Rodent

Rodents Temporal range: Early Paleocene–Recent PreЄ Є O S D C P T J K Pg N
Indian Palm Squirrel (Funambulus palmarum)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Infraclass:	Eutheria
Magnorder:	Boreoeutheria
Superorder:	Euarchontoglires
Order:	Rodentia Bowdich, 1821
Suborders
Sciuromorpha Castorimorpha Myomorpha Anomaluromorpha Hystricomorpha
Combined range of all rodent species

Rodentia is an order of mammals also known as rodents, characterised by two continuously growing incisors in the upper and lower jaws which must be kept short by gnawing.^[1][2]

Forty percent of mammal species are rodents, and they are found in vast numbers on all continents other than Antarctica. Common rodents include mice, rats, squirrels, gerbils, porcupines, beavers, chipmunks, guinea pigs, hamsters, and voles.^[1] Rodents have sharp incisors that they use to gnaw wood, break into food, and bite predators. Most eat seeds or plants, though some have more varied diets. Some species have historically been pests, eating seeds stored by people^[3] and spreading disease.^[4]

The name comes from the Latin word rodens, "gnawing one" (from the verb rodere, "gnaw").

Size and range of order

In terms of number of species—although not necessarily in terms of number of organisms (population) or biomass—rodents make up the largest order of mammals. There are about 2,277 species of rodents (Wilson and Reeder, 2005), with over 40 percent of mammalian species belonging to the order.^[5] Their success is probably due to their small size, short breeding cycle, and ability to gnaw and eat a wide variety of foods. (Lambert, 2000)

Rodents are found in vast numbers on all continents except Antarctica, most islands, and in all habitats except oceans. They are the only non-volant, non-marine placental order—and in particular are the only placental order besides bats (Chiroptera) and Pinnipeds—to have reached Australia without human introduction.

Characteristics

The capybara, the largest living rodent, can weigh up to 65 kg (140 lb).

Many rodents are small; the tiny African pygmy mouse can be as little as 6 cm (2.4 in) in length and 7 g (0.25 oz) in weight at maturity, and the Baluchistan Pygmy Jerboa is of roughly similar or slightly smaller dimensions. On the other hand, the capybara can weigh up to 80 kg (180 lb),^[6] and the largest known rodent, the extinct Josephoartigasia monesi, is estimated to have weighed about 1,000 kg (2,200 lb), and possibly up to 1,534 kg (3,380 lb)^[7] or 2,586 kg (5,700 lb).^[8]

Rodents have two incisors in the upper as well as in the lower jaw which grow continuously and must be kept worn down by gnawing; this is the origin of the name, from the Latin rodere, to gnaw.^[9] These teeth are used for cutting wood, biting through the skin of fruit, or for defense. The teeth have enamel on the outside^[ambiguous] and exposed dentine on the inside^[ambiguous], so they self-sharpen during gnawing. Rodents lack canines, and have a space (diastema) between their incisors and premolars. Nearly all rodents feed on plants, seeds in particular, but there are a few exceptions which eat insects or fish. Some squirrels are known to eat passerine birds like cardinals and blue jays.

Typical rodent tooth system

Rodents are important in many ecosystems because they reproduce rapidly, and can function as food sources for predators, mechanisms for seed dispersal, and as disease vectors. Humans use rodents as a source of fur, as pets, as model organisms in animal testing, for food, and even for detecting landmines.^[10]

Members of non-rodent orders such as Chiroptera (bats), Scandentia (treeshrews), Soricomorpha (shrews and moles), Lagomorpha (hares, rabbits and pikas) and mustelid carnivores such as weasels and mink are sometimes confused with rodents.

Evolution

Masillamys sp. fossil from the Messel Pit fossil site

The fossil record of rodent-like mammals begins shortly after the extinction of the non-avian dinosaurs 65 million years ago, as early as the Paleocene. Some molecular clock data, however, suggest that modern rodents (members of the order Rodentia) already appeared in the late Cretaceous, although other molecular divergence estimations are in agreement with the fossil record.^[11][12] By the end of the Eocene epoch, relatives of beavers, dormice, squirrels, and other groups appeared in the fossil record. They originated in Laurasia, the supercontinent composed of today's North America, Europe, and Asia. Some species colonized Africa, giving rise to the earliest hystricognaths. From Africa hystricognaths rafted to South America, an isolated continent during the Oligocene and Miocene epochs. By the Miocene, Africa collided with Asia, allowing rodents such as porcupines to spread into Eurasia. During the Pliocene, rodent fossils appeared in Australia. Although marsupials are the most prominent mammals in Australia, rodents now make up almost 25% of the continent's mammal species. Meanwhile, the Americas became joined by the Isthmus of Panama and some rodents participated in the resulting Great American Interchange; sigmodontines surged southward and caviomorphs headed north.

