Species
Artiodactyla
IUCN
NCBI
EOL Text
Artiodactyls are an important food source for a number of different carnivores. As artiodactyl populations decline, so too will those animals that depend on them. For example, the decline of cheetahs is often attributed habitat loss. However, cheetahs primarily prey upon small to medium sized ungulates, specifically gazelles. According to the IUCN Red List of Threatened Species, 2 species of gazelle are extinct, while 10 more are listed as vulnerable, endangered or critically endangered. In north Africa, as preferred prey species have declined, more and more cheetahs are turning to livestock for prey. Consequently, these cheetahs are then killed as pests. As a result, one of the major directives for cheetah conservation is restoration of wild prey species, most of which are small to medium-sized artiodactyls.
- Ray, J., K. Redford, R. Steneck, J. Berger. 2005. Large Carnivores and the Conservation of Biodiversity. Washington D. C.: Island Press.
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Extinction threatens nearly half of all artiodactyls and risk of extinction increases in areas with decreased economic development. Humans have hunted many species without regulation to near extinction. One of the greatest threats to artiodactyls is habitat loss. For example, the native swamp habitat of Pere David's deer was largely destroyed 3500 years ago due to the draining and cultivation. Fortunately, large herds of Pere David's deer live in numerous parks and reserves throughout their native range. In some cases, conservation efforts to increase local population growth have been so effective that population control has become necessary (e.g., Giraffa camelopardalis). In addition to habitat loss, climate change has begun to contract species ranges and forced many species move poleward. For example, moose (Alces alces), which are an important ecological component of the boreal ecosystem, are notoriously heat intolerant and are at the southern edge of their circumpolar distribution in the north central United States. Since the mid to late 1980's, demographic studies of this species have revealed sharp population declines at its southernmost distribution in response to increasing temperatures.
The IUCN Red List of Threatened Species lists 168 artiodactyl species. Seven are listed as "extinct" and two are listed as "extinct in the wild". Twenty-six species are listed as “endangered,” one is “near threatened,” and data is lacking for thirteen other species. The remaining 73 species are listed as “lower risk”. Within the United States, the U.S. Fish and Wildlife Service has listed wood bison (Bison bison athabascae), woodland caribou (Rangifer tarandus caribou), Columbian white-tailed deer (Odocoileus virginianus leucurus), key deer (Odocoileus virginianus clavium), Sonoran pronghorn (Antilocapra americana sonoriensis), Peninsular bighorn sheep (Ovis canadensis nelsoni), and Sierra Nevada bighorn sheep (Ovis canadensis sierrae) as endangered throughout at least part of their native U.S. range.
- IUCN, 2010. "Mammals" (On-line). IUCN Red List of Threatened Species. Accessed March 23, 2011 at http://www.iucnredlist.org/initiatives/mammals.
- Lenarz, M., M. Nelson, M. Schrage, A. Edwards. 2009. Temperature mediated moose survival in northeastern Minnesota. Journal of Wildlife Management, 73: 503-510.
- Murray, D., E. Cox, W. Ballard, H. Whitlaw, M. Lenarz, T. Custer, T. Barnett, T. Fuller. 2006. Pathogens, nutritional deficiency, and climate influences on a declining moose population. Wildlife Monographs, 166: 1-30.
- Price, S., J. Gittleman. 2007. "Hunting to extinction: biology and regional economy influence extinction risk and the impact of hunting in artiodactyls" (On-line). Accessed February 07, 2009 at http://journals.royalsociety.org/content/835104w3v3727236/fulltext.pdf.
- U. S. Fish and Wildlife Service, 2011. "Mammalian species report" (On-line). U.S. Fish and Wildlife Service, Endangered Species Program. Accessed March 23, 2011 at http://www.fws.gov/endangered/species/us-species.html.
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ARTIODACTYLA
The mammal order Artiodactyla (even-toed ungulates) includes ten families:
1) Camelidae (camels and relatives)
2) Suidae (pigs)
3) Tayassuidae (peccaries)
4) Hippopotamidae (hippopotamuses)
5) Tragulidae (chevrotains or mouse deer)
6) Moschidae (Musk-deer)
7) Cervidae (deer)
8) Bovidae (hollow-horned ruminants)
9) Antilocapridae (Pronghorn)
10) Giraffidae (Giraffe and Okapi)
Molecular phylogenetic studies and other evidence (some going back to the 1880s) indicate a close relationship between Artiodactyla and Cetacea (whales). The group Cetartiodactyla is a clade composed of Cetacea + Artiodactyla.
