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Oryzomyini is a tribe of in the subfamily of the family . 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 South America, including many offshore islands. It is part of the Oryzomyalia, which includes most of the South American Sigmodontinae.

The name Oryzomyini derives from that of its , , which means "rice rat" or "rice mouse". Many species are also known as rice rats.


Taxonomy

Contents of Oryzomyini
An oryzomyine group was first envisaged by in the early 20th century. He defined it to include pentalophodont species, which have a (id) on the upper and lower molars, with a long (extending past the third molars). Thomas included , , and (which included many species now in other genera), as well as , which is currently classified in the related tribe Thomasomyini instead. In 1944, Hershkovitz proposed a more extended definition of the group, excluding Rhagomys, but including (then including ), , and . Some subsequent authors did not separate the oryzomyines from the thomasomyines, which were distinguished from them by having a short palate, including Vorontsov, who in 1959 was the first to use Oryzomyini as a formal family-group name. He included most current oryzomyines as well as the thomasomyines and , which are now known to be more distantly related.Weksler, 2006, p. 4

The genera (including at the time), , and were not included at the time because of their tetralophodont molars (lacking complete mesoloph(id)s); instead, Holochilus was considered to be a sigmodont, related to and , and Pseudoryzomys and Zygodontomys were considered to be members of Phyllotini, another large South American tribe. Although lacking mesoloph(id)s, these genera share other characters with oryzomyines, and a series of papers by Robert Voss and coworkers in the early 1990s established their membership in Oryzomyini.Weksler, 2006, p. 5

In a 1993 paper, Voss and Carleton proposed the first diagnosis of Oryzomyini. They included twelve genera and proposed five for the tribe: presence of a pair of on the chest; a long palate marked by posterolateral palatal pits, perforations near the third molar; absence of an alisphenoid strut, which in some sigmodontines separates two foramina (openings) in the skull; absence of a suspensory process of the bone attached to the roof of the , the ; and absence of a .Voss and Carleton, 1993, p. 31 Some of these have been reversed in some oryzomyines; for example, an alisphenoid strut is present in several oryzomyines, including .Weksler, 2006, p. 38

The contents of Oryzomyini have been largely stable since,Musser and Carleton, 2005; Weksler, 2006; Weksler et al., 2006 but the allocation of some animals has been contentious. , an extinct giant rat from the Galápagos Islands, has been allocated to both Oryzomyini and Thomasomyini, but its correct classification currently remains unclear.Musser and Carleton, 2005 The genus has been excluded from Oryzomyini on the basis of studies of the mitochondrial gene,Musser and Carleton, 2005, p. 1117 but the IRBP gene provides evidence for its membership in Oryzomyini.Weksler, 2006, p. 124 Handleyomys fuscatus was placed in the thomasomyine genus before its close relationship to H. intectus was recognized in 2002.Voss et al., 2002, p. 2 was described as a transitional form between oryzomyines and , but later allied with OryzomyiniMusser and Carleton, 2005, p. 1126 and even summarily dismissed as an anomalous .Weksler et al., 2006, table 1, footnote d

In the early 2000s, the advent of molecular phylogenetics has led to progress in the understanding of the relationships of oryzomyines. They are currently classified in the family , which includes the , , , and many other species, chiefly in the Americas and Eurasia. Within this family, they are part of the subfamily , which is mainly distributed in South America but also extends into southern North America. Sigmodontinae includes several tribes, most of which cluster into a now known as Oryzomyalia, which includes Oryzomyini, Akodontini, Phyllotini, Thomasomyini, and other, smaller groups, but not the ( Sigmodon) and the .Steppan et al., 2004


Internal classification
The relationships among the oryzomyine genera have long been obscure, although several studies provided insights into the relations of some genera.Weksler, 2006, p. 7 The single most significant problem in oryzomyine taxonomy has been the definition of the , ,Weksler, 2006, p. 10 which in one classification included all animals then recognized as oryzomyines.Ray, 1962 Many groups were later excluded from the genus, but even so it included forty species that did not form a group.Weksler, 2006, p. 10, table 2

In the 2000s, Marcelo Weksler published several studies in which he used evidence from IRBP, a nuclear gene, and morphology to assess the relationships among the members of Oryzomyini. He provided support for several intergeneric relationships and clarified the scale of the Oryzomys problem, as species of Oryzomys appeared in about ten separate .Weksler, 2006, figs. 34–39 In a 2006 publication, he and coworkers described ten new genera for species previously placed in Oryzomys and transferred some others to Handleyomys, leaving only about six species in Oryzomys.