Some prehistoric rodents

Castoroides, a giant beaver

Ceratogaulus, a horned burrowing rodent

Spelaeomys, a rat that grew to a large size on the island of Flores

Giant hutias, a group of rodents once found in the West Indies

Ischyromys, a primitive squirrel-like rodent

Leithia, a giant dormouse

Neochoerus pinckneyi, a large North American capybara that weighed 100 kg (220 lb)

Josephoartigasia monesi, the largest known rodent, with an estimated weight of very roughly 1,000 kg (2,200 lb)

Phoberomys pattersoni, the second largest known rodent, with an estimated weight of 700 kg (1,500 lb)

Telicomys, another giant South American rodent

Classification

2/3 of rodent species are in the superfamily Muroidea (rats, mice, and related species). The families Muridae (blue) and Cricetidae (red) make up the bulk of the Muroidea.

Standard classification

The rodents are part of the clades Glires (along with lagomorphs), Euarchontoglires (along with lagomorphs, primates, treeshrews, and colugos), and Boreoeutheria (along with most other placental mammals). The order Rodentia may be divided into suborders, infraorders, superfamilies and families.

Classification scheme:

ORDER RODENTIA (from Latin, rodere, to gnaw)

• Suborder Anomaluromorpha

  • Family Anomaluridae: scaly-tailed squirrels
  • Family Pedetidae: springhares

• Suborder Castorimorpha

  • Superfamily Castoroidea
    • Family Castoridae: beavers
  • Superfamily Geomyoidea
    • Family Geomyidae: pocket gophers (true gophers)
    • Family Heteromyidae: kangaroo rats and kangaroo mice

• Suborder Hystricomorpha

  • Family incertae sedis Diatomyidae: Laotian rock rat

  • Infraorder Ctenodactylomorphi

    • Family Ctenodactylidae: gundis

  • Infraorder Hystricognathi

    • Family Bathyergidae: African mole rats
    • Family Hystricidae: Old World porcupines
    • Family Petromuridae: dassie rat
    • Family Thryonomyidae: cane rats

    • Parvorder Caviomorpha
      • Family †Heptaxodontidae: giant hutias
      • Family Abrocomidae: chinchilla rats
      • Family Capromyidae: hutias
      • Family Caviidae: cavies, including guinea pigs and the capybara
      • Family Chinchillidae: chinchillas and viscachas
      • Family Ctenomyidae: tuco-tucos
      • Family Dasyproctidae: agoutis
      • Family Cuniculidae: pacas
      • Family Dinomyidae: pacaranas
      • Family Echimyidae: spiny rats
      • Family Erethizontidae: New World porcupines
      • Family Myocastoridae: nutria, coypu
      • Family Octodontidae: octodonts
• Suborder Myomorpha

  • Superfamily Dipodoidea
    • Family Dipodidae: jerboas and jumping mice
  • Superfamily Muroidea
    • Family Calomyscidae: mouse-like hamsters
    • Family Cricetidae: hamsters, New World rats and mice, voles
    • Family Muridae: true mice and rats, gerbils, spiny mice, crested rat
    • Family Nesomyidae: climbing mice, rock mice, white-tailed rat, Malagasy rats and mice
    • Family Platacanthomyidae: spiny dormice
    • Family Spalacidae: mole rats, bamboo rats, and zokors

• Suborder Sciuromorpha

  • Family Aplodontiidae: mountain beaver
  • Family Gliridae (also Myoxidae, Muscardinidae): dormice
  • Family Sciuridae: squirrels, including chipmunks, prairie dogs, & marmots

Alternative classifications

The above taxonomy uses the shape of the lower jaw (sciurognath or hystricognath) as the primary character. This is the most commonly used approach for dividing the order into suborders. Many older references emphasize the zygomasseteric system (suborders Protrogomorpha, Sciuromorpha, Hystricomorpha, and Myomorpha).

