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The large Mindoro forest mouse (''Apomys gracilirostris'') is a species of
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are nat ...
in the family
Muridae The Muridae, or murids, are the largest family of rodents and of mammals, containing approximately 1,383 species, including many species of mice, rats, and gerbils found naturally throughout Eurasia, Africa, and Australia. The name Muridae comes ...
, from the genus '' Apomys''. It is found only in the
Philippines The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no), * bik, Republika kan Filipinas * ceb, Republika sa Pilipinas * cbk, República de Filipinas * hil, Republ ...
. Its natural
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
is subtropical or tropical moist
montane forest Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
s. It is a large mouse with large feet, a long tail and an elongated snout which is morphologically unique within its genus. It is covered in soft fur which is mostly dark brown in colour. Its closest relative is thought to be the Luzon montane forest mouse, based on genetic and morphological similarities.


Discovery and taxonomy

In May and June 1992, an expedition to the Philippines was organised for the purpose of increasing the knowledge on their biodiversity. During this expedition, sixteen examples of a then unknown species of mouse were captured on
Mount Halcon Mount Halcon ( fil, Bundok Halcon) and ( es, Monte Halcón) is the highest mountain in the island of Mindoro in the Philippines, according to the new data release by ''Oriental Mindoro peakvisor'' as of 2022, it has an elevation of above sea lev ...
, on the island
Mindoro Mindoro is the seventh largest and eighth-most populous island in the Philippines. With a total land area of 10,571 km2 ( 4,082 sq.mi ) and has a population of 1,408,454 as of 2020 census. It is located off the southwestern coast of Luz ...
. To this day, these animals, all captured between May 28 and June 12, 1992, remain the only known specimens of this species. In 1995, in the scientific journal ''
Proceedings of the Biological Society of Washington In academia and librarianship, conference proceedings is a collection of academic papers published in the context of an academic conference or workshop. Conference proceedings typically contain the contributions made by researchers at the conferen ...
'', biologist Luis A. Ruedas described the animal as ''Apomys gracilirostris''. At the time it was the ninth known species of the
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
Filipino genus ''Apomys'', and the second found on Mindoro, after ''Apomys musculus'' ( least forest mouse). The editors of a compilation work on the indigenous mammals of the Philippines accepted the animal as a new species in 1998. ''A. gracilirostris'' was the first newly described ''Apomys'' since 1962, when ''Apomys sacobianus'' ( long-nosed Luzon forest mouse) was described. However, it was far from being the only new species discovered in the Philippines. Several new rodents had already been discovered in the eighties, one of which was actually from Mindoro as well (''Anonymomys mindorensis'' or Mindoro climbing rat). A constant supply of new species remained in the years after the description of ''A. gracilirostris''. In ten years time, eight new mammal species were added to the list. In 2006, another new ''Apomys'' was described: '' Apomys camiguinensis'' (Camiguin forest mouse). ''Apomys'' also comprises several species which are yet undescribed, two of which are from Mindoro. Another species from Mindoro, from the predominantly
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guinea. In ...
n genus ''
Maxomys ''Maxomys'' is a genus of rodents, widespread in Southeast Asia. They are mid-sized rodents, similar to rats, that live on the ground of tropical rainforests. There they build nests, padded with fallen leaves from trees. They feed on roots, falle ...
'', has yet to receive its scientific name. In accordance with the
International Code of Zoological Nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the I ...
, the name ''Apomys gracilirostris'' is a binomen. The generic name, ''Apomys'' was proposed by American biologist Edgar Mearns in 1905 and has the meaning of "mouse from
Mount Apo Mount Apo, also known locally as Apo Sandawa, is a large solfataric, dormant stratovolcano on the island of Mindanao, Philippines. With an elevation of above sea level, it is the highest-mountain in the Philippine Archipelago, Mindanao and ...
" (''mys'', μῦς, being the Ancient Greek word for "mouse"); this because the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
of the genus was first found on Mount Apo. The specific name, ''gracilirostris'', is a combination of the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
words ''gracilis'' "slender" and ''rostrum'' "snout" and refers to the animal's long, slender snout.


