across broader elevational range with A. microdon and A. musculus of subgenus Apomys. Monotypic. Distribution. Currently known only from Mt Banahaw and adjacent Mt Banahaw de Lucban, Luzon, Philippines;it is likely to occur on adjacent Mt San Cristobal. Descriptive notes. Head-body 125-154 mm,tail 111-133 mm, ear 22-24 mm, hindfoot 33-37 mm; weight 71-92 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; they are distinguished from other Philippine murines by distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes,relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground; in most forest habitats across Luzon, they are the most abundant terrestrial mammal in any local community. The Banahaw Forest Mouse is one of the smaller species of Megapomys. Fur on upperparts, flanks and outer surfaces of all limbs is dense and moderately long, dark brown with pale rusty tints and diffuse black flecking, paler on flanks; body hairs dark gray basally, guard hairs black and narrowly projecting through fur; fur on underparts and on inside of limbs is dark gray at bases with pale ash-gray tips; upperside and lower parts with sharply defined boundary; ears are darkly pigmented. Dark fur of limbs extends onto upper surfaces of forefeet, but digits are unpigmented and covered with short white hairs; dark fur of hindlimbs stops at ankle, but patch of dark hairs is present on upper surface of each foot; hindfeet relatively elongate and narrow, undersurface with dark gray pigment for proximal two-thirds, surrounding posterior pads, most plantar pads small and well separated but hallucal pad long and narrow, digits relatively short. Tail is relatively short (86-103% of head-body length), dark above and white below for entire length, but sometimes with scattered pigmented scales and dark hairs. Cranium has moderately long and slender rostrum, and relatively small molars for the genus. Chromosomal complement has 2n = 48 and FN> 54, autosomes with two pairs small metacentric orsubmetacentric, remainder telocentric; X chromosome is large and submetacentric, Y is very small and telocentric. Habitat. Evergreen tropical rainforest, including primary montane rainforest and ecotone with higher-elevation “mossy” forest, at elevations of 1465-1750 m. Banahaw Forest Mice have notbeen encountered in disturbed habitats. Food and Feeding. [.. R Heaney and colleagues reported in 2016 that Banahaw Forest Mice “showed a slight preference for live earthworm bait rather than fried coconut” and “are probably omnivorous.” Breeding. R. Heaney and coworkers reported that “nine pregnant females carried from one to two embryos each (mean = 1:9).” Activity patterns. Heaney and coworkers reported that “nearly all were captured at night, and nearly all were on the ground surface (not in trees).” Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The entire known geographic range of the Banahaw Forest Mouse falls within the Mts Banahaw—San Cristobal National Park. Bibliography. Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Heaney, Balete, Rosell-Ambal et al. (2013), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 664, DOI: 10.5281/zenodo.6887260

203. Least Philippine Forest Mouse Apomys musculus French: Petit Apomys / German: Zwerg-Philippinenwaldmaus / Spanish: Raton de bosque minimo de Filipinas Other common names: Least Forest Mouse, Least Philippine Apomys Taxonomy. Apomys musculus G. S. Miller, 1910, “Camp John Hay, Baguio, Benguet, Luzon (altitude 5000 ft [= 1524 m]),” Philippines. Apomys musculus is a member ofsubgenus Apomys and is most closely related to A. microdon. It is locally sympatric with latter on Luzon, but often replaces it at higher elevations;it is presumably sympatric with A. gracilirostris on Mindoro and with A. insignis on Dinagat. G. G. Musser and M. D. Carleton in 2005 gave range of A. musculus as Luzon, Dinagat, and Mindoro, but commented (citing unpublished work by LL. R. Heaney as a source) that what is now referred to as A. musculus may in fact represent three species: typical musculus in northern Luzon, a separate species in southern Luzon, and a furtherspecies on Mindoro. Heaney and others in 2016 gave distribution of A. musculus as “only on Luzon, where they are widespread.” Currently regarded as monotypic but in need offurther assessment. Distribution. Luzon, Mindoro, and DinagatIs, Philippines. Descriptive notes. Head-body 76-124 mm, tail 97-121 mm, ear 14-17 mm, hindfoot 20-25 mm; weight 16-24 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet with sharp, recurved claws on all digits, thinly furred tail with weakly overlapping scales and three hairs per tail scale, unreduced eyes, relatively large and thinly furred ears,vibrissae on snout very elongate and,folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed species active primarily in canopy. The Least Philippine Forest Mouse is smallest member of both subgenus and genus. Fur on its upperparts, flanks, and outer surfaces of limbs is overall bright tawny brown,lightly flecked with black on back and with brighterorange tints on flanks and cheeks, while that on underparts and inside oflimbs is rich ocherous buff or buffy gray, often with white patches on midline of chest and abdomen; abrupt border between upperside and underside. Dark hues of foreand hindlimbs extend to ankle and wrist, giving way to buffy white on foreand hindfeet, and white hairs on digits; hindfeet elongate and narrow, with six plantar pads. Tail is equal to or slightly longer than head-body length, brown above to tip and white or mottled with pale brown below, with small but conspicuous scales and short hairs. Cranium has globular braincase and relatively short, narrow rostrum. Chromosomal complement has 2n = 42, FN = 52; autosomes include one pair of medium-sized and two pairs of small metacentric, one pair of large submetacentric, one pair oflarge subtelocentric, and 16 pairs oftelocentric chromosomes graded from large to small. Sex chromosomes not obvious, hence X and Y chromosomes presumably both telocentric. Habitat. Typically found in evergreen tropical forests, including both montane and mossy communities. Least Philippine Forest Mice occur also in secondary forest, and sometimes in shrubby and grassy habitats. Elevational range on Luzon is 300-2370 m, but most records are from above 1460 m; on Mindoro recorded at ¢.1370 m. Record from Dinagat is from 750 m. Food and Feeding. Reported as “omnivorous, feeding on seeds, insects and other invertebrates, and earthworms.” Breeding. In sample of females from Mount Isarog (S Luzon), estimated litter size from counts of embryos and uterine scars was 1-3 (average 2). Males in this population had small testes, measuring up to 5 mm x 11 mm. Activity patterns. Least Philippine Forest Mice are generally nocturnal, though occasionally active during day in rainy season. They spend most time in forest canopy, but sometimes descend to ground or close toit. Movements, Home range and Social organization. On Mount Isarog in 1993-1994, Least Philippine Forest Mice occurred at density of 8-8 ind/ha, the highest density for any small terrestrial mammal at the site. Average distance moved between captures was 42 m, maximum distances varying from 150 m to 226 m. Home-range size was estimated at 0-22 ha (range 0-01-0-56 ha). Status and Conservation. Classified as Least Concern on The IUCN RedList. Bibliography. Balete & Heaney (1997), Danielsen et al. (1994), Ellerman (1941), Heaney (2016d), Heaney, Balete, Dolar et al. (1998), Heaney, Balete, Gee et al. (2005), Heaney, Balete & Rickart (2016), Heaney, Tabaranza, Rickart et al. (2006), Hollister (1913), Justiniano et al. (2015), Miller (1910), Musser (1982a), Musser & Carleton (1993, 2005), Rickart et al. (1991)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 669, DOI: 10.5281/zenodo.6887260

