Luzon montane forest mouse
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The Luzon montane forest mouse (''Apomys datae'') is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Muridae The Muridae, or murids, are either the largest or second-largest family of rodents and of mammals, containing approximately 870 species, including many species of mice, rats, and gerbils found naturally throughout Eurasia, Africa, and Australia. ...
, from the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
'' Apomys''. It occurs only in the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
, where it has been found on the large northern island
Luzon Luzon ( , ) is the largest and most populous List of islands in the Philippines, island in the Philippines. Located in the northern portion of the List of islands of the Philippines, Philippine archipelago, it is the economic and political ce ...
(in the Cordillera Central and on the coast of
Ilocos Norte Ilocos Norte (), officially the Province of Ilocos Norte (; ), is a Provinces of the Philippines, province of the Philippines located in the Ilocos Region. It is located in the northwest corner of Luzon island, bordering Cagayan and Apayao to t ...
). It is most closely related to the large Mindoro forest mouse, which occurs on
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 ), it has a population of 1,408,454, as of the 2020 census. It is located off the southwestern coast of ...
. There may be another related species in the Sierra Madre, but this species is yet undescribed. The Luzon montane forest mouse is a relatively large, ground-dwelling rat with a tail that is quite short for its genus.


Discovery

The Luzon montane forest mouse was the first species of ''Apomys'' ever to be discovered. In 1895, an expedition was organised which brought to Europe the first specimens of several genera, including ''
Carpomys ''Carpomys'' is a genus of rodent in the family Muridae. It contains two extant species, and one extinct species: * '' Carpomys melanurus'' * '' Carpomys phaeurus'' *'' Carpomys dakal'' It was first described by Oldfield Thomas Michael Roger ...
'', '' Rhynchomys'' and '' Crunomys''. During this expedition, in February,
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
explorer Exploration is the process of exploring, an activity which has some Expectation (epistemic), expectation of Discovery (observation), discovery. Organised exploration is largely a human activity, but exploratory activity is common to most organis ...
John Whitehead captured a number of unknown rats on a site called ''Lepanto'' on Mount Data, at an altitude of approximately . In 1898, British biologist
Oldfield Thomas Michael Rogers Oldfield Thomas (21 February 1858 – 16 June 1929) was a British zoologist. Career Thomas worked at the Natural History Museum, London, Natural History Museum on mammals, describing about 2,000 new species and subspecies for ...
described these animals as an "interesting species", but identified them as ''Mus chrysocomus'', a species from
Sulawesi Sulawesi ( ), also known as Celebes ( ), is an island in Indonesia. One of the four Greater Sunda Islands, and the List of islands by area, world's 11th-largest island, it is situated east of Borneo, west of the Maluku Islands, and south of Min ...
that is now known as
yellow-haired hill rat The yellow-haired hill rat (''Bunomys chrysocomus'') is a species of rodent in the family Muridae. It is found only in Sulawesi, Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, betwee ...
(''Bunomys chrysocomus'') and reckoned among the genus ''
Bunomys ''Bunomys'' is a genus of rodent from Sulawesi and Buton Island. Currently, eight species are recognised in two species-groups. Species Genus ''Bunomys'' ''chrysocomus''-group: * Yellow-haired hill rat, ''Bunomys chrysocomus'' Hoffmann, 1887 * ...
'', which is not actually closely related to ''Apomys''. Thomas sent a specimen to the Staatliches Museum für Tierkunde Dresden, where
Adolf Bernard Meyer Adolf Bernhard Meyer (11 October 1840, Hamburg – 22 August 1911, Dresden) was a German anthropologist, ornithologist, entomologist, and Herpetology, herpetologist. He served for nearly thirty years as director of the Königlich Zoologisches und ...
concluded that the animal did not resemble ''Mus chrysocomus''. Meyer described th animal as ''Mus datae'' in 1899, after its type locality – Mount Data (at the time the generic name '' Mus'' was used more broadly than it is now). For a long time, little was known about ''Mus datae'', until 1913, when American biologist Ned Hollister described eight rats from Luzon under the name ''Epimys datae'' ("''Epimys''" was the name of the genus that would later become ''
Rattus ''Rattus'' is a genus of muroid rodents, all typically called rats. However, the term rat can also be applied to rodent species outside of this genus. Species and description The best-known ''Rattus'' species are the black rat (''R. rattus'') ...
''). These were in fact examples of the Himalayan field rat (''Rattus nitidus''), but they were only identified as such in 1977, by
Guy Musser Guy Graham Musser (August 10, 1936 – October 2019) was an American zoologist. His main research was in the field of the rodent subfamily Murinae, in which he has described many new species. Musser was born in Salt Lake City, Utah. He attended ...
, another American biologist. Meanwhile, in Britain, John Ellerman had finally placed ''Mus datae'' with its relatives in ''Apomys'', in 1941. Eleven years later, in 1952, American
zoologist Zoology ( , ) is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one ...
Colin Campbell Sanborn announced that he had captured 54 specimens of ''A. datae'' on Mount Data. A good part of this catch, however, was later found to consist of specimens of the Luzon Cordillera forest mouse (''A. abrae''), a species which had been described by Sanborn in the same article in which he had made his announcement. In a 1982 article, Musser defined the genus ''Apomys'' and gave the first modern description of ''A. datae'', while also correcting Sanborn's mistake in the identification of his collection. It was revealed that Sanborn had not been the only one to get the major species of ''Apomys'' from Luzon confused: the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of the species ''Apomys Major'', described by Gerrit Smith Miller Jr. of the
Smithsonian Institution The Smithsonian Institution ( ), or simply the Smithsonian, is a group of museums, Education center, education and Research institute, research centers, created by the Federal government of the United States, U.S. government "for the increase a ...
in 1910, turned out to have been a specimen of ''A. datae'', while the other animals to have been identified as ''A. major'' were discovered to be examples of ''A. abrae''. Since that time, ''Apomys major'' has been considered a subjective synonym of ''A. datae''. Musser identified ''A. datae'' as the only member of the "''Apomys datae'' group" within the genus, being different from all other species. In 1993 and 1994, the species was observed in the Sierra Madre, at an altitude between , but this probably concerns a population of a separate, undescribed species. A second species within the ''A. datae'' group was described by Luis Ruedas, in 1995: '' Apomys gracilirostris''. In the 21st century, the knowledge about ''A. datae'' was expanded with data from genetic research. In 2002, the
karyotype A karyotype is the general appearance of the complete set of chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is discerned by de ...
was revealed, and in 2003, the
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationship with ''A. gracilirostris'' was confirmed, based on common features in the species'
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
. Most recently, the animal has been found on several new locations in North Luzon.