Several molecular phylogenetic studies have used gene sequences to determine the relationships among rodents, but these studies are yet to produce a single consistent and well-supported taxonomy. Some clades have been consistently produced such as:

• Ctenohystrica contains:
  • Ctenodactylidae (gundis)
  • Hystricognathi containing:
    • Hystricidae
    • An unnamed clade containing:
      • Phiomorpha
      • Caviomorpha

• An unnamed clade contains:
  • Gliridae
  • Sciuroidea containing:
    • Aplodontiidae
    • Sciuridae

• Myodonta includes:
  • Dipodoidea
  • Muroidea

The positions of the Castoridae, Geomyoidea, Anomaluridae, and Pedetidae are still being debated.

Monophyly or polyphyly?

In 1991, a paper submitted to Nature proposed that caviomorphs should be reclassified as a separate order (similar to Lagomorpha), based on an analysis of the amino acid sequences of guinea pigs.^[13] This hypothesis was refined in a 1992 paper, which asserted the possibility that caviomorphs may have diverged from myomorphs prior to later divergences of Myomorpha; this would mean caviomorphs, or possibly hystricomorphs, would be moved out of the rodent classification into a separate order.^[14] A minority scientific opinion briefly emerged arguing that guinea pigs, degus, and other caviomorphs are not rodents,^[15][16] while several papers were put forward in support of rodent monophyly.^[17][18][19] Subsequent studies published since 2002, using wider taxon and gene samples, have restored consensus among mammalian biologists that the order Rodentia is monophyletic.^[20][21]

References

1. ^ ^{a b} "rodent - Encyclopedia.com". http://www.encyclopedia.com/doc/1E1-rodent.html. Retrieved 2007-11-03. 
2. ^ "Rodents: Gnawing Animals". http://www.kidport.com/RefLib/Science/Animals/Rodents.htm. Retrieved 2007-11-03. 
3. ^ Meerburg BG, Singleton GR, Leirs H (2009). "The Year of the Rat ends: time to fight hunger!". Pest Manag Sci 65 (4): 351–2. doi:10.1002/ps.1718. PMID 19206089. http://www3.interscience.wiley.com/journal/121686000/abstract. 
4. ^ Meerburg BG, Singleton GR, Kijlstra A (2009). "Rodent-borne diseases and their risks for public health". Crit Rev Microbiol 35 (3): 221–70. doi:10.1080/10408410902989837. PMID 19548807. http://www.informahealthcare.com/doi/pdf/10.1080/10408410902989837. 
5. ^ Myers, Phil (2000). "Rodentia". Animal Diversity Web. University of Michigan Museum of Zoology. http://animaldiversity.ummz.umich.edu/site/accounts/information/Rodentia.html. Retrieved 2006-05-25. 
6. ^ Capybaras (Hydrochaeridae): Information and Much More from Answers.com
7. ^ Millien, Virginie (05 2008). "The largest among the smallest: the body mass of the giant rodent Josephoartigasia monesi". Proceedings of the Royal Society B 1 (1646): -1. doi:10.1098/rspb.2008.0087. PMC 2596365. PMID 18495621. http://journals.royalsociety.org/content/pg31525230323q27/?p=35f8c90fe97d44c9b32766e547837566&pi=0. Retrieved 2008-05-27. 
8. ^ Rinderknecht, Andrés; Blanco, R. Ernesto (01 2008). "The largest fossil rodent" (pdf). Proceedings of the Royal Society B 275 (1637): 923–928. doi:10.1098/rspb.2007.1645. PMC 2599941. PMID 18198140. http://journals.royalsociety.org/content/34j867846u164624/fulltext.pdf. Retrieved 2008-05-27. 
9. ^ Pearsall, J., ed (2002). The Concise Oxford English Dictionary, 10th ed. rev.. Oxford: Oxford University Press. p. 1,239. ISBN 0-19-860572-2. 
10. ^ Wines, Michael (2004-05-19). "Gambian rodents risk death for bananas". The Age (The Age Company Ltd.). http://www.theage.com.au/articles/2004/05/18/1084783512636.html. Retrieved 2006-05-25.  "A rat with a nose for landmines is doing its bit for humanity" Cited as coming from the New York Times in the article.
11. ^ Douzery, E.J.P., F. Delsuc, M.J. Stanhope, and D. Huchon (2003). "Local molecular clocks in three nuclear genes: divergence times for rodents and other mammals and incompatibility among fossil calibrations". Journal of Molecular Evolution 57: S201. doi:10.1007/s00239-003-0028-x. PMID 15008417. 
12. ^ Horner, D.S., K. Lefkimmiatis, A. Reyes, C. Gissi, C. Saccone, and G. Pesole (2007). "Phylogenetic analyses of complete mitochondrial genome sequences suggest a basal divergence of the enigmatic rodent Anomalurus". BMC Evolutionary Biology 7: 16. doi:10.1186/1471-2148-7-16. PMC 1802082. PMID 17288612. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=1802082. 
13. ^ Graur, D.; Hide, W.; Li, W. (1991). "'Is the guinea-pig a rodent?'". Nature 351 (6328): 649–652. doi:10.1038/351649a0. PMID 2052090. 
14. ^ Li, W.; Hide, W.; Zharkikh, A.; Ma, D.; Graur, D. (1992). "'The molecular taxonomy and evolution of the guinea pig.'". Journal of Heredity 83 (3): 174–81. PMID 1624762. 
15. ^ D'Erchia, A.; Gissi, C.; Pesole, G.; Saccone, C.; Arnason, U. (1996). "'The guinea-pig is not a rodent.'". Nature 381 (6583): 597–600. doi:10.1038/381597a0. PMID 8637593. 
16. ^ Reyes, A.; Pesole, G.; Saccone, C. (2000). "'Long-branch attraction phenomenon and the impact of among-site rate variation on rodent phylogeny.'". Gene 259 (1-2): 177–87. doi:10.1016/S0378-1119(00)00438-8. PMID 11163975. 
17. ^ Cao, Y.; Adachi, J.; Yano, T.; Hasegawa, M. (1994). "'Phylogenetic place of guinea pigs: No support of the rodent-polyphyly hypothesis from maximum-likelihood analyses of multiple protein sequences.'". Molecular Biology and Evolution 11 (4): 593–604. PMID 8078399. 
18. ^ Kuma, K.; Miyata, T. (1994). "'Mammalian phylogeny inferred from multiple protein data.'". Japanese Journal of Genetics 69 (5): 555–66. doi:10.1266/jjg.69.555. PMID 7999372. 
19. ^ Robinson-Rechavi, M.; Ponger, L.; Mouchiroud, D. (2000). "'Nuclear gene LCAT supports rodent monophyly.'". Molecular Biology and Evolution 17 (9): 1410–1412. PMID 10960041. 
20. ^ Lin, Y-H; McLenachan, PA; Gore, AR; Phillips, MJ; Ota, R; Hendy, MD; Penny, D (2002). "Four new mitochondrial genomes and the increased stability of evolutionary trees of mammals from improved taxon sampling". Molecular Biology and Evolution 19 (12): 2060–2070. PMID 12446798. 
21. ^ Carleton, Michael D., and Musser, Guy G. "Order Rodentia". Mammal Species of the World, 3rd edition, 2005, vol. 2, p. 745. (Concise overview of the literature)