201. Small Luzon Forest Mouse Apomys microdon French: Apomys de Catanduanes / German: Kleinzahn-Philippinenwaldmaus / Spanish: Raton de bosque pequefo de Luzon Other common names: Small Luzon Apomys Taxonomy. Apomys microdon Hollister, 1913, “Biga [= Viga], CatanduanesIsland,” Catanduanes Province, Bicol Region, Philippines. Apomys microdon, a member of subgenus Apomys, 1s most closely related to A. musculus. G. B. Corbet and |]. E. Hilllisted this species in 1992 as Apomys hollisteri, a name proposed by J. R. Ellerman within Rattus as a replacement for microdon on grounds that latter name was preoccupied by Mus microdon (a member of African genus Mastomys that Ellerman also placed within Rattus). As correctly assessed by G. G. Musser in 1982, removal of both of these names from Rattus dissociates them from each other and thus allows use of the combination Apomys microdon, with: Rattus (Apomys) hollisteri as a junior synonym. A. microdon is very likely a species complex,as yet incompletely sampled or understood across its geographic range. Musser in 1982 associated samples from Dinagat and Leyte with A. microdon, whereas E. A. Rickart and others referred samples from Biliran, Bohol, and Leyte to A. lttoralis, these populations probably represent one or more undescribed species, related to A. microdon. On Luzon, A. microdon is sympatric with various members of subgenus Megapomys which are more strictly terrestrial, contrasting with the scansorial to arboreal members of subgenus Apomys. Currently regarded as monotypic but in need offurther assessment. Distribution. Luzon and Catanduanes Is, Philippines. Descriptive notes. Head-body 90-107 mm, tail 124-149 mm, ear 15-19 mm, hindfoot 26-31 mm; weight 28-42 g. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet with sharp, recurved claws on all digits, thinly furred tail with weakly overlapping scales and three hairs per tail scale, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed, and active primarily in canopy. The Small Luzon Forest Mouse, one of smallest and more brightly colored members of subgenus, has fur on upperparts, flanks, and outer surfaces of limbs overall tawny brown and lightly flecked with black, guard hairs black and narrowly projecting through fur; fur on underparts and inside of limbs of variable coloration, some bright orange brown to bases, others buff but with pale to dark gray bases to hairs, occasional individuals have white patches on chest and midline of abdomen; upperside and underside with sharply defined boundary. Tail is considerably longer than head-body length, brown above to tip but variably brown, mottled, or pale brown to white below; scales small; upper surfaces of foreand hindfeet pale brown, sparsely haired. Habitat. Recorded fromprimary evergreen tropical rainforest, ranging from lowland rainforest and hill forest to montane forest; extends into mossy forest only when other species of subgenus Apomys are absent. Occurs also in patches of secondary forest, regenerating after natural or anthropogenic disturbance, but not outside forest cover. On Luzon, Small Luzon Forest Mice are found from sea level to 2025 m, but with lower elevational cut-off where A. musculus (also a member of subgenus Apomys) is present in the montane forests. Food and Feeding. Dietary focus of the Small Luzon Forest Mouseis said to be seeds, with occasional consumption of insects and earthworms. Breeding. Pregnant females are reported with 1-3 embryos (modally 2). Adult males reportedly have small testes, measuring 4 mm x 8 mm. Reports of nests constructed from “dry bamboo leaves, with a few leaves from broad-leafed trees,” and located in axils of Pandanus palms and in clumps of climbing bamboos. Activity patterns. The Small Luzon Forest Mouse is said to be active at night, most often in trees but sometimes on ground. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Small Luzon Forest Mice are recorded from numerous localities on Luzon and can be moderately common in suitable habitat. Lowland parts of this species’ range have suffered severe deforestation, but remaining habitats at higher elevations are not under any immediate threat. Bibliography. Corbet & Hill (1992), Danielsen et al. (1994), Ellerman (1941), Heaney (2016¢), Heaney, Balete, Dolar et al. (1998), Heaney Balete, Gee et al. (2005), Heaney, Balete & Rickart (2016), Hollister (1913), Justiniano etal. (2015), Musser (1982a), Musser & Carleton (1993, 2005), Rickart et al. (1991)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 668, DOI: 10.5281/zenodo.6887260

198. Mindanao Lowland Forest Mouse Apomys lttoralis French: Apomys cétier / German: Kiisten-Philippinenwaldmaus / Spanish: Raton de bosque de tierras bajas de Mindanao Other common names: Mindanao Lowland Apomys Taxonomy. Rattus (Apomys) littoralis Sanborn, 1952, “Bugasan, altitude 50 feet [= 15 m], Cotabato Province, Mindanao Island, Philippine Islands.” Apomys littoralis is a member of subgenus Apomys. Its relationship with A. hylocoetes, the type species of Apomys, was first recognized by D. H.Johnson in 1962. Monotypic. Distribution. Known with certainty only from Mindanao I (Lanao del Sur and Maguindanao provinces), Philippines; populations on Leyte, Biliran, and Bohol Is are tentatively referred to this species. A related population on Negros I is considered to represent an undescribed species. Descriptive notes. Head-body 100-103 mm, tail 122-126 mm, ear 16-17 mm, hindfoot 26-27 mm. No specific data are available for body weight. Species of Apomys are softfurred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomysare small, longtailed, and typically most active in canopy. The Mindanao Lowland Forest Mouse, a medium-sized member of this subgenus, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, without spinesor stiff hairs, tawny brown on back and head, paler on flanks; fur on underparts and inside of limbsis mostly gray with white tips, but with some patches of all-white fur on chest; ears relatively short. Upper surfaces offoreand hindfeet “light brown,” which suggests that dark fur of limbs extends onto extremities; hindfeet relatively long and narrow, as in other members of subgenus Apomys, plantar pads relatively small and well separated, digits moderately long and slender,all with sharply pointed claws. Tail is relatively long (120-125% of head-body length), grayish brown above and below for entire length, tail scales conspicuous, with short hairs and no terminal brush. Available crania are damaged, but do not show any exceptional features. A karyotype reported by E. A. Rickart and G. G. Musser in 1993 for a sample from Leyte Island (chromosomal complement 2n = 44, FN = 88) represents an undescribed species. Habitat. Capture localities indicate probable occurrence in evergreen tropical rainforest, including lowland rainforest and montane forest. Elevational range on Mindanao is 15 m to ¢.1400 m. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Lowland to mid-elevation rainforests on Mindanao Island have been severely impacted by forestry activities, and populations ofthis species are likely to be highly fragmented. Bibliography. Heaney et al. (1998), Johnson (1962), Kennerley (2016g), Musser (1982a), Musser & Carleton (2005), Musser & Heaney (1992), Rickart & Musser (1993), Rickart et al. (1993), Sanborn (1952a). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015). 198. Mindanao Lowland Forest Mouse Apomys lttoralis French: Apomys cétier / German: Kiisten-Philippinenwaldmaus / Spanish: Raton de bosque de tierras bajas de Mindanao Other common names: Mindanao Lowland Apomys Taxonomy. Rattus (Apomys) littoralis Sanborn, 1952, “Bugasan, altitude 50 feet [= 15 m], Cotabato Province, Mindanao Island, Philippine Islands.” Apomys littoralis is a member of subgenus Apomys. Its relationship with A. hylocoetes, the type species of Apomys, was first recognized by D. H.Johnson in 1962. Monotypic. Distribution. Known with certainty only from Mindanao I (Lanao del Sur and Maguindanao provinces), Philippines; populations on Leyte, Biliran, and Bohol Is are tentatively referred to this species. A related population on Negros I is considered to represent an undescribed species. Descriptive notes. Head-body 100-103 mm, tail 122-126 mm, ear 16-17 mm, hindfoot 26-27 mm. No specific data are available for body weight. Species of Apomys are softfurred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomysare small, longtailed, and typically most active in canopy. The Mindanao Lowland Forest Mouse, a medium-sized member of this subgenus, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, without spinesor stiff hairs, tawny brown on back and head, paler on flanks; fur on underparts and inside of limbsis mostly gray with white tips, but with some patches of all-white fur on chest; ears relatively short. Upper surfaces offoreand hindfeet “light brown,” which suggests that dark fur of limbs extends onto extremities; hindfeet relatively long and narrow, as in other members of subgenus Apomys, plantar pads relatively small and well separated, digits moderately long and slender,all with sharply pointed claws. Tail is relatively long (120-125% of head-body length), grayish brown above and below for entire length, tail scales conspicuous, with short hairs and no terminal brush. Available crania are damaged, but do not show any exceptional features. A karyotype reported by E. A. Rickart and G. G. Musser in 1993 for a sample from Leyte Island (chromosomal complement 2n = 44, FN = 88) represents an undescribed species. Habitat. Capture localities indicate probable occurrence in evergreen tropical rainforest, including lowland rainforest and montane forest. Elevational range on Mindanao is 15 m to ¢.1400 m. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Lowland to mid-elevation rainforests on Mindanao Island have been severely impacted by forestry activities, and populations ofthis species are likely to be highly fragmented. Bibliography. Heaney et al. (1998), Johnson (1962), Kennerley (2016g), Musser (1982a), Musser & Carleton (2005), Musser & Heaney (1992), Rickart & Musser (1993), Rickart et al. (1993), Sanborn (1952a). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 667-668

