Eremoryzomys Polius
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''Eremoryzomys polius'', also known as the gray rice rat or the Marañon oryzomys,Musser and Carleton, 2005, p. 1153 is a
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 n ...
species in the tribe
Oryzomyini Oryzomyini is a tribe of rodents in the subfamily Sigmodontinae of the family Cricetidae. It includes about 120 species in about thirty genera,Weksler et al., 2006, table 1 distributed from the eastern United States to the southernmost parts of ...
of the family
Cricetidae The Cricetidae are a family of rodents in the large and complex superfamily Muroidea. It includes true hamsters, voles, lemmings, muskrats, and New World rats and mice. At almost 608 species, it is the second-largest family of mammals, and h ...
. Discovered in 1912 and first described in 1913 by Wilfred Osgood, it was originally placed in ''
Oryzomys ''Oryzomys'' is a genus of semiaquatic rodents in the tribe Oryzomyini living in southern North America and far northern South America. It includes eight species, two of which—the marsh rice rat (''O. palustris'') of the United States and ...
'' and named ''Oryzomys polius''. In 2006, a
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis found that it was not closely related to ''Oryzomys'' in the strict sense or to any other oryzomyine then known, so that it is now placed in its own
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
, ''Eremoryzomys''. The Brazilian genus ''
Drymoreomys ''Drymoreomys'' is a rodent genus in the tribe Oryzomyini that lives in the Atlantic Forest of Brazil. The single species, ''D. albimaculatus'', is known only from the states of São Paulo and Santa Catarina and was not named until 2011. I ...
'', named in 2011, is probably the closest relative of ''Eremoryzomys''. ''Eremoryzomys'' has a limited distribution in the dry upper valley of the
Marañón River , name_etymology = , image = Maranon.jpg , image_size = 270 , image_caption = Valley of the Marañón between Chachapoyas ( Leimebamba) and Celendín , map = Maranonrivermap.png , map_size ...
in central Peru, but may yet contain more than one species. A large, long-tailed rice rat, with head and body length of , ''E. polius'' has gray fur and short ears. There are well-developed
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 ...
of hair on the hindfeet. Females have eight
mammae 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 primat ...
. The rostrum (front part of the skull) is long and robust and the braincase is rounded. The bony
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 separ ...
is relatively short. The IUCN assesses the
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservatio ...
of the species as " Data Deficient"; it is poorly known but may be threatened by
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
.