Evolution and phylogenetic relationships

The Luzon montane 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 The genus ''Chrotomys'' contain a unique group of rodents found only in the Philippines, specifically the islands of Luzon, Mindoro, and Sibuyan. Instead of being predominantly herbivorous or omnivorous like other murines, these rats feed pre ...
'' 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. ''Apomys'' itself was divided into two groups, in the aforementioned article by Musser from 1982: the ''datae'' group, containing only ''A. datae'', and 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 Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood is com ...
from
arteries An artery () is a blood vessel in humans and most other animals that takes oxygenated blood away from the heart in the systemic circulation to one or more parts of the body. Exceptions that carry deoxygenated blood are the pulmonary arteries in ...
. Since the publication of Musser's article, another species has been described that falls into the ''datae'' group: ''A. gracilirostris''. 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 study was published which compared DNA sequences from the
cytochrome b Cytochrome b is a protein found in the membranes of aerobic cells. In eukaryotic mitochondria (inner membrane) and in aerobic prokaryotes, cytochrome b is a component of respiratory chain complex III () — also known as the bc1 complex or ubiq ...
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''), as well as the status of the ''datae'' group as a sister group of the other species of ''Apomys''. The phylogenetic relationships of the Luzon montane forest mouse can be summarized as follows. According to this study, the split between the Luzon montane forest mouse and the large Mindoro forest mouse took place some three million years ago, as calculated using a
molecular 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 nucleot ...
, putting it in the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
n group including ''Mus'', '' Otomys'' and ''
Mastomys ''Mastomys'' is a genus of rodent in the family Muridae endemic to Africa. It contains eight species: * Angolan multimammate mouse (''M. angolensis'') * Awash multimammate mouse or Awash mastomys (''M. awashensis'') * Southern multimammate mouse ...
'', took place some sixteen million years ago. Seeing as ''Apomys'' probably originated on Luzon, the Luzon montane forest mouse likely developed directly from the ancestor species of the ''datae'' group, while the large Mindoro forest mouse is the result of a pliocene migration to Mindoro.


Identification

The genus ''Apomys'', of which the Luzon montane forest mouse is a member, can be identified by its small size, long tail, elongate, narrow hind feet, the presence of four abdominal
mammary glands A mammary gland is an exocrine gland that produces milk in humans and other mammals. Mammals get their name from the Latin word ''mamma'', "breast". The mammary glands are arranged in organs such as the breasts in primates (for example, human ...
, and a large number of
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
characteristics. The Luzon montane forest mouse is a large, thickset species with a tail that is about the same length as the body. The soft, thick dorsal fur is dark brown, while the ventral side of the body is a cream white. The hind feet are partly brown in colour on the dorsal side, but otherwise white. The tail is brown on the dorsal side, and a cream white ventrally. The animal has a large skull with a square-shaped
neurocranium In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the cal ...
. Some of its skull characteristics are so special that they set the animal apart from most every other species of ''Apomys''. One specimen has a head-torso-length of 143 mm, a tail length of 144 mm and a hind feet length of 34 mm., specifies these measures as belonging to example USNM 151513, the holotype of ''Apomys major'', but mentions in a footnote that it concerns the holotype of ''Mus datae''. The specimens captured by Sanborn on Mount Data have a
mean A mean is a quantity representing the "center" of a collection of numbers and is intermediate to the extreme values of the set of numbers. There are several kinds of means (or "measures of central tendency") in mathematics, especially in statist ...
skull length of 39.2 mm (37.0 to 40.6 mm with a
standard deviation In statistics, the standard deviation is a measure of the amount of variation of the values of a variable about its Expected value, mean. A low standard Deviation (statistics), deviation indicates that the values tend to be close to the mean ( ...
of 1.1 mm). The Luzon montane forest mouse's holotype has a skull length of 39.9 mm; that of ''A. major'' has a length of 38.4 mm. The animal has a karyotype of 2n=44 and FN=54, which consists of five pairs of four-armed chromosomes and sixteen pairs of two-armed ( telocentric) chromosomes. The X-chromosome is a large, and the Y-chromosome a small telocentric chromosome. This karyotype has much in common with that of the least forest mouse (''Apomys musculus''), while at the same time being very different from the karyotypes of species form other parts of the Philippines (the karyotype of the large Mindoro forest mouse, on a side note, is unknown).


Notes and references


Further reading

* * * * * {{Taxonbar, from=Q307898 Apomys Fauna of Luzon Mammals described in 1899 Taxonomy articles created by Polbot