199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 667-668

205. Sierra Madre Forest Mouse Apomys sierrae French: Apomys de la Sierra Madre / German: Sierra-Madre-Philippinenwaldmaus / Spanish: Raton de bosque de Sierra Madre Other common names: North Sierra Madre Forest Mouse, Sierra Apomys Taxonomy. Apomys sierrae Heaney et al., 2011, “Philippines: Luzon [Island]: Cagayan Province: 3-5 km SW Mt. Cetaceo peak, 1400 m elevation, 17-69561°N, 122-01683°E.” Apomys sierrae 1s a member of subgenus Megapomys. It seems not to have any particularly close affinities, but nearest phylogenetic relatives may be A. magnus, A. aurorae, A. zambalensis, and A. iridensis. A. sierrae was tentatively reported as A. datae by F. Danielsen and colleagues in 1994; it has been newly collected over much of its geographic range only since the mid-2000s. Currently regarded as monotypic but in need of further assessment. Distribution. Luzon I (Sierra Madre, Mt Lataan, Mungiao Mts, and Mt Palali) and adjacent Palaui I, Philippines. Descriptive notes. Head—body 122-171 mm,tail 118-154 mm, ear 18-21 mm, hindfoot 31-40 mm; weight 64-111 g. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground; in most forest habitats across Luzon they are the most abundant terrestrial mammal in any given community. The Sierra Madre Forest Mouse is a medium-sized Megapomys with fur on upperparts variable, dark brown with rusty tints dorsally in Sierra Madre populations and medium brown with yellowish tints in Caraballo Mountains populations; in both populations, fur on underparts and inside of limbsis white or nearly white with orange wash, hairs medium to pale gray at bases; border between upperside and undersideis slightly diffuse; head has whitish-gray furon throat but cheek and sides of snout brown. Dark fur on foreand hindlimbs terminates at wrist and ankle, upper surfaces of foreand hindfeet mostly white with scattered dark hairs; hindfeet relatively long and narrow, undersurface heavily pigmented for most of length though not on plantar pads, hallucal pad long and narrow, other plantar pads relatively small, digits relatively long and narrow, and bearing sharp, curved claws. Tail is relatively long (95-100% of head-body length) and sharply bicolored, dark brown above and white below for all or most of length, sometimes with short white tip. Cranium is similar to those of other Megapomys but rostrum relatively slender, braincase globular, and molars relatively narrow. Habitat. Evergreen tropical rainforest, including lowland and montane rainforest types, and narrowly extending into “mossy forest.” At lower elevations, it occurs in lightly to heavily disturbed forest, as well as inprimary forest. Elevational range on Luzon 475-1700 m; recorded at 153 m on Palaui I. Food and Feeding. L.. R. Heaney and others stated in 2016 that Sierra Madre Forest Mice “showed a strong preference for earthworm bait rather than fried coconut on Mount Cetaceo, but a roughly equal preference on Mount Palali; they appear to be omnivorous overall.” Breeding. Seven pregnant females had average of 1-7 embryos (range 1-2). Activity patterns. According to Heaney and others, nearly all captures of Sierra Madre Forest Mouse were made at night, and nearly all on ground surface. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List but its wide distribution and elevational range, local abundance, and occurrence in regrowth habitat following heavy disturbance all attest to its resilience. Sierra Madre Forest Mice were the most commonly captured small ground mammal between 780 m and 1707 m on Mount Palali inthe Caraballo Mountains, and they were similarly common at 1300-1500 m on Mount Cetaceo in Cagayan Province. Bibliography. Alviola et al. (2011), Danielsen et al. (1994), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 670, DOI: 10.5281/zenodo.6887260

198. Mindanao Lowland Forest Mouse Apomys lttoralis French: Apomys cétier / German: Kiisten-Philippinenwaldmaus / Spanish: Raton de bosque de tierras bajas de Mindanao Other common names: Mindanao Lowland Apomys Taxonomy. Rattus (Apomys) littoralis Sanborn, 1952, “Bugasan, altitude 50 feet [= 15 m], Cotabato Province, Mindanao Island, Philippine Islands.” Apomys littoralis is a member of subgenus Apomys. Its relationship with A. hylocoetes, the type species of Apomys, was first recognized by D. H.Johnson in 1962. Monotypic. Distribution. Known with certainty only from Mindanao I (Lanao del Sur and Maguindanao provinces), Philippines; populations on Leyte, Biliran, and Bohol Is are tentatively referred to this species. A related population on Negros I is considered to represent an undescribed species. Descriptive notes. Head-body 100-103 mm, tail 122-126 mm, ear 16-17 mm, hindfoot 26-27 mm. No specific data are available for body weight. Species of Apomys are softfurred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomysare small, longtailed, and typically most active in canopy. The Mindanao Lowland Forest Mouse, a medium-sized member of this subgenus, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, without spinesor stiff hairs, tawny brown on back and head, paler on flanks; fur on underparts and inside of limbsis mostly gray with white tips, but with some patches of all-white fur on chest; ears relatively short. Upper surfaces offoreand hindfeet “light brown,” which suggests that dark fur of limbs extends onto extremities; hindfeet relatively long and narrow, as in other members of subgenus Apomys, plantar pads relatively small and well separated, digits moderately long and slender,all with sharply pointed claws. Tail is relatively long (120-125% of head-body length), grayish brown above and below for entire length, tail scales conspicuous, with short hairs and no terminal brush. Available crania are damaged, but do not show any exceptional features. A karyotype reported by E. A. Rickart and G. G. Musser in 1993 for a sample from Leyte Island (chromosomal complement 2n = 44, FN = 88) represents an undescribed species. Habitat. Capture localities indicate probable occurrence in evergreen tropical rainforest, including lowland rainforest and montane forest. Elevational range on Mindanao is 15 m to ¢.1400 m. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Lowland to mid-elevation rainforests on Mindanao Island have been severely impacted by forestry activities, and populations ofthis species are likely to be highly fragmented. Bibliography. Heaney et al. (1998), Johnson (1962), Kennerley (2016g), Musser (1982a), Musser & Carleton (2005), Musser & Heaney (1992), Rickart & Musser (1993), Rickart et al. (1993), Sanborn (1952a). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015). 198. Mindanao Lowland Forest Mouse Apomys lttoralis French: Apomys cétier / German: Kiisten-Philippinenwaldmaus / Spanish: Raton de bosque de tierras bajas de Mindanao Other common names: Mindanao Lowland Apomys Taxonomy. Rattus (Apomys) littoralis Sanborn, 1952, “Bugasan, altitude 50 feet [= 15 m], Cotabato Province, Mindanao Island, Philippine Islands.” Apomys littoralis is a member of subgenus Apomys. Its relationship with A. hylocoetes, the type species of Apomys, was first recognized by D. H.Johnson in 1962. Monotypic. Distribution. Known with certainty only from Mindanao I (Lanao del Sur and Maguindanao provinces), Philippines; populations on Leyte, Biliran, and Bohol Is are tentatively referred to this species. A related population on Negros I is considered to represent an undescribed species. Descriptive notes. Head-body 100-103 mm, tail 122-126 mm, ear 16-17 mm, hindfoot 26-27 mm. No specific data are available for body weight. Species of Apomys are softfurred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomysare small, longtailed, and typically most active in canopy. The Mindanao Lowland Forest Mouse, a medium-sized member of this subgenus, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, without spinesor stiff hairs, tawny brown on back and head, paler on flanks; fur on underparts and inside of limbsis mostly gray with white tips, but with some patches of all-white fur on chest; ears relatively short. Upper surfaces offoreand hindfeet “light brown,” which suggests that dark fur of limbs extends onto extremities; hindfeet relatively long and narrow, as in other members of subgenus Apomys, plantar pads relatively small and well separated, digits moderately long and slender,all with sharply pointed claws. Tail is relatively long (120-125% of head-body length), grayish brown above and below for entire length, tail scales conspicuous, with short hairs and no terminal brush. Available crania are damaged, but do not show any exceptional features. A karyotype reported by E. A. Rickart and G. G. Musser in 1993 for a sample from Leyte Island (chromosomal complement 2n = 44, FN = 88) represents an undescribed species. Habitat. Capture localities indicate probable occurrence in evergreen tropical rainforest, including lowland rainforest and montane forest. Elevational range on Mindanao is 15 m to ¢.1400 m. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Lowland to mid-elevation rainforests on Mindanao Island have been severely impacted by forestry activities, and populations ofthis species are likely to be highly fragmented. Bibliography. Heaney et al. (1998), Johnson (1962), Kennerley (2016g), Musser (1982a), Musser & Carleton (2005), Musser & Heaney (1992), Rickart & Musser (1993), Rickart et al. (1993), Sanborn (1952a). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015). 199. Lubang Forest Mouse Apomys lubangensis French: Apomys de Lubang / German: Lubang-Philippinenwaldmaus / Spanish: Raton de bosque de Lubang Other common names: Lubang Apomys Taxonomy. Apomys lubangensis Heaney el al., 2014, “Philippine Islands: Lubang Is- land, Mindoro Occidental Province: L.ooc Municipality: Barangay Burol: Sitio Bubuy- og, 13°48’N, 120°7-8’E.” Apomys lubangensis, newly collected in 2001, is a member of subgenus Megapomys, and may be most closely related to A. sacobianus, A. banahao, and A. brownorum. On Lubang Island, A. lubangensis was sympatric with a small, as yet unidentified arboreal species of Apomys at 300-425 m. Distribution. Endemic to Lubang I, Philippines, recorded at elevations of 130-425 m on Mt Ambulong. Descriptive notes. Head—body 143-160 mm, tail 128-154 mm, ear 22-27 mm, hindfoot 38-42 mm; weight 102-128 g. Males average slightly larger than females in all external measurements; no chromatic sexual dimorphism. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Lubang Forest Mouse, one of largest Megapomys, has fur on upperparts, flanks, and outer surfaces of limbs dense and very soft, brownish gray with reddish wash, and a “variegated” appearance probably caused by slightly shaggy texture; fur on underparts and inside of limbs grayish white; boundary between colors of upperside and underside abrupt; earslarge, thinly furred. Dark fur of limbs does not extend onto upper surfaces of foreand hindfeet, which are unpigmented and clothed in white hairs; hindfeet relatively elongate, undersurface with dark gray pigmentation except on plantar pads. Tail is relatively long (c.88-95% of head-body length) compared with other Megapomys, sharply bicolored, nearly black above and white below for entire length, tail scales large and prominent, tail hairs relatively short, no terminal brush. Cranium is robust, with broad interorbital region and relatively short, deep, broad rostrum. Habitat. Natural habitat is evergreen tropical rainforest. Lubang Forest Mice have been recorded in “secondary lowland forest with few mature trees on steep terrain” at 130 m, in “mixed second-growth lowland forest” at ¢.180 m, and in “well-developed, regenerating forest” at 300 m and 450 m. These records indicate a degree of tolerance of habitat disturbance, but this is unlikely to extend to occupation offully converted land. Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species rest on the extent of future forestry activity on Lubang Island. Bibliography. Balete, Heaney & Rickart (2013), Heaney, Balete, Rickart et al. (2011), Heaney, Balete, Veluz et al. (2014), Justiniano et al. (2015).