Identification

The genus ''Apomys'', of which the large Mindoro forest mouse is a member, can be identified by its small size, a long tail, elongate, narrow hind feet, the presence of four
abdominal The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
mammary glands A mammary gland is an exocrine gland in humans and other mammals that produces milk to feed young offspring. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in prima ...
, and a large number of
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
characteristics. Within this genus, the large Mindoro forest mouse has several specific identifying features, the most important of which are its very thin and short upper
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wh ...
s, long lower incisors and long snout. The tail and feet are also relatively long, even within the genus ''Apomys''. The characteristics of the incisors are so peculiar that when Ruedas discovered the species, he initially thought he had found a new genus. The fur of the large Mindoro forest mouse is soft and smooth. The hairs of the dorsal fur are light gray near the root, and dark brown at the tip. Between the normal hairs are many black tactile hairs. This makes the dorsal fur appear dark brown, although recently captured animals have a dark blue-green hue. Some animals are darker around the midline of the back, and all are a lighter colour at the flanks. The colour of the ventral fur varies between animals, with it being a yellow-brown colour in some, slightly lighter coloured than the dorsal fur in others, still others having gray hairs with brown or silver-coloured tips in the ventral fur, and finally some animals showing no difference between the dorsal and ventral fur at all. The tail is usually an even, dark colour, but in some animals the ventral side is somewhat darker. In some animals the tail ends in a white tip of 2 to 10 mm in length. The tail has fourteen
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
per centimeter. The long, dark hind feet end in long claws (about 4 mm). The claws on the front feet are about 3 mm in length. The animal has seven
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In sa ...
, thirteen
thoracic vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
, six
lumbar vertebrae The lumbar vertebrae are, in human anatomy, the five vertebrae between the rib cage and the pelvis. They are the largest segments of the vertebral column and are characterized by the absence of the foramen transversarium within the transverse p ...
, three
sacral vertebrae The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
and thirty-two to thirty-five caudal vertebrae.


Evolution and phylogenetic relationships

The large Mindoro forest mouse belongs to the ''Chrotomys'' division, a group within the
Murinae The Old World rats and mice, part of the subfamily Murinae in the family Muridae, comprise at least 519 species. Members of this subfamily are called murines. In terms of species richness, this subfamily is larger than all mammal families excep ...
that occurs exclusively on the Philippines and in addition to ''Apomys'' also includes '' Rhynchomys'', '' Chrotomys'' and '' Archboldomys''. Animals in this division share several morphological and genetic features. Within this group, ''Apomys'' is by far the biggest and most extensive genus, containing small, inconspicuous wood mice which are common to the whole of the Philippines, while the other, more specialized genera are barely ever found outside
Luzon Luzon (; ) is the largest and most populous island in the Philippines. Located in the northern portion of the Philippines archipelago, it is the economic and political center of the nation, being home to the country's capital city, Manila, a ...
. ''Apomys'' itself was divided in two groups by American biologist Guy Musser in 1982, the first being the ''datae'' group, containing only the Luzon montane forest mouse (''A. datae''), and the second the ''abrae-hylocetes'' group, containing all other species. Animals in these two groups differ in the way in which the head is supplied with blood from arteries. Ruedas placed the large Mindiro forest mouse in the ''datae'' group. This relationship is further supported by other similarities: both species are relatively large for the genus and have a relatively long snout. In 2003, a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
study was published which compared DNA sequences from the cytochrome b gene of thirteen species of ''Apomys''. This study confirmed the proposed relationship between the large Mindoro forest mouse (''A. gracilirostris'') and the Luzon montane forest mouse (''A. datae''). The phylogenetic relationships of the large Mindoro forest mouse can be summarized as follows. This study also found that the large Mindoro forest mouse was the only ''Apomys'' species that displayed deep within-species divergences, which were calculated to date back some 400.000 years. The split between the large Mindoro forest mouse and the Luzon montane forest mouse dates back further still at an estimated three million years ago, putting it in the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotid ...
. Another, more elaborate phylogenetic study on all of the endemic Filipino genera and species of
Murinae The Old World rats and mice, part of the subfamily Murinae in the family Muridae, comprise at least 519 species. Members of this subfamily are called murines. In terms of species richness, this subfamily is larger than all mammal families excep ...
, of which the large Mindoro forest mouse is a member, produced different results. According to this study, the ''Chromotys'' division originated some ten million years ago (rather than the estimate of six million years ago from the previous study). It was further calculated that the split between the ''Chromotys'' division and its closest relatives, a predominantly
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
n group including '' Mus'', ''
Otomys African vlei rats (''Otomys''), also known as groove-toothed rats, live in many areas of sub-Saharan Africa. Most species live in marshlands, grasslands, and similar habitats and feed on the vegetation of such areas, occasionally supplementin ...
'' and ''
Mastomys ''Mastomys'' is a genus of rodent in the family Muridae endemic to Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square&nbs ...
'', had happened about sixteen million years ago. However, since the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
of ''Murinae'' from the Philippines is non-existent, and controversial for Murinae from other areas, these estimates can only be based on the molecular clock, and not on direct
paleontological Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
data. The large Mindoro forest mouse is the third known rodent endemic to Mindoro, after the Mindoro black rat and the Mindoro climbing rat, and not including the Ilin Island cloudrunner, the occurrence of which has not been confirmed on the island. However, in a compilation work on the indigenous mammals of the Philippines, published in 1998, it was announced that there is a second ''Apomys'' on Mindoro. This species has yet to be described and has temporarily been given the name of "''Apomys'' sp. E". It may be related to two other undescribed species from
Sibuyan Island Sibuyan is a crescent-shaped island, the second largest in an archipelago comprising Romblon Province, Philippines. Located in the namesake Sibuyan Sea, it has an area of and has a total population of 62,815 as of 2020 census. The island has t ...
and Greater Negros-Panay, ''Apomys'' sp. A/C and ''Apomys'' sp. B. The local population of ''A. musculus'', the only other ''Apomys'' known on the island, may also represent a separate species. Several other species occur exclusively on Mindoro (in addition to the undescribed ''Maxomys'' species mentioned earlier), including Oliver's warty pig, the
tamaraw The tamaraw or Mindoro dwarf buffalo (''Bubalus mindorensis'') is a small hoofed mammal belonging to the family Bovidae. It is endemic to the island of Mindoro in the Philippines, and is the only endemic Philippine bovine. It is believed, h ...
and the
megabat Megabats constitute the family Pteropodidae of the order Chiroptera (bats). They are also called fruit bats, Old World fruit bats, or—especially the genera '' Acerodon'' and ''Pteropus''—flying foxes. They are the only member of the ...
'' Styloctenium mindorensis''. The relatively large number of endemic species on Mindoro can be explained from the fact that, in all probability, Mindoro was never connected to any other landmass. This enabled the animals present on the island to develop in isolation from their relatives. A remarkable
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
aspect of Mindoro is its status as a transition area between Greater Palawan and the rest of the Philippines. On the one hand, species like Oliver's warty pig and the tamaraw are clearly related to animals from Greater Palawan and the rest of South-East Asia, but on the other, Mindoro also harbours animals from genera like ''Apomys'' and ''Chrotomys'', which are clearly Filipino. The latter animals reached Mindoro from Luzon, or possibly in some cases Greater Negros-panay. The large Mindoro forest mouse, with its close relationship to the Luzon montane forest mouse, probably falls into the group of animals that reached the island from Luzon.Musser and Carleton, 2005, p. 1282


