Diversity of Living Camelids
The Camelidae (camels and relatives) are sufficiently distinct from other families in the mammal order Artiodactyla that they are placed in their own suborder, Tylopoda ("padded foot"). This suborder includes only the family Camelidae, which is divided into two tribes, the Camelini and Lamini. The Camelini includes just two living representatives, the Bactrian Camel (Camelus bactrianus) and the Dromedary (Camelus dromedarius). The Lamini includes four species: Guanaco (Lama guanicoe), Llama (Lama glama), Viçuna (Vicugna vicugna), and Alpaca (Vicugna pacos).
The earliest camelids evolved in North America, where they flourished for forty to fifty million years, with the last known North American camelids disappearing only around 10,000 years ago. Six to seven million years ago, camels that would give rise to the Dromedary and Bactrian Camel spread into Asia, across Europe, and as far west as Spain. Three to four million years ago, camelids that would give rise to the current day South American species spread south across the Panamanian Land Bridge. Because of their their ability to thrive under tough conditions of extreme temperature and scarce food and water, domesticated camelids have been extremely important to the development of human cultures in the steppes of Eurasia, the deserts of Africa, and the arid Andes of South America.
All camelids are diurnal and are adapted to harsh and dry climates. They are all highly social.
Six living species of camelids are recognized by Franklin (2011), three of which are domesticated forms.
1) Bactrian Camel (Camelus bactrianus). The Bactrian Camel is associated with Central Asia. The wild form (known to science only since 1878) is critically endangered in its remote Gobi Desert homeland, but the domesticated form is found widely in Asia. The relationship between the wild and domesticated forms remains unclear. Bactrian Camels were domesticated 4,000-6,000 years ago. Some authorities recognize the wild form as a distinct species, Camelus ferus, in part based on preliminary genetic evidence that the wild and domestic "Bactrian Camel" lineages actually diverged hundereds of thousands of years ago.
2) Dromedary (Camelus dromedarius). This one-humped camel is associated with northern Africa and southwestern Asia. This camel is entirely domesticated (except for a free-ranging feral population in central Australia that was introduced in the late 1800s). The Dromedary was domesticated around 4000-5000 years ago; the wild form is believed to have gone extinct by 2000-5000 years ago. The Dromedary's hump contains a large fat reservoir that sustains it through times of scarcity. When the animal is well fed, the hump is plump and erect, but in starved animals it shrinks or flops to one side. Even when food is plentiful, Dromedaries may travel 30 km/day. A range of domesticated Dromedary varieties have been bred as pack or riding animals or for milk or meat. In the wet season, when lush, green plants are available, Dromedaries can go eight months without drinking. They can tolerate water loss greater than 30% of body weight (half this loss is fatal for most mammals).
3) Guanaco (Lama guanicoe). The heart of this species' range is Patagonia, in southern South America, but this is the most widely distributed wild artiodactyl in South America and occurs in four of the continent's 10 major habitats: desert and xeric shrublands, montane and lowland grasslands, savannas and shrublands, and wet temperate forests. Guanacos are both grazers and browsers and range from coastal forests at sea level to sparsely vegetated alpine deserts at 4500 m. In some areas, they may be found in isolated but productive meadows. Galloping Guanacos can exceed 55 km/h.
4) Llama (Lama glama). This species, domesticated largely as a pack animal and meat source (although there is a less common long-wooled form valued for its wool), is derived from the Guanaco lineage. Llamas were domesticated between 4000 and 6000 years ago. With a mass of up to 220 kg, Llamas are the largest of the four South American camelids and the tallest Neotropical mammals.
5) Viçuna (Vicugna vicugna). This species is far more specialized in its habitat needs than is the Guanaco, being restricted to the high Andean Puna at 3200-4800 m. The Viçuna and Guanaco lineages are believed to have diverged between two and three million years ago. The unique equatorial high-altitude short grassland in which the viçuna lives is found above the treeline but below the snowline. With their gazelle-like build, viçuna are the smallest camelids. The Viçuna and Alpaca are the only living ungulates with continuously growing incisors, like rodents. Viçuna and Alpacas need to drink frequently, unlike other camelids.
