Macrotarsomys Petteri
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Petter's big-footed mouse (''Macrotarsomys petteri''), is a Madagascan
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 na ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
'' Macrotarsomys''. With a head and body length of 150 mm (5.9 in) and body mass of 105 g (3.7 oz), it is the largest
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of its genus. Its upper body is brown, darkest in the middle of the back, and the lower body is white to yellowish. The animal has long
whiskers Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to touch, sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are c ...
, short forelimbs, and long hindfeet. The tail ends in a prominent tuft of long, light hairs. The skull is robust and the
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
are low-crowned and
cuspidate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
. Petter's big-footed mouse is now found only in the
Mikea Forest The Mikea Forest (or Forêt des Mikea) is a forested region of southwestern Madagascar, between Manombo and Morombe. More specifically, it stretches from Route Nationale 9 to the west coast and from Mangoky south to the Manombo River. It lies at ...
of southwestern Madagascar, but
subfossil 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 ...
records indicate that it used to be more widely distributed in southern Madagascar. Climatic changes and competition with
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...
may have led to the shift in its distribution. The Mikea Forest, the only place where it is still known to occur, is
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
by human development.


Taxonomy

During a 2003 biological inventory of the
Mikea Forest The Mikea Forest (or Forêt des Mikea) is a forested region of southwestern Madagascar, between Manombo and Morombe. More specifically, it stretches from Route Nationale 9 to the west coast and from Mangoky south to the Manombo River. It lies at ...
, a forest region of southwestern Madagascar, a single specimen of the rodent genus '' Macrotarsomys'' was collected.Goodman and Soarimalala, 2005, p. 452 This animal turned out to be distinct from both previously known species of the genus—''
bastard big-footed mouse The bastard big-footed mouse (''Macrotarsomys bastardi'') is a species of rodent in the family Nesomyidae. It is found only in Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagas ...
'', which is widespread in western Madagascar, and the larger ''
greater big-footed mouse The greater big-footed mouse or long-tailed big-footed mouse (''Macrotarsomys ingens'') is a nocturnal rodent found only in north west Madagascar. It was first described by F. Petter in 1959. It is listed as an endangered species as a result of ...
'', known only from the
Ankarafantsika National Park Ankarafantsika National Park is a national park near Andranofasika in the Boeny Region of Madagascar. The closest city is Majunga north of the park. Ankarafantsika is mostly tropical in climate type. The Sakalava people are the predominant ethnic ...
. Accordingly, Steven Goodman and Voahangy Soarimalala named it in 2005 as a new species, ''Macrotarsomys petteri''.Goodman and Soarimalala, 2005, p. 453 The specific name, ''petteri'', honors French zoologist
Francis Petter Francis may refer to: People *Pope Francis, the head of the Catholic Church and sovereign of the Vatican City State and Bishop of Rome * Francis (given name), including a list of people and fictional characters *Francis (surname) Places * Rural ...
for his contributions to the study of Malagasy rodents.Goodman and Soarimalala, 2005, p. 457 Petter's big-footed mouse, the largest species in the genus, is most similar to greater big-footed mouse, which may be its closest relative.Goodman and Soarimalala, 2005, p. 459