Further reading

• Adkins RM, Gelke R. M. E. L., Rowe D., Honeycutt R. L. (2001). "Molecular phylogeny and divergence time estimates for major rodent groups: Evidence from multiple genes". Molecular Biology and Evolution 18 (5): 777–791. PMID 11319262. 
• Carleton, M. D. and G. G. Musser. 2005. "Order Rodentia," pp. 745–752 in Mammal Species of the World: A Taxonomic and Geographic Reference. Johns Hopkins University Press, Baltimore.
• David Lambert and the Diagram Group. The Field Guide to Prehistoric Life. New York: Facts on File Publications, 1985. ISBN 0-8160-1125-7
• Leung, LKP; Cox, Peter G.; Jahn, Gary C.; Nugent, Robert (2002). "Evaluating rodent management with Cambodian rice farmers". Cambodian Journal of Agriculture 5: 21–26. 
• McKenna, Malcolm C., and Bell, Susan K. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, 631 pp. ISBN 0-231-11013-8
• Nowak, R. M. 1999. Walker's Mammals of the World, Vol. 2. Johns Hopkins University Press, London.
• Steppan, S. J.; Adkins, R. A.; Anderson, J. (2004). "Phylogeny and divergence date estimates of rapid radiations in muroid rodents based on multiple nuclear genes". Systematic Biology 53: 533–553. doi:10.1080/10635150490468701. PMID 15371245. 
• University of California Museum of Paleontology (UCMP). 2007 "Rodentia". [1]
• Wilson, D. E. and D. M. Reeder, eds. 2005. Mammal Species of the World: A Taxonomic and Geographic Reference. Johns Hopkins University Press, Baltimore.

External links

• Website on African rodentia
• Rodent Photos
• Rodent Pests chapter in United States Environmental Protection Agency‎ and University of Florida/Institute of Food and Agricultural Sciences National Public Health Pesticide Applicator Training Manual
• Can Rodents be trained not to gnaw?