Taxonomy

The first two specimens of ''Eremoryzomys polius'' were collected by Wilfred Osgood and M.P. Anderson in 1912. The next year, Osgood described these animals as a new species in the genus ''
Oryzomys ''Oryzomys'' is a genus of semiaquatic rodents in the tribe Oryzomyini living in southern North America and far northern South America. It includes eight species, two of which—the marsh rice rat (''O. palustris'') of the United States and ...
'', ''Oryzomys polius''.Osgood, 1913, p. 97 Osgood wrote that he was unable to find any species closely related to ''O. polius'' and compared it with ''O. xanthaeolus'' (currently '' Aegialomys xanthaeolus'') "for convenience". Its relationships remained obscure ever afterward and it was never assigned to any of the several groups of species recognized within ''Oryzomys''. In 2006, Marcelo Weksler published a large-scale
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis of
Oryzomyini Oryzomyini is a tribe of rodents in the subfamily Sigmodontinae of the family Cricetidae. It includes about 120 species in about thirty genera,Weksler et al., 2006, table 1 distributed from the eastern United States to the southernmost parts of ...
("rice rats"), the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
to which ''O. polius'' belongs. He used both morphological data and molecular characters from the IRBP gene. In all of his analyses, ''O. polius'' was found to be part of clade D, one of four large groups within Oryzomyini, as the sister group to a clade containing all the other species of clade D. Clade D was supported by two shared derived ( synapomorphic) molecular characters and by seven morphological synapomorphies—the tail has a different color above and below; the
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is nam ...
extends to the side of the skull; 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
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 separ ...
) extend back between the first molars; the
posterolateral palatal pits In anatomy, posterolateral palatal pits are gaps at the sides of the back of the bony palate, near the last molars.Weksler, 2006, p. 34 Posterolateral palatal pits are present, in various degrees of development, in several members of the roden ...
(perforations of the palate near the third molars) are complex; the sphenopalatine vacuities (openings in the mesopterygoid fossa, the gap behind the end of the palate) are large; the pattern of the
arterial An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pu ...
circulation in the head is
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
; and the posteroloph (a crest at the back) is present on the third upper molar. Two other molecular synapomorphies supported the clade of all members of clade D except ''O. polius'', coupled with three morphological traits—in these species, but not in ''O. polius'', the first upper molar has an additional small root at the outer (labial) side; the first lower molar has additional small roots; and the second upper molar has the mesoflexus (one of the valleys between the cusps and crests) divided in two.Weksler, 2006, p. 130 In Weksler's analysis, species placed in ''Oryzomys'' did not form a coherent ( monophyletic) group, but instead were found at various positions across the oryzomyine tree, and he suggested that most of these species, including ''O. polius'', should be placed in new genera. Later in 2006, Weksler and others described ten new genera for species formerly placed in ''Oryzomys'',Weksler et al., 2006, p. 1 including ''Eremoryzomys'' for ''polius''; thus, the species is now known as ''Eremoryzomys polius''.Weksler et al., 2006, p. 10 In reference to its "isolated distribution", they incorporated the Greek word ''eremia'' "lonely place" into the generic name.Weksler et al., 2006, p. 11 The 2008
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 ...
, citing Pacheco, commented that ''Eremoryzomys'' may in fact include more than one species. In 2011, a new oryzomyine, '' Drymoreomys albimaculatus'', was described from southeastern Brazil, and phylogenetic analysis of morphological and molecular data suggested that this animal is the closest known relative of ''Eremoryzomys''.Percequillo et al., 2011, p. 372 ''Eremoryzomys'' is now one of about 28 genera in the tribe Oryzomyini, which includes well over a hundred species distributed mainly in South America, including nearby islands such as the
Galápagos Islands The Galápagos Islands (Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuador ...
and some of the
Antilles The Antilles (; gcf, label=Antillean Creole, Antiy; es, Antillas; french: Antilles; nl, Antillen; ht, Antiy; pap, Antias; Jamaican Patois: ''Antiliiz'') is an archipelago bordered by the Caribbean Sea to the south and west, the Gulf of Mex ...
. Oryzomyini is one of several tribes recognized within the subfamily
Sigmodontinae The rodent subfamily Sigmodontinae includes New World rats and mice, with at least 376 species. Many authorities include the Neotominae and Tylomyinae as part of a larger definition of Sigmodontinae. When those genera are included, the specie ...
, which encompasses hundreds of species found across South America and into southern North America. Sigmodontinae itself is the largest subfamily of the family
Cricetidae The Cricetidae are a family of rodents in the large and complex superfamily Muroidea. It includes true hamsters, voles, lemmings, muskrats, and New World rats and mice. At almost 608 species, it is the second-largest family of mammals, and h ...
, other members of which include
vole Voles are small rodents that are relatives of lemmings and hamsters, but with a stouter body; a longer, hairy tail; a slightly rounder head; smaller eyes and ears; and differently formed molars (high-crowned with angular cusps instead of lo ...
s,
lemming A lemming is a small rodent, usually found in or near the Arctic in tundra biomes. Lemmings form the subfamily Arvicolinae (also known as Microtinae) together with voles and muskrats, which form part of the superfamily Muroidea, which also includ ...
s,
hamster Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The ...
s, and deermice, all mainly from Eurasia and North America.Musser and Carleton, 2005