Description

Petter's big-footed mouse is a
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
rodent with short forelimbs and long hindfeet.Goodman and Soarimalala, 2005, p. 454 With a head and body length of 150 mm (5.9 in) and body mass of 105 g (3.7 oz) in the only known complete specimen,Goodman and Soarimalala, 2006, table 1 it is much larger than ''M. bastardi'', and its measurements fall at or above the upper end of the known range of variation in ''M. ingens''. The upperparts are covered with soft and short, brown fur. Most cover hairs (the main part of the fur) are dark brown for the two-thirds closest to the base, then light brown, with a short dark brown tip. The middle of the back appears darker, because the cover hairs there are entirely dark brown. The hairs are 6 to 8 mm (0.2 to 0.3 in) long on the shoulders and 7 to 9 mm (0.3 to 0.4 in) on the back. The
guard hair Guard hair or overhair is the outer layer of hair of most mammals, which overlay the fur. Guard hairs are long and coarse and protect the rest of the pelage (fur) from abrasion and frequently from moisture. They are visible on the surface of the fu ...
s are gray. Because the flanks lack entirely dark cover hairs, they are slightly lighter than the rest of the upperparts. They are sharply separated in color from the underparts, which are entirely white to buffish. The mystacial vibrissae (whiskers above the mouth) are long, up to 60 mm (2.4 in), and white or black in color. The
pinnae The auricle or auricula is the visible part of the ear that is outside the head. It is also called the pinna (Latin for "wing" or "fin", plural pinnae), a term that is used more in zoology. Structure The diagram shows the shape and location of ...
(external ears) are dark brown and covered with fine gray hairs,Goodman and Soarimalala, 2005, p. 455 and ear length is 32 mm (1.3 in). Hindfoot length is 37 mm (1.5 in). The upper sides of the feet are covered with grayish white fur, which extends around the claws to form
ungual tufts In mammals, ungual tufts are tufts of hairs at the base of claws of the forefeet and hindfeet. Their presence has been used as a character in cladistic studies of the Cricetidae, a large family of rodents.Weksler, 2006, p. 19 Rice rats Membe ...
. On the hindfeet, the fifth digit is relatively short at 6 mm (0.2 in); the hallux (first digit) is 8 mm (0.3 in) long, and the other digits 11 to 12 mm (0.4 to 0.5 in). The tail is 238 mm (9.4 in) long and naked in part. At the base, it is dark brown both above and below, but slightly lighter below. The upper side remains dark brown for much of its length, though the color does become lighter towards the tip. The lower side becomes mottled at about 55 mm (2.2 in) from the tip and then whitish at about 65 mm (2.6 in). It has a well-developed tuft at its tail tip, consisting of whitish and occasional light brown hairs. This tuft commences at about 130 mm (5.1 in) from the base with fairly short hairs and becomes more pronounced at 180 mm (7.1 in).Goodman and Soarimalala, 2005, p. 456 In contrast, the greater big-footed mouse has a weaker, dark brown tuft. Petter's big-footed mouse has a large and robust skull with well-developed
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygomati ...
es (cheekbones). The
interorbital region The interorbital region of the skull is located between the eyes, anterior to the braincase. The form of the interorbital region may exhibit significant variation between taxonomic groups. In oryzomyine rodents, for example, the width, form, an ...
of the skull (between the eyes) is smooth, as in greater big-footed mouse, and lacks the shelves characteristic of the bastard big-footed mouse. The
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
is broad and the
incisive foramina In the human mouth, the incisive foramen (also known as: "''anterior palatine foramen''", or "''nasopalatine foramen''") is the opening of the incisive canals on the hard palate immediately behind the incisor teeth. It gives passage to blood vesse ...
(openings in the front portion of the palate) are long and broad. In the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
(lower jaw), the root of the lower
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, whe ...
is housed in a distinct
capsular process In rodents, the capsular process or projection is a bony capsule that contains the root of the lower incisor. It is visible on the labial (outer) side of the mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest ...
, a protuberance at the back of the jawbone. The lower masseteric ridge (a crest on the outer side of the mandible) is prominent.Goodman et al., 2006, p. 957 As is typical of ''Macrotarsomys'', the
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
are
cuspidate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
and low-crowned.