191. Brown's Forest Mouse Apomys brownorum French: Apomys des Brown / German: Brown-Philippinenwaldmaus / Spanish: Raton de bosque de Brown Other common names: Brown's Apomys, Tapulao Apomys, Tapulao Forest Mouse Taxonomy. Apomys brownorum Heaney et al, 2011, “Philippines: Luzon Island: Zambales Province: Palauig Munic.: Brangay Salasa: Mt. Tapulao peak, 2024 m; 15°28’54-8”S, 120°07°10-4’E.” Apomys brownorum, newly collected in 2005, is a member of subgenus Megapomys and most closely related to A. banahao. It is not known to be sympatric with any other Megapomys species, but occurs with Apomys microdon and A. musculus of subgenus Apomys. Monotypic. Distribution. Recorded only from type locality, at 2024 m on Mt Tapulao, Zambales Mts, W Luzon, Philippines; it was not trapped at lower elevation (1690 m) on Mt Tapulao, where A. zambalensis was encountered instead, but it may occur on other, hitherto unsurveyed high peaks in the Zambales Mts. Descriptive notes. Head-body 123-140 mm, tail 107-116 mm, ear 21-22 mm, hindfoot 31-36 mm; weight 60-84 g. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species that are active primarily on ground; in most forest habitats across Luzon, they are the most abundant terrestrial mammal in any local community. Brown's Forest Mouseis one of the smaller species of Megapomys.Fur on upperparts, flanks, and outer surfaces of all limbs is dense and very soft, rich dark brown, sometimes with rusty-orange tints between eye and ear and below ear, body hairs dark gray basally, guard hairs black and narrowly projecting through fur; fur on underparts and inside of limbs dark gray at bases with ocherous-gray tips; coloration of upperside merges gradually with lower parts or is separated by band of rusty-orange fur; ears darkly pigmented. Dark fur of limbs extends onto upper surfaces of feet, but digits are unpigmented and covered with short white hairs; hindfeet relatively elongate but somewhat broader than those of other members of Megapomys, undersurface mostly pale but some with pale gray blotches proximally, most plantar pads small and well separated but hallucal pad long and quite broad, digits relatively long but stocky. Tail is relatively short (70-88% of head-body length), sharply bicolored, dark above and white below for entire length. Cranium with moderately long rostrum, relatively robust incisors, and relatively small molars for genus. Habitat. Known only from high-elevation evergreen tropical rainforest, called “mossy forest” in Philippine ecological literature. Brown’s Forest Mouseis recorded from primary forest and also from regenerating habitat after mining operation;it is the most abundant terrestrial mammal at these sites. Food and Feeding. I.. R. Heaney and coworkers reported in 2016 that these mice “showed a slight preference for fried coconut bait rather than live earthworms” and “are probably omnivorous.” Breeding. Heaney and colleagues reported that “two pregnant females each carried a single embryo.” Activity patterns. Heaney and coworkers reported that “Nearly all were captured at night, and nearly all were on the ground surface (not in trees).” Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The long-term prospects of this species may rest on whetherit occurs more widely at high elevation in the Zambales Mountains. Bibliography. Balete et al. (2009), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 664, DOI: 10.5281/zenodo.6887260

206. Zambales Forest Mouse Apomys zambalensis French: Apomys des Zambales / German: Zambales-Philippinenwaldmaus / Spanish: Raton de bosque de Zambales Other common names: Zambales Apomys Taxonomy. Apomys zambalensis Heaney et al., 2011, “Philippines: Luzon I[slan]d.: Bataan Province: 0-1 km N Mt. Natib peak, 1150 m, 14-71513°N,120-39892°E.” Newly collected in 2005, A. zambalensis is a member of subgenus Megapomys and its nearest phylogenetic relatives may be A. wridensis, A. aurorae, and A. magnus. It is sympatric in the Zambales Mountains with A. sacobianus, but is found only at lower elevations than A. brownorum on Mount Tapulao. Monotypic. Distribution. Restricted to the Zambales Mts of W Luzon, Philippines. Descriptive notes. Head-body 127-163 mm,tail 123-158 mm, ear 20-23 mm, hindfoot 35-40 mm; weight 67-112 g. Apomys species are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite offeatures, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears,vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground; in most forest habitats across Luzon they are the most abundant terrestrial mammal in any given community. The Zambales Forest Mouse is the second largest species of Megapomys and has fur on upperparts soft but relatively short and bright rusty orange, fur on underparts and inside of limbs pale to medium gray at bases and white with slight to heavy ocherous wash at tips; border between upperside and underside is slightly diffuse; head with whitish-gray fur on throat but cheek and sides of snout orange brown; ears are relatively large, thinly furred, and less heavily pigmented than those of other Megapomys. Dark fur on forelimb terminates well above wrist, followed by a band of pale orange fur and then bywhite fur on wrist and forefoot; dark fur on hindlimb terminates at ankle, hindfeet mostly white, sometimes with scattered dark hairs; hindfeet are relatively long and broad, undersurface pigmented over most of length including some plantar pads, hallucal pad relatively short and placed well forward, other plantar pads relatively large but well separated; hallux long, reaching almost to top of adjacent interdigital pad, other digits also relatively long and narrow,all digits bearing sharp, curved claws. Tail is relatively long (94-100% of head-body length) and sharply bicolored, dark brown above and white below for all or most of length. Cranium is similar to those of other Megapomys but rostrum relatively long and robust, braincase somewhat elongate and flattened. Chromosomal complement has 2n = 44, FN> 58, autosomes include two pairs of large submetacentric, two pairs of medium-sized metacentric or submetacentric, and 15 pairs of small to large telocentric or subtelocentric chromosomes. The X chromosome is a medium-sized submetacentric and the Y chromosome a small telocentric. Habitat. Evergreen tropical rainforest, including lowland and montane rainforest types, and narrowly extending into mossy forest, at elevations of 365-1690 m. The Zambales Forest Mouse occurs in lightly to heavily disturbed forest, as well as in primary forest, and on Mount Pinatubo was abundant—though less so than the Longnosed Luzon Forest Mouse (A. sacobianus)—in sparse regrowth vegetation after the violent eruption of 1991. It was the most commonly captured small ground mammal on Mount Tapulao from 925 m to 1690 m and on Mount Natib from 900 m to 1250 m. Food and Feeding. Zambales Forest Mice are reported as avidly consuming live earthworm bait and fried coconut and are “best considered to be omnivorous.” Breeding. A pregnant female carried two embryos. Activity patterns. The Zambales Forest Mouse is active almost exclusively at night and rarely, if ever, climbs above ground surface (and never more than 1 m above ground). Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List butits relatively wide distribution and elevational range, local abundance, and occurrence in regrowth habitat following heavy disturbance suggest a high degree ofresilience. Bibliography. Balete et al. (2009), Engelbrektsson & Kennerley (2017), Johnson (1962), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Heaney, Balete, Veluz et al. (2014), Musser (1982a), Musser & Carleton (1993, 2005)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 670, DOI: 10.5281/zenodo.6887260