Weksler's analyses suggested that oryzomyines fall into four major clades, which were largely congruent across his analyses of morphology and IRBP, but support for all of those was limited and the placement of some genera remained unclear. He dubbed these clades "clade A" through "clade D". Some analyses supported a relationship between clades C and D, which in turn were related to clade B, with clade A at a basal position, but other analyses could not resolve the relationships among the major clades. The four clades are as follows:

  • Clade A includes only and , but support for a relationship between those morphologically and dissimilar genera was not strong.Weksler, 2006, p. 75
  • Clade B includes at least , , , , , and , the latter four of which were included in Oryzomys until 2006.Weksler et al., 2006 and were also recovered in the clade in some analyses,Weksler, 2006, figs. 34, 35, 38 but the former, for which most morphological characters are unknown, is more likely related to Nectomys within clade DWeksler, 2006, p. 133 and the relations of the latter, a poorly known yet unique animal, are obscure as some analyses suggest it is a basal member of Oryzomyini.Weksler et al., 2006, p. 17 There is little support for all intergeneric relationships within the clade.
  • Clade C includes , , , and , which was placed in Oryzomys until 2006. No intergeneric relationships within this clade received substantial support.
  • Clade D includes ,Percequillo et al., 2011, p. 372 , , , , , , , , , , , , most likely , and the extinct ,Weksler et al., 2006, p. 18 ,Zijlstra et al., 2010, p. 869 and .Turvey et al., 2010, p. 766 Eremoryzomys, Cerradomys, Sooretamys, and Aegialomys were included in Oryzomys before 2006. There is some support for a basal placement of Eremoryzomys within this clade and for two major groups of related genera, one including Holochilus, Lundomys, and Pseudoryzomys and the other Nectomys, Amphinectomys, Sigmodontomys, Melanomys, Aegialomys, and Nesoryzomys,Weksler, 2006, figs. 37–39 with the extinct Megalomys and Pennatomys.Turvey et al., 2010, pp. 759–760 The HolochilusLundomysPseudoryzomys group also includes the extinct and .Pardiñas, 2008 Oryzomys may be its closest relative, but it is possible that this arrangement is the result of a convergent development of adaptations to a life in the water in the two groups.Weksler, 2006, p. 131 Within the group of Nectomys and related genera, there is some support for a core group that excludes Aegialomys and Nesoryzomys and for two subclades within the core group, one including Nectomys and Amphinectomys and the other Melanomys and Sigmodontomys.

The affinities of some species remain unclear. Many oryzomyines are known from the , including " " and species of Megalomys and Oligoryzomys, but most remain undescribed.Turvey, 2009


Description
Most oryzomyines are nondescript rodents that look like common and , but the tribe also includes some spectacularly specialized forms. The smallest members, mainly in clade C, may have a head-body length of as little as and mass of , but the largest living oryzomyine, Nectomys, reaches head and body lengths over and mass of about ; Lundomys and Holochilus are only a little smaller.Weksler, 2006, p. 3, table 8 Some of the extinct species from the Lesser Antilles, such as " Ekbletomys hypenemus" and Megalomys desmarestii, were even larger.Ray, 1962, tables 7, 11


Distribution and ecology
Oryzomyines range from in the north, where the Marsh Rice Rat ( Oryzomys palustris) is found, to Tierra del Fuego in the south, where Oligoryzomys magellanicus occurs. Extinct species are known from ( Oryzomys antillarum), the Galápagos Islands ( Nesoryzomys and Aegialomys galapagoensis), Fernando de Noronha ( Noronhomys), and the north to ( Megalomys, Oligoryzomys victus, and several unidentified genera). They are abundant in many environments, from rainforest to grassland.Weksler, 2006, p. 3 Most live in the forest, but Zygodontomys, Lundomys, Pseudoryzomys, Aegialomys, and Nesoryzomys live exclusively in open vegetation and some other genera include both forest and non-forest forms.Weksler, 2006, pp. 81–82 Most oryzomyines are relatively unspecialized animals that live on the ground, but Oecomys is specialized to and various members of clade D, including Holochilus, Oryzomys, and Nectomys, are , spending at least some of their time in the water.Weksler, 2006, pp. 78–79


Literature cited

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