Ecology and behaviour

The large Mindoro forest mouse has been found at three different sites in the municipality of San Teodoro, on the North flank of Mount Halcon. These sites are located at altitudes between . There is a good possibility, however, that the large Mindoro forest mouse also occurs on other mountains on Mindoro, especially on the northern part of the island, which is home to large areas of undamaged
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level clou ...
. The lowest forests in which the large Mindoro forest mouse has been found, at some above sea level, are mostly between in height, with the very tallest reaching . The dominant species of tree are from the genera '' Leptospermum flavescens'', '' Tristaniopsis'' and ''
Lithocarpus ''Lithocarpus'' is a genus in the beech family, Fagaceae. Trees in this genus are commonly known as the stone oaks and differ from ''Quercus'' primarily because they produce insect-pollinated flowers on erect spikes and the female flowers hav ...
'', from the families
Myrtaceae Myrtaceae, the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All specie ...
and
Fagaceae The Fagaceae are a family of flowering plants that includes beeches, chestnuts and oaks, and comprises eight genera with about 927 species. Fagaceae in temperate regions are mostly deciduous, whereas in the tropics, many species occur as everg ...
. These very humid forests have a dense
understory In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abov ...
vegetation, which consists, amongst other plants, of
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hornw ...
es,
tree ferns The tree ferns are arborescent (tree-like) ferns that grow with a trunk elevating the fronds above ground level, making them trees. Many extant tree ferns are members of the order Cyatheales, to which belong the families Cyatheaceae (scaly tree ...
and species of ''
Pandanus ''Pandanus'' is a genus of monocots with some 750 accepted species. They are palm-like, dioecious trees and shrubs native to the Old World tropics and subtropics. The greatest number of species are found in Madagascar and Malaysia. Common names ...
''. At higher altitudes (between ) the forest is dominated by
bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, but ...
, while the only tree species present is '' Agathis philippinensis''. Here, vegetation reaches a height between . There are many smaller plants present, such as
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s,
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Along with the Asteraceae, they are one of the two largest families of flowering ...
s and species of ''Pandanus''. The forest floor is covered with a layer of leaves with a thickness between . Nothing is known with any certainty about the behaviour of the large Mindoro forest mouse. The long snout, small upper incisors and long lower incisors suggest that the animal feeds on soft
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s, but this is not certain. Furthermore, the long tail and elongated hind feet with well-developed claws point towards a climbing lifestyle. Very little is also known about the animal's
procreation Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual org ...
habits. A female specimen was found to carry three
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm c ...
s. Another female had a swollen
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
, and
testicle A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testostero ...
s were visible in several of the male specimens.


References


Bibliography

* *Musser, G.G. 1982. Results of the Archbold Expeditions. No. 108. The definition of ''Apomys'', a native rat of the Philippine islands. ''American Museum Novitates'' 2746:1–43. * {{Taxonbar, from=Q307840 Rats of Asia Apomys Endemic fauna of the Philippines Fauna of Mindoro Rodents of the Philippines Mammals described in 1995 Taxonomy articles created by Polbot