194. Large Mindoro Forest Mouse Apomys gracilirostris French: Apomys de Mindoro / German: Schmalschnauzen-Philippinenwaldmaus / Spanish: Raton de bosque de Mindoro Other common names: Large Mindoro Apomys : Taxonomy. Apomys gracilirostris Ruedas, 2 1995, “Philippines: Mindoro Island; Mindoro Occidental Province; Municipality of San Teodoro, North Ridge approach to Mount Halcon, ca. 1580 m; ca. 13°16’48’N, 121 °B9°19°E.” Although currently placed in subgenus Megapomys, Apomys gracilirostris is morphologically divergent and appears to be the earliest derivative of this lineage. Monotypic. Distribution. Currently recorded only on N ridge of Mt Halcon, Mindoro I, Philippines; likely widespread in mountains of Mindoro, an area that remains poorly surveyed for small mammals. Descriptive notes. Head-body 137-147, tail 135-185 mm, ear 14-22 mm, hindfoot 33-45 mm; weight 71-140 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground. The Large Mindoro Forest Mouseis one of the bigger and more unusual species of Megapomys. Fur on upperparts is dark brown with fine black flecking, the latter produced by numerous short black guard hairs, body hairs with light gray bases; fur on underparts and inside of all limbs usually similar in tones to back but paler, some individuals with silvery tipping; colors of upper and lower parts blend without sharply defined boundary; ears darkly pigmented on interior surface, somewhat paler externally. Dark fur of limbs extends onto upper surfaces of foreand hindfeet, digits pigmented and with short dark hairs; hindfeet relatively short and broad compared with those of other Megapomys, undersurface with dark gray pigment apart from on pads, digits relatively short and stocky, plantar pads moderately large but well separated, first digit very long, reaching to front of first interdigital pad. Tail is relatively long (on average 105% of head-body length), dark above and only slightly paler below for entire length, some individuals with short white tip (2-10 mm), scales in 14 rows per cm near base and at middle oftail. Cranium is larger but quite gracile compared with most Megapomys, with long, slender rostrum, relatively small molars, and very narrow incisors (lowers described as lacking enamel). Habitat. The Large Mindoro Forest Mouse was abundant in 1992 in evergreen tropical montane rainforest on Mount Halcon. At elevations between 1250 m and ¢.1600 m the forest canopy was floristically diverse, with heavy growth of moss, and understory generally dense, with areas oftree ferns, climbing bamboos and palms including pandans (Pandanus, Pandanaceae). Above 1600 m, the canopy consisted entirely of the pine Agathis philippinensis, with dense understory of bamboos, pandans and gingers. Recorded at elevations of 1250-1950 m. Food and Feeding. I.. A. Ruedas speculated that it might feed on insects on account of its gracile dentition. Breeding. Ruedas reported in 1995 that “one female had three embryos;” largest scrotal testes measured 7 mm x 4 mm. Activity patterns. Ruedas offered reasonable speculation that this species was scansorial on basis of the form ofits feet and relative length oftail. Apparent absence of any member of subgenus Apomys on Mindoro may allow the Large Mindoro Forest Mouse a greater ecological breadth thanother members of the Megapomys lineage. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Least Concern on The IUCN Red List. Montane habitats on Mindoro Island are currentlystable, but the island has been subject to past episodes of deforestation. The status of this species has not been assessed in the field since its initial discovery. Bibliography. Heaney, Balete, Dolar et al. (1998), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Justiniano et al. (2015), Kennerley (2016f), Musser & Carleton (2005), Ruedas (1995)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 665-666

189. Aurora Forest Mouse Apomys aurorae French: Apomys dAurora / German: Aurora-Philippinenwaldmaus / Spanish: Raton de bosque de Aurora Other common names: Aurora Apomys Taxonomy. Apomys aurorae Heaney et al., 2011, “Philippines: Luzon Island: Aurora Province: Dingalan Munic.: 2 km S, 2 km W Mingan peak, 1305 m, 15-46456°N, 121-38421°L,” Newly collected in 2006, A. auroraeis a member of subgenus Megapomys and may be most closely related to A. iridensis and A. zambalensis. It is narrowly sympatric at upper end of its elevational range with A. minganensis (of subgenus Megapomys), and more broadly sympatric with each of A. microdon and A. musculus (of subgenus Apomys). Monotypic. Distribution. Currently known only from Mingan Mts of S Sierra Madre, E Luzon I, Philippines. Descriptive notes. Head-body 126-152 mm, tail 129-153 mm, ear 18-21 mm, hindfoot 33-37 mm; weight 58-92 g. Males average slightly larger than females in all measurements and in weight (81-2 g vs. 73-8 g). Species of Apomys are soft-furred, small to mediums-sized murines with relatively unspecialized body form; they are distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species that are active primarily on ground; in most forest habitats across Luzon, they are the most abundant terrestrial mammal in any local community. The Aurora Forest Mouse is one of the smaller species of Megapomys. Fur on upperparts, flanks, and outer surfaces of all limbs is rich rusty reddish brown with diffuse black flecking, body hairs medium gray basally, guard hairs black and narrowly projecting through fur; fur on underparts and inside of foreand hindlimbs pale gray basally with white tips, sometimes with pale ocherous wash; upper and lower parts with sharply defined boundary. Dark fur of limbs does not extend ontoupper surfaces of foreand hindfeet, which are unpigmented and covered with short white hairs, sometimeslightly peppered with black; hindfeet relatively elongate and narrow, undersurface with gray pigment for proximal half, often surrounding posterior pads, plantar pads small and well separated, first digit short but others relatively long and slender. Tail is relatively long (98-100% of head-body length), dark above and white below for entire length, but sometimes with scattered pigmented scales and dark hairs. Cranium has rounded braincase, moderately short and slender rostrum, and average-sized molars for genus. Habitat. Evergreen tropical rainforest, including primary lowland and montane rainforest and extending into ecotone of montane and “mossy” forest types. It is recorded also in lightly disturbed habitats across its elevational range of 733-1677 m, but is most abundant between 902 m and 1476 m. Food and Feeding. I.. R Heaney and coworkers reported in 2016 that Aurora Forest Mice are “omnivorous, consuming seeds, insects, and earthworms.” Breeding. Four pregnant females in 2016 had average of two embryos (range 1-3). Activity patterns. Aurora Forest Mice are nocturnal and forage on surface of ground. Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. Current known range is very small, but it may occur more widely in the Mingan Mountains. Bibliography. Balete et al. (2011), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Justiniano etal. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 663-664, DOI: 10.5281/zenodo.6887260

192. Camiguin Forest Mouse Apomys camiguinensis French: Apomys de Caminguin / German: Camiguin-Philippinenwaldmaus / Spanish: Ratén de bosque de Camiguin Other common names: Camiguin Apomys Taxonomy. Apomys camiguinensis Heaney & Tabaranza, 2006, “Barangay Kital-is, Sagay Municipality, 2 km N, 62 kw W Mahinog, 1000 m elevation, Camiguin Province, Camiguin Island. 9°9.5°N, 124°43.5’E.” A member of subgenus Apomys, A. camiguinensis was newly collected in 1994 and, prior to formal description, was referred to as“ Apomys sp. D.” Molecular evidence suggests possible special affinity with A. Aylocoetes and A. insignis from Mindanao Island, and with an undescribed species of Apomys from Leyte and Biliran. Monotypic. Distribution. Camiguin I, S Philippines. Descriptive notes. Head-body 106 mm, tail 140-160 mm, ear 18-20 mm, hindfoot 31-34 mm; weight 33-48-5 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond,relatively large and thinly furred ears, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Apomys are small, long-tailed species active primarily in canopy. The Camiguin Forest Mouse is a medium-sized species of subgenus Apomys. Fur on upperparts, flanks, and outer surfaces of limbs is dense and very soft, without spines or stiff hairs, rich brownish russet with small amount of “salt-andpepper” speckling; fur on underparts and inside of limbs is usually nearly white with a buffy or pale russet wash, sometimes with pure white blaze on chest; colors of upperside and lower parts merge gradually; ears are darkly pigmented and with short hairs on outer surface; eyes ringed with thinly furred pale skin. Dark fur of limbs extends onto upper surfaces of foreand hindfeet, but digits are unpigmented and covered with shorter white hairs; hindfeet relatively elongate but somewhat broader than those of some other members of subgenus Apomys, undersurface lightly pigmented, plantar pads small and well separated, hallucal pad short but well forward, digits moderately long. Tail is relatively long (averaging slightly longer than head-body length), sharply bicolored, dark brown above and white below for entire length, tail scales conspicuous and relatively large, in 12-125 rows per cm near tail base, and usually with three short hairs per scale, hairiness increasing toward tail tip but without terminal brush. Cranium very similar to those of other members of subgenus Apomys, but braincase and orbit unusually long, and rostrum long but robust. Habitat. Known only from evergreen tropical rainforest on Mount Timpoong,at elevations of ¢.1000-1400 m. It was common in 1994-1995 in disturbed lowland rainforest at 1000 m and in undisturbed montane forest at 1275 m, and less so in primary mossy forest at 1200-1400 m. It is likely to occur on other forested peaks on Camiguin Island, but is probably absent from extensively deforested areas below 1000 m. Food and Feeding. No information. Breeding. A pregnant female captured in May 1994 had a single embryo, while two other adult females captured in May 1994 and March 1995 each had two placental scars. Adult females with enlarged teats weighed 37-5-42 g, whereas nulliparous females weighed up to 35 g. Adult males with scrotal testes weighed 36-5-48 g, whereas immature males with abdominal testes weighed 34 g or less. Activity patterns. All individuals captured in 1994-1995 were trapped on the ground among roots or beneath fallen and rotten logs, and at night. The Camiguin Forest Mouse is likely to be scansorial or arboreal, as are other species of subgenus Apomys. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Vulnerable on The IUCN Red List. The Camiguin Forest Mouse was locally abundant in 1994-1995 in both primary and disturbed habitats but its current status is unknown. Its known population is within Timboong Hibok-Hibok Natural Monument. Bibliography. Heaney (2017), Heaney & Tabaranza (1997 2006a, 2006b), Heaney et al. (2011), Heaney, Tabaranza, Balete & Rigertas (2006), Justiniano et al. (2015), Musser & Carleton (2005)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 664-665