(Lewison 2011 and references therein)
- Lewison, R.L. 2011. Family Hippopotamidae (hippopotamuses). Pp. 308-319. in: Wilson, D.E. & Mittermeier, R.A., eds. Handbook of the Mammals of the World. Volume 2. Hoofed Mammals. Lynx Edicions, Barcelona.
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Various forms of zoonotic pathogens use artiodactyls during critical portions of their life or viral cycle. For example, pigs can harbor several influenza virus strains simultaneously, which can hybridize and result in new and virulent strains of influenza (e.g., H1N1). In addition, artiodactyls can transmit zoonotic diseases (e.g. Mad Cow disease) to humans through meat, milk, or direct physical contact. Artiodactyls also present a potential threat to various forms of agriculture by damaging and consuming crops, serving as a potential vector of zoonotic diseases for domestic artiodactyl populations (e.g., brucellosis), and competing with livestock for resources.
Negative Impacts: injures humans (carries human disease); crop pest; causes or carries domestic animal disease
- Pulliam, J., J. Dushoff. 2009. "Ability to Replicate in the Cytoplasm Predicts Zoonotic Transmission of Livestock Viruses" (On-line). Chicago Journals. Accessed March 11, 2009 at http://www.journals.uchicago.edu/doi/full/10.1086/596510?cookieSet=1.
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Humans and their ancestors have subsisted by hunting and gathering for the majority of their evolutionary history. Artiodactyls likely served as an important food source during a significant majority of this time and continue to be important parts of the human diet. Between 72,000 and 42,000 years ago, humans began wearing clothes, which probably included the skins of many artiodactyl species. In the near east, around 10,000 years ago, goats and sheep were domesticated for subsistence purposes, followed by the domestication of cows (7,500 years ago), pigs (7,500 years ago), llamas and alpacas (6,500 years ago), and camels (3,500 years ago). The domestication of artiodactyls for subsistence purposes lead to one of the most important cultural changes in human history, the transition from a purely hunter-gatherer society to a pastoral and agricultural societies.
Economically, cattle are the most important domesticated animal world wide. In 2001, the global population of domestic artiodactyls was greater than 4.1 billion, more than 31% of which consisted of cattle. In the United States, one of the worlds top 4 beef producers, beef production is the country's fourth largest industry. In addition to meat production, artiodactyls are used for their milk, fur, skin, bone, and feces and sport hunting generates millions of dollars in North America and Europe annually. However, trophy hunting can alter the evolutionary dynamics of wild populations by imposing unnatural selective pressures for decreased ornamentation. Finally, artiodactyls play an important role in the global ecotourism movement as various species of ungulates are readily observable throughout much of their native habitat.
Positive Impacts: pet trade ; food ; body parts are source of valuable material; ecotourism ; research and education; produces fertilizer
- Bates, D. 2005. Human Adaptive Strategies: Ecology, Culture, and Politics (Third Edition). Boston, MA: Pearson, Allyn, and Bacon.
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Artiodactyls are the most diverse, large, terrestrial mammals alive today. They are the fifth largest order of mammals, consisting of 10 families, 80 genera, and approximately 210 species. Although the majority of artiodactyls live in relatively open habitats, they can be found in all habitat types, including some aquatic systems, and are native to every continent, excluding Australia and Antarctica. As would be expected in such a diverse group, artiodactyls exhibit exceptional variation in body size and structure. Body mass ranges from 4000 kg in hippos to 2 kg in lesser Malay mouse deer. Height ranges from 5 m in giraffes to 23 cm in lesser Malay mouse deer.
Artiodactyls are paraxonic, that is, the plane of symmetry of each foot passes between the third and fourth digits. In all species, the number of digits is reduced by the loss of the first digit (i.e., pollex), and many species have second and fifth digits that are reduced in size. The third and fourth digits, however, remain large and bear weight in all artiodactyls. This pattern has earned them their name, Artiodactyla, which means "even-toed". In contrast, the plane of symmetry in perissodactyls (i.e., odd-toed ungulates) runs down the third toe. The most extreme toe reduction in artiodactyls, living or extinct, can be seen in antelope and deer, which have just two functional (weight-bearing) digits on each foot. In these animals, the third and fourth metapodials fuse, partially or completely, to form a single bone called a cannon bone. In the hind limb of these species, the bones of the ankle are also reduced in number, and the astragalus becomes the main weight-bearing bone. These traits are probably adaptations for running fast and efficiently.