Description

''Eremoryzomys polius'' is a large, long-tailed rice rat that in color resembles some North American
woodrat A pack rat or packrat, also called a woodrat or trade rat, are any species in the North and Central American rodent genus ''Neotoma''. Pack rats have a rat-like appearance, with long tails, large ears, and large, black eyes. Pack rats are notice ...
s (''
Neotoma A pack rat or packrat, also called a woodrat or trade rat, are any species in the North and Central American rodent genus ''Neotoma''. Pack rats have a rat-like appearance, with long tails, large ears, and large, black eyes. Pack rats are notice ...
''). The fur is grayish above and lighter below, where the hairs are gray at the bases but white at the tips. The external ears (
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 ...
) are short and the tail is dark above and light below. The hindfeet have well-developed
ungual tuft 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 ...
s (patches of hair) along the
plantar Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
margins and between all of the digits, a character shared only with '' Sooretamys angouya'' among oryzomyines.Weksler, 2006, p. 24 The squamae, small structures resembling scales that cover the soles of the hindfeet in many oryzomyines, are well developed. The claw of the first digit extends nearly to the end of the first
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly ...
of the second toe and the claw of the fifth toe extends slightly beyond the first phalanx of the fourth toe. As in most oryzomyines, the female has eight
mammae 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 primat ...
. Head and body length is . In Osgood's original two specimens, an old female and an adult female, tail length is , respectively; hindfoot length is ; and greatest skull length is . ''E. polius'' has 12 
thoracic The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the crea ...
, 7 or 8  lumbar, and 35 or 36  caudal vertebrae; the presence of 12 thoracic vertebrae is a putative
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
of Oryzomyini.Weksler, 2006, p. 52


Skull

In the
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 ...
, the rostrum (front part) is long and robust. The nasal bones are short, not extending further back than the
lacrimals The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the Orbit (anatomy), orbit. It has two surfaces and four borders. Several ...
, and the premaxillaries extend about as far back as the nasals. The zygomatic notch, an extension at the front of the
zygomatic plate In rodent anatomy, the zygomatic plate is a bony plate derived from the flattened front part of the zygomatic arch (cheekbone). At the back, it connects to the front (maxillary) root of the zygomatic arch, and at the top it is connected to the ...
, is present. The plate's back margin is level with the front of the first upper molar. A strong
jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anato ...
is present in the
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 zygo ...
(cheekbone), so that the maxillary and
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
bones, which form the front and back parts of the arch, respectively, do not overlap when seen from the side.Weksler et al., 2006, p. 10; Percequillo et al., 2011, p. 388. Weksler, 2006, table 5, scores ''Eremoryzomys'' (as ''Oryzomys polius'') as having overlapping squamosals and maxillaries (see character state definitions for character 30, p. 32). The narrowest part of 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 ...
(located between the eyes) is to the front and the region's margins exhibit strong beading. Various crests develop on the rounded braincase, especially in old animals. The parietal bones form part of the roof of the braincase and, unlike in some other rice rats, also extend to the sides of the braincase. The
interparietal bone An interparietal bone (os interparietale or Inca bone or ''os inca var.'') is a dermal bone situated between the parietal and supraoccipital. It is homologous to the postparietal bones of other animals. In humans, it corresponds to the upper p ...
at the back of the braincase is narrow and wedge-shaped, so that the parietal and
occipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
bones meet extensively. The incisive foramina are very long, extending well between the molars. The posterolateral palatal pits are well-developed and recessed into a fossa (depression). The bony palate is relatively short, with the mesopterygoid fossa extending forward to the end of the molar row or even between the third molars. The roof of the fossa is perforated by large sphenopalatine vacuities. Usually, an
alisphenoid strut In some rodents, the alisphenoid strut is an extension of the alisphenoid bone that separates two foramina in the skull, the masticatory–buccinator foramen and the foramen ovale accessorium. The presence or absence of this strut is variable be ...
is present; this extension of the
alisphenoid bone The greater wing of the sphenoid bone, or alisphenoid, is a bony process of the sphenoid bone; there is one on each side, extending from the side of the body of the sphenoid and curving upward, laterally, and backward. Structure The greater wi ...
separates two
foramina In anatomy and osteology, a foramen (;Entry "foramen"
in
(openings) in the skull, the masticatory–buccinator foramen and the foramen ovale accessorium. The condition of various grooves and foramina of the skull indicates that the pattern of the arterial circulation of the head is derived. The subsquamosal fenestra, an opening at the back of the skull determined by the shape of the squamosal bone, is large and the
mastoid bone The mastoid part of the temporal bone is the posterior (back) part of the temporal bone, one of the bones of the skull. Its rough surface gives attachment to various muscles (via tendons) and it has openings for blood vessels. From its borders, ...
is perforated by a fenestra (opening). The squamosal lacks a suspensory process that contacts the
tegmen tympani The tympanic cavity is a small cavity surrounding the bones of the middle ear. Within it sit the ossicles, three small bones that transmit vibrations used in the detection of sound. Structure On its lateral surface, it abuts the external auditor ...
, the roof of the
tympanic cavity The tympanic cavity is a small cavity surrounding the bones of the middle ear. Within it sit the ossicles, three small bones that transmit vibrations used in the detection of sound. Structure On its lateral surface, it abuts the external auditory ...
, a defining character of oryzomyines. In the mandible, the
mental foramen The mental foramen is one of two foramina (openings) located on the anterior surface of the mandible. It is part of the mandibular canal. It transmits the terminal branches of the inferior alveolar nerve and the mental vessels. Structure The ...
, an opening in the mandible just before the first molar, opens to the outside, not upwards as in a few other oryzomyines. The upper and lower masseteric ridges, which anchor some of the chewing muscles, usually join into a single crest at a point below the first molar and do not extend forward beyond the molar. There is no 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 ...
of the lower incisor, a trait ''Eremoryzomys'' shares with only a few other oryzomyines.