202. Mount Mingan Forest Mouse Apomys minganensis French: Apomys des Mingan / German: Mount-Mingan-Philippinenwaldmaus / Spanish: Ratén de bosque de Mingan Other common names: Mount Mingan Apomys Taxonomy. Apomys minganensis Heaney et al, 2011, “Philippines: Luzon [Island]: Aurora Province: Dingalan Munic.: 1-5 km S, 0-5 km W Mingan peak, 1,681 m elevation, 15-46802°N, 121-40039°E.” Apomys minganensis, newly collected in 2006, is a member of subgenus Megapomys, within which it occupies an isolated position but may be most closely related to A. sierrae, A. magnus, A. zambalensis, A. iridensis, and A. aurorae. At lower end ofits elevational range A. minganensis is sympatric with A. aurorae. Monotypic. Distribution. Known only from the Mingan Mts, E LuzonI, Philippines, at elevations of 1540-1785 m; it may be present also in adjacent S Sierra Madre and the poorly surveyed Dingalan Mts. Descriptive notes. Head-body 246-279 mm, tail 116-138 mm, ear 18-19 mm, hindfoot 31-35 mm; weight 66-92 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite offeatures, includinglong, narrow hindfeet with sharp, recurved claws on all digits, thinly furred tail with weakly overlapping scales and three hairs per tail scale, unreduced eyes,relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species active primarily on ground; in most forest habitats across Luzon, they are the most abundant terrestrial mammal in any local community. The Mount Mingan Forest Mouse, a relatively small Megapomys species, has fur on upperparts “dark brown with rusty-orange tips” over dark gray bases, fur on underparts and inside of limbs dark gray at base with paler tips that usually have ocherous wash, upperside and underside with abrupt boundary. Dark fur on foreand hindlimbs extends onto ankle and wrist; upper surfaces of foreand hindfeetis dark brown, with pigmented skin and brown hairs even on digits; hindfeet relatively short and broad compared with those of other species of Megapomys, undersurface darkly pigmented at rear and surrounding but not including posterior pads, post-hallucal pad moderately elongate, other pads moderately large but well separated, digits relatively short and robust. Tail is moderately long (87-101%, average 94%, of head-body length), dark brown above and pure white or with scattered dark scales and hairs below for entire length. Cranium is similar to those of other Megapomys but molars unusually small and narrow, and molar rows parallel rather than diverging posteriorly. Habitat. Restricted to evergreen tropical forest at elevations of 1540-1785 m. Itis the most abundant small ground mammal in the “mossy” forest type on Mount Mingan, but is somewhat less abundant in transitional montane—mossy forest at ¢.1540 m. Above 1680 m it was the only Megapomys species captured, and it likely extends to summit of Mount Mingan at 1910 m. The mossy forest on Mount Mingan is naturally disturbed by typhoons and features abundant climbing pandan palms (Pandanus, Pandanaceae) and bamboos. Food and Feeding. L.. R. Heaney and coworkers reported in 2016 that Mount Mingan Forest Mice “show a strong preference for live earthwormbait rather than fried coconut and probably are at least somewhat omnivorous.” Breeding. Heaney and colleagues reported that “four adultfemales carried one embryo each, and one hadthree.” Activity patterns. Heaney and coworkers reported that “nearly all were captured at night, and all were captured on the ground surface (not in trees).” Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. The Mount Mingan range is not contained within a national park or protected area. Bibliography. Balete et al. (2011), Engelbrektsson (2017), Heaney, Balete & Rickart (2016), Heaney, Balete, Rickart, Alviola et al. (2011), Justiniano et al. (2015), Musser (1982a), Musser & Carleton (2005)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 668-669, DOI: 10.5281/zenodo.6887260

200. Large Forest Mouse Apomys magnus French: Grand Apomys / German: GroRe Philippinenwaldmaus / Spanish: Raton de bosque grande Other common names: Large Apomys, Large Banahaw Forest Mouse Taxonomy. Apomys magnus Heaney et al., 2011, “Philippines: Luzon Island: Quezon Province: Tayabas Munic.: Barangay Lalo: Mt. Banahaw, Hasaan, 1250 m; 14°3’44”N, 121°31'S"E.” Apomys magnus, newly collected in 2005,is a member of subgenus Megapomys. Genetic evidence indicates that it is well differentiated from all other known Megapomys; its closest relatives may be A. awrorae and A. zambalensis. It is narrowly sympatric with A. banahao at ¢.1465 m on Mount Banahaw. Monotypic. Distribution. Recorded only from Mt Banahaw, Luzon I, Philippines; possibly present on adjacent Mt San Cristobal. Descriptive notes. Head-body 137-160 mm, tail 133-154 mm, ear 21-23 mm, hindfoot 37-41 mm; weight 92-128 g. Species of Apomys are soft-furred, small to medium-sized murines with relatively unspecialized body form; distinguished from other Philippine murines by a distinctive suite of features, including long, narrow hindfeet, thinly furred tail, unreduced eyes, relatively large and thinly furred ears, vibrissae on snout very elongate and, folded back, reach to shoulder or beyond, reduction to two mammae on each side, both inguinal, and a simplified molar pattern with cusps united into transverse laminae. Members of subgenus Megapomys are larger, shorter-tailed species that are active primarily on ground; in most forest habitats across Luzon Island, they are the most abundant terrestrial mammal in any local community. The Large Forest Mouse is the largest Megapomys, with fur on upperparts moderately long and dark brown with conspicuous black tipping, further accentuated by short black guard hairs, body hairs with gray bases; fur on underparts and inside of limbs medium gray at base with white tipping; upperside and underside with sharply defined boundary. Dark fur of forelimbs extends forward as narrowing strip to forefeet, but digits are unpigmented and covered with short white hairs; dark fur on forelimb stops above ankle, hindfeet unpigmented and entirely covered with short white hairs; hindfeet relatively long and stout, undersurface darkly pigmented except on some pads, post-hallucal pad elongate, other pads moderately large but well separated, digits relatively long and robust. Tail is moderately long (averaging 96-99% of head-body length), dark brown above and unpigmented or white below for entire length. Cranium is larger than that of all other Megapomys, with robust, flaring zygomatic arches, long,stout rostrum, and long molar rows. Habitat. Abundant in evergreen tropical montane forests, including both lowland and montane types distinguished in Philippine ecological literature, at elevations of 765— 1465 m. Not captured at or below 600 m or above 1465 m, despite intensive trapping in these elevational zones Food and Feeding. [.. R. Heaney and colleagues reported in 2016 that Large Forest Mice “showed a weak preference for live earthworms rather than fried coconut but probably are best considered omnivorous.” Breeding. One pregnant female carried three embryos. No other information. Activity patterns. Heaney and coworkers reported that “all were captured at night, and all were captured on the ground surface (not in trees).” Movements, Home range and Social organization. No information. Status and Conservation. Not assessed on The IUCN Red List. Most of the known distribution ofthis species lies with the Mounts Banahaw-San Cristobal Protected Landscape. The Large Forest Mouse is the most commonly trapped small mammal in oldgrowth forest or mature regrowth at mid-elevations on Mount Banahaw. In heavily disturbed forest, at same elevations,it is less abundant than the Philippine Forest Rat (Rattus everetti). Bibliography. Heaney, Balete, Rickart et al. (2011), Heaney, Balete & Rickart (2016), Heaney, Balete, Rosell-Ambal et al. (2013), Justiniano et al. (2015)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 668, DOI: 10.5281/zenodo.6887260

Aim Our aims were to document the extent of diversification in an endemic clade of small mammals (Apomys, subgenus Megapomys) on a large oceanic island and to test whether speciation occurred primarily (1) along isolated elevational gradients or (2) among montane areas, as would be expected if diversification were driven by glacial cycles. Location The Philippine archipelago, particularly Luzon Island and two smaller adjacent islands (Mindoro and Lubang). Methods We analysed newly generated nuclear DNA sequences from five unlinked genes and mitochondrial cytochrome b using concatenation (likelihood and Bayesian) and coalescent-based methods to estimate the species tree for all 13 species. We tested a priori models of speciation using both topological constraints and reconstructed elevational ranges of ancestors. Results All individual gene trees recovered at least four of the morphologically defined Megapomys species as monophyletic, while the concatenated approaches recovered all 13 species as monophyletic. Each species was confined to a single mountain range or off-shore island. Most mountain ranges had two species, but these species pairs usually were not sisters. Megapomys originated at medium to high elevation followed by three transitions into lower elevations and possibly one to high elevation. Both a priori models of speciation (elevational gradient and glacial cycle) were rejected by topology tests. The speciation rate was approximately constant through time. Main conclusions Nuclear sequence data strongly corroborated the species status of recently described taxa. A well-supported phylogeny showed that Megapomys diversified by splitting into a predominantly high-elevation clade and an entirely low-elevation clade. Neither adaptation along elevational gradients on single mountain ranges nor vicariance of high-elevation species following glacial cycle-induced dispersals fitted the data. Rather, the most likely process explaining species distributions is repeated colonization of isolated mountain ranges by distantly related species.