Artiodactyls are divided into 3 suborders. Suiformes includes the suids, tayassuids and hippos, including a number of extinct families. These animals do not ruminate (chew their cud) and their stomachs may be simple and one-chambered or have up to three chambers. Their feet are usually 4-toed (but at least slightly paraxonic). They have bunodont cheek teeth, and canines are present and tusk-like. The suborder Tylopoda contains a single living family, Camelidae. Modern tylopods have a 3-chambered, ruminating stomach. Their third and fourth metapodials are fused near the body but separate distally, forming a Y-shaped cannon bone. The navicular and cuboid bones of the ankle are not fused, a primitive condition that separates tylopods from the third suborder, Ruminantia. This last suborder includes the families Tragulidae, Giraffidae, Cervidae, Moschidae, Antilocapridae, and Bovidae, as well as a number of extinct groups. In addition to having fused naviculars and cuboids, this suborder is characterized by a series of traits including missing upper incisors, often (but not always) reduced or absent upper canines, selenodont cheek teeth, a 3 or 4-chambered stomach, and third and fourth metapodials that are often partially or completely fused.
- Feldhamer, G., L. Drickamer, S. Vessey, J. Merritt. 2004. Mammalogy: Adaptation, Diversity, Ecology. New York: McGraw Hill.
- Grzimek, B. 2003. Artiodactyla (Even-toed ungulates). Pp. 263-417 in M Hutchins, D Kleiman, V Geist, M McDade, eds. Grzimek's Animal Life Encyclopedia, Vol. 15, Mammals IV, 2nd Edition. Farmington Hills, Michigan, USA: Gale Group.
- Grzimek, B. 1990. Artiodactyla. Pp. 1-639 in S Parker, ed. Grzimek’s Encyclopedia of Mammals, Vol. 5, 1st Edition. New York: McGraw-Hill.
- Nowak, R. 1999. Walker’s Mammals of the World. Baltimore and London: The Johns Hopkins University Press.
- Savage, R., M. Long. 1986. Mammal Evolution, an Illustrated Guide. New York: Facts of File Publications.
- Simpson, C. 1984. Artiodactyls. Pp. 686 in S Anderson, J Jones, Jr., eds. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons.
- Vaughn, T., J. Ryan, N. Czaplewski. 2000. Mammalogy, Fourth Edition. Fort Worth: Brooks/Cole.
- Wilson, D., D. Reeder. 1993.
Mammal Species of the World, A Taxonomic and Geographic Reference. 2nd edition
. Washington D. C.: Smithsonian Institution Press.
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Artiodactyls are distributed nearly worldwide and are native to all continents except Antarctica and Australia. Numerous introductions, consisting mainly of domestic species, have occurred in areas outside their normal range. Where introduced in areas with suitable forage, artiodactyls usually thrive.
Biogeographic Regions: nearctic (Introduced , Native ); palearctic (Introduced , Native ); oriental (Introduced , Native ); ethiopian (Introduced , Native ); neotropical (Introduced , Native ); australian (Introduced ); oceanic islands (Introduced )
Other Geographic Terms: cosmopolitan
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In artiodactyls, the structure of the foot is especially diagnostic, specifically the number of toes and the morphology of the astragalus. Most species have either 2 or 4 toes on each foot (for exceptions see Pecari and Tayassu) as the first digit, present in most ancestral mammals, has been lost through evolution and the second and fifth digits have been significantly reduced. As a result, artiodactyls are paraxonic. The unique structure of the astragalus, which consists of a "double-pulley" arrangement of the articular surfaces, completely restricts lateral motion and allows for greater flexion and extension of the hind limb. The astragalus, in conjunction with springing ligaments in the limbs, hard hooves, relatively small feet, and elongated lightweight limbs, allows for highly developed cursorial locomotion in more derived species. In the families Camelidae, Cervidae, Giraffidae, Antilocapridae, and Bovidae, the third and fourth metapodials have become fused to create the cannon bone, which serves as the insertion point for the springing ligament in each of the four limbs. Throughout all of Artiodactyla, the range of fusion between the third and fourth metapodials varies from none to complete. Finally, residents of sandy or snowy habitats often have splayed toes, which distributes an individual's weight over a greater surface area, thereby decreasing movement costs in more fluid terrestrial substrates.