Molars

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
bunodont 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 ...
(with the cusps higher than the connecting crests) and
brachydont 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 ...
(low-crowned). On the upper first and second molar, the outer and inner valleys between the cusps and crests do not interpenetrate. Many accessory crests are present, including the mesolophs and mesolophids. The anterocone and anteroconid, the front cusps on the upper and lower first molar, are not divided into smaller outer and inner cusps. Small accessory roots are absent from the molars, so that each of the three upper molars has two roots on the outer side and one on the inner side and each of the lower molars has one root at the front and one at the back.


Distribution and status

As far as now known, ''Eremoryzomys polius'' is confined to a small area in central Peru, at an altitude of ,Percequillo et al., 2011, p. 378 but the species may range more widely. It occurs in forest in the dry lowlands of the upper parts of the basin of the
Marañón River , name_etymology = , image = Maranon.jpg , image_size = 270 , image_caption = Valley of the Marañón between Chachapoyas ( Leimebamba) and Celendín , map = Maranonrivermap.png , map_size ...
, east of the main mountain range of the Andes.Musser and Carleton, 2005, p. 1153; Pacheco et al., 2008 The biogeographical pattern indicated by the relationship between ''Eremoryzomys'' and the Brazilian ''Drymoreomys'' is unusual. While there are some similar cases of relationships between Andean and Atlantic Forest animals, these involve inhabitants of humid forests in the Andes; ''Eremoryzomys'', in contrast, lives in an arid area.Percequillo et al., 2011, p. 379 Because ''E. polius'' is so poorly known, the 2008 IUCN Red List assesses it as " Data Deficient". It is threatened by
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
for cattle pasture and is not known from any protected areas.


Notes


References


Literature cited

* * Osgood, W.H. 1913
New Peruvian mammals
Field Museum of Natural History, Zoölogical Series 10:93–100. * * *Weksler, M. 2006
Phylogenetic relationships of oryzomyine rodents (Muroidea: Sigmodontinae): separate and combined analyses of morphological and molecular data
''Bulletin of the American Museum of Natural History'' 296:1–149. *Weksler, M., Percequillo, A.R. and Voss, R.S. 2006
Ten new genera of oryzomyine rodents (Cricetidae: Sigmodontinae)
''American Museum Novitates'' 3537:1–29. {{Featured article Mammals of Peru Oryzomyini Monotypic rodent genera Mammals described in 1913 Taxa named by Marcelo Weksler Taxa named by Alexandre Reis Percequillo Taxonomy articles created by Polbot