We describe two new species of mice of the endemic Philippine genus Apomys, subgenus Megapomys. One is from Lubang Island, a small oceanic island off the southwest coast of Luzon, and the other is from Mt. Irid, a peak in the Southern Sierra Madre that lies northeast of Manila on Luzon Island. We also report the first specimens of Apomys sacobianus to be seen since the holotype was captured in 1956 at the foot of Mt. Pinatubo; our specimens were obtained subsequent to the eruption of Mt. Pinatubo in 1991, demonstrating the continued existence of this species. Speciation within Apomys (Megapomys) has contributed substantially to the high level of mammalian diversity and fine-scaled endemism observed on Luzon and adjacent Philippine islands.

The high diversity of native Philippine murid rodents includes an old endemic group, the chrotomyines, which are the sister group of the Australasian hydromyines. Herein we detail their interspecific diversity of relative testes mass (RTM) and sperm morphology. We find that in chrotomyines, as in the Australasian hydromyines, testes mass relative to body mass differs by an order of magnitude across the species and ranges from a large RTM in Soricomys and Chrotomys species to a small RTM in Apomys. Sperm morphology is associated with these findings, with individuals in species of Soricomys and Chrotomys producing relatively larger spermatozoa with a prominent apical hook and long tail, whereas, by contrast, the Apomys species have a sperm head that either has a very short or no apical hook and a shorter tail. These findings indicate coevolution of RTM with sperm morphological traits across the species, with the marked interspecific differences in RTM suggesting differences in the intensity of intermale sperm competition and hence breeding system. Thus, we hypothesise that species of Soricomys and Chrotomys that produce more streamlined spermatozoa with longer tails have a polyandrous or promiscuous mating system, whereas the Apomys species, which produce smaller and less streamlined spermatozoa, may exhibit monogamy.

Excavations in Callao Cave, in the lowland (ca. 85 m elevation) Cagayan River Valley of northeastern Luzon, Philippines, have produced the first fossils of any endemic genera of Philippine murid rodents. Three dentaries dated to the Late Pleistocene, between ca. 50,000 and 68,000 BP, are referred to the genera Batomys and Apomys; the former is a member of the endemic “Phloeomys Division” of Philippine murids, and the latter of the “Chrotomys Division,” also endemic to the Philippines. Batomys is currently known from five extant species from Luzon, Mindanao, and Dinagat islands; the two species known from Luzon differ in size and dental and mandibular morphology from the two fossil mandibles, and both occur only at elevations above 1350 m. Apomys is currently known from two subgenera on Luzon; the fossil is a member of the nominate subgenus, which contains two species on Luzon, one of which, Apomys microdon, is conspecific with one fossil. We hypothesize that the Batomys fossils represent a differ...

We surveyed non-volant mammals in montane (1300 and 1400 m) and mossy forest (1500 and 1550 m) on Mount Cetaceo in the northern Sierra Madre of northeast Luzon in 2004 and 2005. We recorded a total of 12 species of mammals, including one shrew (Soricidae), seven murid rodents (Muridae), one deer (Cervidae), one wild pig (Suidae), one macaque (Cercopithecidae), and one civet (Viverridae). In this paper, we present the first ecological data on two recently discovered species endemic to northeast Luzon, Archboldomys musseri and Apomys sierrae. We captured only three species at 1300 m; captured five species at 1400 m; captured or inferred six species at 1500 m; and documented four species at 1550 m. Combined with an earlier study at 960 m (3 species), these results imply that species richness might increase from the lowlands to ca. 1500 m. The small number of sampling localities prevented strong statistical inference, but percent trap success with roasted coconut bait declined with increased elevatio...

In 2005 and 2007, we conducted surveys of mammals along an elevational transect on Mount Palali (peak 1707 m) in the Caraballo Mountains, a poorly known mountain range on Luzon Island, Philippines. The surveys covered eight localities representing habitats from lowland agroforest and regenerating disturbed lowland rainforest at 780 m to mossy forest near the peak. We recorded 24 species, including one native shrew, one non-native shrew, five fruit bats, seven insectivorous bats, one monkey, six native rodents, two civets, and one pig. One species of Apomys is the newly described A. sierrae, and two species of Chrotomys are potentially undescribed species. Elevational patterns varied among mammals: bats were most diverse in the lowlands, native nonvolant small mammals had almost equal richness along the entire elevational transect, and most species of large mammals were present at all elevations. Bait attractiveness and diel activity pattern differed among native nonvolant small mammals: Apomys mi...

Surveys of small mammals on carefully selected mountains and mountain ranges on Luzon Island, Philippines, since 2000 have led to the discovery of seven previously unknown species of forest mice, Apomys, a remarkable radiation on just a portion of one island. On the basis of morphological and cytochrome (cyt) b DNA sequence data presented here, we propose a new subgenus, Megapomys, to include the large-bodied members of the genus, which form a monophyletic unit of relatively large mice (averaging ca. 65–110 g) with tails about as long as or slightly shorter than the length of the head and body; all of these species forage on the ground. Other members of the genus are assigned to the subgenus Apomys; they are smaller (ca. 18–41 g), have long tails, and usually or often forage above the ground surface. Members of the subgenus Megapomys include four previously recognized species (A. abrae, A. datae, A. gracilirostris, and A. sacobianus) and the seven new species described here (A. aurorae, A. banaha...

In 2004 and 2005, we conducted a survey of the small mammals on Mt. Tapulao (=Mt. High Peak, 2037 m) in the Zambales Mountains, Luzon Island, Philippines in order to obtain the first information on the mammals of this newly discovered center of endemism. We also tested two hypotheses regarding the relationship of species richness with elevation and the impact of alien species on native mammals. The survey covered five localities representing habitats from regenerating lowland rain forest at 860 m to mossy rain forest near the peak at 2024 m. We recorded 11 species, including 1 native shrew, 1 alien shrew, 8 native rodents, and 1 alien rodent. Two species of Apomys and one species of Rhynchomys are endemic to Zambales; this establishes the Zambales Mountains as a significant center of mammalian endemism. Species richness of native small mammals increased with elevation, from five species in the lowlands at 925 m to seven species in mossy forest at 2024 m; total relative abundance of native small mammals increased from 925 to 1690 m, then declined at 2024 m. Alien small mammals were restricted to highly disturbed areas. Our results support the prediction that maximum species richness of small mammals would occur in lower mossy forest near the peak, not near the center of the gradient. Our results also support the hypothesis that when a diverse community of native Philippine small mammals is present in either old-growth or disturbed forest habitat, “invasive” alien species are unable to penetrate and maintain significant populations in forest.

Breed, W.G. and Leigh, C.M. 2010. The spermatozoon of the Old Endemic Australo-Papuan and Philippine rodents – its morphological diversity and evolution.—Acta Zoologica (Stockholm) 91: 279–294 The spermatozoon of most murine rodents contains a head in which there is a characteristic apical hook, whereas most old endemic Australian murines, which are part of a broader group of species that also occur in New Guinea and the Philippines, have a far more complex sperm form with two additional ventral processes. Here we ask the question: what is the sperm morphology of the New Guinea and Philippines species and what are the trends in evolutionary changes of sperm form within this group? The results show that, within New Guinea, most species have a highly complex sperm morphology like the Australian rodents, but within the Pogonomys Division some species have a simpler sperm morphology with no ventral processes. Amongst the Philippines species, many have a sperm head with a single apical hook, but in three Apomys species the sperm head contains two additional small ventral processes, with two others having cockle-shaped sperm heads. When these findings are plotted on a molecular phylogeny, the results suggest that independent and convergent evolution of highly complex sperm heads containing two ventral processes has evolved in several separate lineages. These accessory structures may support the sperm head apical hook during egg coat penetration.

We analysed the phylogenetic relationships of ten of the 13 known species of the genus Apomys using DNA sequences from cytochrome b. Apomys, endemic to oceanic portions of the Philippine archipelago, diversified during the Pliocene as these oceanic islands arose de novo. Several of the speciation events probably took place on Luzon or Mindanao, the two largest, oldest, and most topographically complex islands. Only one speciation event is associated with vicar- iance due to Pleistocene sea-level fluctuation, and a Pleistocene diversification model in which isolation is driven by sea-level changes is inconsistent with the data. Tectonic vicariance is nearly absent from the Philippines, in which tectonic coalescence plays a significant role. Most speciation events (about two-thirds) are associated with dispersal to newly developed oceanic islands. The data imply that the species have persisted for long periods, measured in mil- lions of years after their origins; further implications therefore are that faunal turnover is very slow, and persistence over geological time spans is more prominent than repeated colonization and extinction. Neither the equilibrium nor the vicariance model of biogeography adequately encompasses these results; a model incorporating colonization, extinction, and speciation is necessary and must incorporate long-term persistence to accommodate our observations. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80, 699-715. ADDITIONAL KEYWORDS: biogeographical models - cytochrome b - dispersal - island biogeography - Pleistocene - speciation - vicariance.