Although exceptions exist (pigs and peccaries), the vast majority of artiodactyls are obligate herbivores, consisting of browsers, grazers and mixed feeders. Although plants provide an abundant and diverse food source, mammals do not possess the enzymes necessary to break down cellulose or lignin. As a result, most artiodactyls rely on microorganisms to help break down these plant compounds. In addition to their true stomach, all artiodactyls have at least one additional chamber in which bacterial fermentation occurs. This chamber, or "false stomach", is located just before the true stomach along the gastrointestinal tract. Cervids and bovids have three false stomachs, hippos, camels, and tragulids have two, while pigs and peccaries have only one small chamber.
A majority of artiodactyls having selenodont cheek teeth, however, many species also exhibit lophodont tooth morphology. In general, browsers tend to have brachydont teeth (i.e., low crowned) while grazers have hypsodont teeth (i.e., high crowned). Within Artiodactyla, the families Suidae (pigs) and Tayassuidae (peccaries) are omnivores and have quadrate, bunodont teeth. Often, a diastema occurs between the canine and first premolar, which is especially prevalent in the lower jaw. Bovidae, Cervidae, and Giraffidae have lost their upper incisors, and several groups have lost their upper canines. However, many have retained their incisors (pigs, peccaries, hippos, and camels) and some have developed them as weapons or indicators of mate quality (some suids, cervids and musk deer). While most families have incisiform lower canines, pigs, peccaries, hippos, and camels have conically shaped canines.
Artiodactyls exhibit a great deal of variation in physical appearance. Body mass ranges from 4000 kg in hippos to 2 kg in lesser Malay mouse deer. Height ranges from 5 m in giraffes to 23 cm in lesser Malay mouse deer. Most artiodactyls have laterally positioned eyes, often with long eyelashes. They commonly have rotating ears that are round or pointed at the tips and are relatively large in relation to skull size. Most artiodactyls also have elongated and powerful legs. Many families have horns, antlers, or tusks. Horns, always consisting of bone or having a bony core, are common in many families and most often stem from the frontals which are usually larger than the parietals. Similar to horns, antlers arise from the base of the frontals and are entirely bony. Unlike horns, however, antlers are deciduous and used during the breeding season. Horns and antlers are often used in ritualized social interactions, such as male-male competition within species.
The pelage of artiodactyls typically consists of guard hair and under fur, which together help control heat exchange. Under fur tends to be short and fine and is efficient at trapping heat. Guard hairs are longer and more stout than underfur and act as a barrier against wind, rain, and snow. Pelage color varies from black to white with many shades of brown. Color patterns within the pelage vary from spots to stripes, while most young have distinctly different coats than adults. In some species, males have a ventral ridge of long hairs referred to as a ruff or dewlap and male coat color is often linked to age or social status. Species living in temperate and arctic regions shed their winter coats on a seasonal basis.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: sexes alike; female larger; male larger; sexes shaped differently; ornamentation
- Eisenberg, J. 1983. The Mammalian Radiations: An Analysis of Trends in Evolution, Adaptation, and Behavior. Chicago, IL: The University of Chicago Press.
- Rose, K., D. Archibald. 2005. The Rise of Placental Mammals. Baltimore, Maryland: The John Hopkins University Press.
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Artiodactyls are exceptionally diverse and globally distributed. Consequently, they inhabit a broad range of habitat types and can be found anywhere sufficient forage exists. Although artiodactyls occur from deserts to tropical forests to tundra, preferred habitat types fall into four major categories, which are linked to forage abundance and predator defense. Open grasslands provide abundant forage while allowing for early detection of approaching predators. Grasslands or meadows near steep cliffs provide forage while offering safety from potential predators in adjacent rocky ledges and steep terrain. Forests and shrublands provide abundant forage while offering cover from potential predators in dense vegetation. Finally, many species inhabit the ecotone between open areas and forests. While open areas provide abundant forage, adjacent forests provide dense cover from potential predators. Habitat-use patterns in artiodactyls are often linked with body size and taxonomy, with small to medium-sized artiodactyls found mainly in habitats with tall, dense vegetation. Most goat and sheep species (Caprinae) are found in open habitats adjacent to rocky cliffs, where they are specialized for navigating uneven terrain.
Habitat Regions: temperate ; tropical ; polar ; terrestrial
Terrestrial Biomes: tundra ; taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains
Wetlands: marsh ; swamp ; bog
Other Habitat Features: urban ; suburban ; agricultural ; riparian
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