Distribution and ecology

The single known living specimen, a young adult male, was collected at 80 m (260 ft) altitude in the Andaladomo forest (part of the Mikea Forest) in 2003. The Andaladomo forest is different in vegetation from the rest of the Mikea Forest, and is similar to forests further to the north on Madagascar.Goodman and Soarimalala, 2005, p. 460 The animal was found in an isolated fragment of dry deciduous forest amid land cleared for
maize Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. Th ...
cultivation. The trap was set at the foot of a tree surrounded by bushes and succulent plants. Other small mammals known from the Mikea Forest include the bastard big-footed mouse, the introduced
black rat The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is n ...
(''Rattus rattus''), several species of
tenrec A tenrec is any species of mammal within the afrotherian family Tenrecidae endemic to Madagascar. Tenrecs are wildly diverse; as a result of convergent evolution some resemble hedgehogs, shrews, opossums, rats, and mice. They occupy aquatic, a ...
s, and the shrew ''
Suncus madagascariensis The Madagascan pygmy shrew (''Suncus madagascariensis'') is a species of mammal in the family Soricidae. It is the only known Malagasy shrew. Taxonomy Some taxonomists regard it as conspecific with the widely distributed Etruscan shrew, the s ...
''. Although only a single individual of Petter's big-footed mouse was caught during Goodman and Soarimalala's survey, which accrued 3100 trap-nights, they argue that this does not necessarily mean the species is rare, since trapping rates for rodents in the dry forests of Madagascar are often variable depending on year and season. Nothing is known of its behavior, but the animal's morphology suggests it lives on the ground.Garbutt, 2007, p. 242 Subsequent to its discovery at the Mikea Forest, Petter's big-footed mouse was also found as a
subfossil 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 ...
in cave deposits at Andrahomana in far southeastern Madagascar, a find reported in 2006.Goodman et al., 2006, p. 753 There, it was found together with more abundant remains of the introduced black rat and
house mouse The house mouse (''Mus musculus'') is a small mammal of the order Rodentia, characteristically having a pointed snout, large rounded ears, and a long and almost hairless tail. It is one of the most abundant species of the genus '' Mus''. Althoug ...
(''Mus musculus''), as well as indigenous rodents such as the bastard big-footed mouse. Two Petter's big-footed mouse bones were
radiocarbon dated Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was dev ...
to 790–410 BCE and 150–390 CE, respectively, a period when the local climate became drier and humans first appeared. ''Macrotarsomys'' species are thought to burrow in sandy ground and would not be expected to enter caves; therefore, the subfossils are probably remains of animals eaten by birds of prey. Although Petter's big-footed mouse could conceivably persist in remnant pockets of wet habitat in southeastern Madagascar, searches at two sites near Andrahomana failed to confirm its presence. It may have become locally extinct in the area because of the drying climate and competition with the black rat.Goodman et al., 2006, p. 960 In 2009, it was also recorded from the cave of Ankilitelo in southwestern Madagascar. Remains of a large ''Macrotarsomys'' had previously been reported from other southern Madagascar sites, and at least some of these may be ''M. petteri''.Goodman and Soarimalala, 2005, p. 459; Goodman et al., 2006, p. 960 A
karst Karst is a topography formed from the dissolution of soluble rocks such as limestone, dolomite, and gypsum. It is characterized by underground drainage systems with sinkholes and caves. It has also been documented for more weathering-resistant ro ...
ic deposit near
Lake Tsimanampetsotsa Lake Tsimanampetsotsa (french: Lac Tsimanampetsotsa) (also called Lake Tsimanampesotse) is a moderately Soda lake, alkaline lake in the Toliara Province, in the southwestern part of Madagascar. It is located at around . The lake is protected wit ...
(dated to the Late
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
on unclear grounds) contained three species of ''Macrotarsomys'', including a very large one that may well be ''M. petteri''. Remains identified as greater big-footed mouse have been reported from a cave at
Ankazoabo Ankazoabo is a town in Atsimo-Andrefana Region, Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in ...
in southern Madagascar; these may also be Petter's big-footed mouse.


Conservation status

The
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
assesses Petter's big-footed mouse as "
Data Deficient A data deficient (DD) species is one which has been categorized by the International Union for Conservation of Nature (IUCN) as offering insufficient information for a proper assessment of conservation status to be made. This does not necessaril ...
", but notes that the species will very probably qualify as
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
if its current distribution turns out to be restricted to primary forest in the Mikea Forest. The Mikea Forest is one of the largest remaining forests of southwestern Madagascar, but it is not protected and is threatened by logging, pasture, and conversion to agricultural land.


References


Literature cited

*Garbutt, N. 2007. Mammals of Madagascar: A Complete Guide. A & C Black, 304 pp. *Goodman, S.M. and Soarimalala, V. 2005
A new species of ''Macrotarsomys'' (Rodentia: Muridae: Nesomyinae) from southwestern Madagascar
(subscription required). Proceedings of the Biological Society of Washington 118(2):450–464. *Goodman, S.M., Vasey, N. and Burney, D.A. 2006
The subfossil occurrence and paleoecological implications of ''Macrotarsomys petteri'' (Rodentia: Nesomyidae) in extreme southeastern Madagascar
(subscription required). Comptes Rendus Palevol 5:753–762 * *Muldoon, K.M., De Blieux, D.D., Simons, E.L. and Chatrath, P.S. 2009
The subfossil occurrence and paleoecological significance of small mammals at Ankilitelo Cave, southwestern Madagascar
(subscription required). Journal of Mammalogy 90(5):1111–1131. {{Featured article Macrotarsomys Mammals of Madagascar Mammals described in 2005