Small mammals were surveyed along an elevational transect on Mount Isarog (summit at 1,966 m) in southeastern Luzon, Philippines. Removal trapping was conducted at six sites ranging from remnant dipterocarp forest (475 m) to mossy forest (1,750 m). A total of 255 small mammals was captured in 9,001 trap nights. Species captured were one shrew ( Crocidura grayi ) and seven rodents ( Apomys musculus, Apomys cf. microdon, Archboldomys luzonensis, Batomys sp., Chrotomys gonzalesi, Rattus everetti , and Rhynchomys isarogensis ). Five species are nocturnal, one diurnal, and two have variable diel activity. On the basis of trap success with two kinds of bait, analysis of stomach contents, and captive feeding trials, we recognize four trophic groups: omnivorous (three species), insectivorous (two species), granivorous-frugivorous (one species), and vermivorous (two species). Species richness and relative abundance followed predicted patterns of increase with elevation. Four species endemic to Mount Isarog were found only at high elevations.

Biodiversity surveys in the 1960s and 1990s on Camiguin Island, a geologically young, volcanically active oceanic island surrounded by deep water, have demonstrated the presence of 24 species of land mammals. Three species (one insectivore and two rodents) are not native to the Philippines, but all others (one insectivore, 12 bats, one monkey, four rodents, two small carnivores, and one ungulate) are indigenous. Among those captured in the 1990s were two previously unknown species of murid rodents in the genera Apomys and Bullimus that are endemic to Camiguin. The discovery of two new species on such a small island (ca. 265 km2) is remarkable; Camiguin is currently the smallest island in the Philippines known to have unique species of mammals. Total species richness of nonvolant mammals on Camiguin is low, but relative to islands that were once part of Pleistocene Greater Mindanao, Camiguin is not depauperate. However, its fauna is not ecologically balanced in the same way as the faunas of the is...

Apomys abrae (Sanborn, 1952). Fieldiana Zool., 33(2):133. TYPE LOCALITY: Philippines, Luzon Isl, Abra Prov, Abra, 3500 ft. DISTRIBUTION: Known only from middle and high elevations on Luzon. COMMENTS: Included within the " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098

Apomys hylocoetes Mearns, 1905. Proc. U.S. Natl. Mus., 28:456. TYPE LOCALITY: Philippines, Mindanao Isl, Mt Apo, 6000 ft. DISTRIBUTION: Known only from 6000-7600 ft on Mindanao. SYNONYMS: petraeus. COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098

Apomys musculus Miller, 1911. Proc. U.S. Natl. Mus., 38:403. TYPE LOCALITY: Philippines, Luzon Isl, Benguet, Baguio, Camp John Hay, 5000 ft. DISTRIBUTION: Luzon and Mindoro, Philippines. COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098

Apomys datae (Meyer, 1899). Abh. Mus. Dresden, ser. 7, 7:25. TYPE LOCALITY: Philippines, N Luzon Isl, Lepanto. DISTRIBUTION: Known only from W highlands in N Luzon. SYNONYMS: major. COMMENTS: The sole member of " Apomys datae Group " (Musser, 1982b)., 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 575, DOI: 10.5281/zenodo.7353098

Apomys littoralis (Sanborn, 1952). Fieldiana Zool., 33(2):134. TYPE LOCALITY: Philippines, Mindanao Isl, Bugasan, Cotabato, 50 ft. DISTRIBUTION: Formerly recorded from Negros and Mindanao (Musser, 1982b), but revised view excludes it from Negros, and records it from lowlands of Mindanao, Bohol, Biliran, Dinagat, and Leyte (Musser and Heaney, 1992). COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098, {"references":["Musser, G. G., and L. R. Heaney. 1992. Philippine rodents: Definitions of Tarsomys and Limnomys plus a preliminary assessment of phylogenetic patterns among native Philippine murines (Murinae, Muridae). Bulletin of the American Museum of Natural History, 211: 1 - 138."]}

Apomys sacobianus Johnson, 1962. Proc. Biol. Soc. Washington, 75:318. TYPE LOCALITY: Philippines, Luzon Isl, Pampanga Prov, Sacobia River, Clark Air Base. DISTRIBUTION: Known only from the type locality. COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098

Apomys insignis Mearns, 1905. Proc. U.S. Natl. Mus., 28:459. TYPE LOCALITY: Philippines, S Mindanao Isl, Mt Apo, 6000 ft. DISTRIBUTION: Known only from middle to high elevations on Mindanao (Musser and Heaney, 1992). SYNONYMS: bardus. COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098, {"references":["Musser, G. G., and L. R. Heaney. 1992. Philippine rodents: Definitions of Tarsomys and Limnomys plus a preliminary assessment of phylogenetic patterns among native Philippine murines (Murinae, Muridae). Bulletin of the American Museum of Natural History, 211: 1 - 138."]}

Apomys microdon Hollister, 1913. Proc. U.S. Natl. Mus., 46:327. TYPE LOCALITY: Philippines, Cataduanes Isl, Biga. DISTRIBUTION: Cataduanes and lowlands of S Luzon (Musser and Heaney, 1992). Formerly and incorrectly thought to also occur on Leyte and Dinagat (Musser, 1982b). SYNONYMS: hollisteri. COMMENTS: Included within " Apomys abrae-hylocetes Group " by Musser (1982b)., 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 575, DOI: 10.5281/zenodo.7353098, {"references":["Musser, G. G., and L. R. Heaney. 1992. Philippine rodents: Definitions of Tarsomys and Limnomys plus a preliminary assessment of phylogenetic patterns among native Philippine murines (Murinae, Muridae). Bulletin of the American Museum of Natural History, 211: 1 - 138."]}

Apomys Mearns, 1905. Proc. U.S. Natl. Mus., 28:455. TYPE SPECIES: Apomys hylocoetes Mearns, 1905. COMMENTS: At one time included in Rattus, but is a distinct genus and forms a monophyletic group of its own within the assemblage of Philippine Old Endemics (Musser and Heaney, 1992). Taxonomic history of the genus and preliminary systematic revision provided by Musser (19826); additional taxonomic notes and phylogenetic relationships outlined by Musser and Heaney (1992). Undescribed species recorded from Negros and Sibuyan islands (Musser and Heaney, 1992); actual distribution in archipelago and number of species still unknown., 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 574, DOI: 10.5281/zenodo.7353098, {"references":["Musser, G. G., and L. R. Heaney. 1992. Philippine rodents: Definitions of Tarsomys and Limnomys plus a preliminary assessment of phylogenetic patterns among native Philippine murines (Murinae, Muridae). Bulletin of the American Museum of Natural History, 211: 1 - 138."]}

Apomys Mearns, 1905. Proc. U.S. Nat. Mus., 28:455. REVIEWED BY: B. R. Stein (BRS). COMMENT: This genus was included in Rattus by Corbet and Hill, 1980:172; but see Musser, 1977, Am. Mus. Novit., 2624:11, and Musser, 1981, Bull. Am. Mus. Nat. Hist., 169(2): 136, who considered it a distinct genus., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 508, DOI: 10.5281/zenodo.7353037, {"references":["Corbet, G. B., and J. E. Hill. 1980. A world list of mammalian species. British Museum (Natural History), London, 226 pp."]}

Apomys insignis Mearns, 1905. Proc. U.S. Nat. Mus., 28:459. TYPE LOCALITY: Philippines, S. Mindanao, Mt. Apo. DISTRIBUTION: Mindanao (Philippines)., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 509, DOI: 10.5281/zenodo.7353037

Apomys hylocoetes Mearns, 1905. Proc. U.S. Nat. Mus., 28:456. TYPE LOCALITY: Philippines, Mindanao, Mt. Apo, 2150 m. DISTRIBUTION: Mindanao (Philippines)., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 509, DOI: 10.5281/zenodo.7353037

Apomys datae (Meyer, 1899). Abh. Mus. Dresden, ser. 7, 7:25. TYPE LOCALITY: Philippines, N. Luzon, Lepanto. DISTRIBUTION: Luzon (Philippines). COMMENT: Includes major; see Sanborn, 1950, Fieldiana Zool., 33: 133., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 508, DOI: 10.5281/zenodo.7353037

Apomys petraeus Mearns, 1905. Proc. U.S. Nat. Mus., 28:458. TYPE LOCALITY: Philippines, Mindanao, Mt. Apo, 2340 m. DISTRIBUTION: Mindanao (Philippines)., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 509, DOI: 10.5281/zenodo.7353037

Apomys musculus Miller, 1911. Proc. U.S. Nat. Mus., 38:403. TYPE LOCALITY: Philippines, Luzon, Benguet, Baguio, 1500 m. DISTRIBUTION: Luzon (Philippines)., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 509, DOI: 10.5281/zenodo.7353037

Apomys abrae (Sanborn, 1952). Fieldiana Zool., 33(2): 133. TYPE LOCALITY: Philippines, Luzon Isl., Abra Prov., Abra, 3500 ft. (1067 m). DISTRIBUTION: Luzon (Philippines)., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 5), pp. 504-560 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 508, DOI: 10.5281/zenodo.7353037