6) Alpaca (Vicugna pacos). This species, domesticated largely as a wool producer, is derived from the Viçuna lineage. Its long, thick wool can give an alpaca the bulky appearance of a Llama (in fact, it was at one time placed in the genus Lama, before its much closer relationship with the Viçuna was recognized). More than a million small producers in the central Andes depend upon the Alpaca and Llama as their pricipal means of subsistence. Most Alpacas are kept in herds of fewer than 50 individuals, but some commercial herds can reach 50,000 animals. Alpacas are believed to have been domesticated between 5500 and 6500 years ago.
Llamas and Alpacas have hybridized extensively.
Camels and Humans
Camelids have played critical roles in a range of human societies. For thousands of years in Central Asia and Asia Minor, domestic Bactrian Camels have served as pack animals in times of peace and war and as draft animals for ploughing and pulling carts. They connected the East and West over the Silk Road, bringing Siberian and Indian goods to the Persians. They have also provided meat, milk, and wool, without which the Gobi Desert would have been uninhabitable for humans. Because these camels are intolerant of high temperature for long periods while they are working, caravans traveling westward from China across the Gobi Desert always traveled in winter. Since at least 1000 BC, Bactrian Camels have been intentionally hybridized with Dromedaries, yielding especially strong pack animals known as Bukhts. Hybrids are also produced in an effort to combine the higher milk production of the Dromedary with the cold tolerance of the Bactrian. The world population of domestic Bactrians is estimated to be somewhere between around 750,000 and one million (according to some sources, as many as two million), with most of these animals found in Mongolia and China.
In the hot deserts of North Africa, Dromedaries have served as beasts of burden and provided meat, milk, fat, and fuel, as well as soft wool for clothing and tents. In some areas, live Dromedaries are bled for consumption of the blood (with a single individual yielding up to 35 liters per year).
Around 95% of the camelids in the Old World are Dromedaries, with nearly all of the remaining 5% being domestic Bactrian Camels (there are no more than a few thousand wild Bactrian Camels). In the New World, around 47% of camelids are Llamas, 41% Alpacas, 8% Guanacos, and 4% Viçunas.
(Franklin 2011 and references therein)
- Franklin, W.L. 2011. Family Camelidae (Camels). Pp. 206-246. in: Wilson, D.E. & Mittermeier, R.A., eds. Handbook of the Mammals of the World. Volume 2. Hoofed Mammals. Lynx Edicions, Barcelona.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimen Records: 42
Specimens with Sequences: 54
Specimens with Barcodes: 42
Species With Barcodes: 7
Public Records: 39
Public Species: 7
Public BINs: 5
Camelids are members of the biological family Camelidae, the only currently living family in the suborder Tylopoda. The extant members of this group are: dromedary camel, Bactrian camels, wild or feral camels, llamas, alpacas, vicuñas, and guanacos.
Camelids are large animals with slender necks and long legs, and are strictly herbivorous. They differ from true ruminants in a number of ways. Their dentition show traces of vestigial central incisors in the upper jaw, and the third incisors are developed into canine-like tusks. Camelids also have true canine teeth and tusk-like premolars, which are separated from the molars by a gap. The musculature of the hind limbs differs from those of other ungulates in that the legs are attached to the body only at the top of the thigh, rather than attached by skin and muscle from the knee upwards. Because of this, camelids have to lie down by resting on their knees with their legs tucked underneath their bodies. They are pseudoruminants with three-chambered stomachs, rather than four-chambered ones; their upper lips are split in two, with each part separately mobile; and, uniquely among mammals, their red blood cells are elliptical. They also have a unique type of antibodies, lacking the light chain, in addition to the normal antibodies found in other mammals. These so-called heavy-chain antibodies are being used to develop single-domain antibodies with potential pharmaceutical applications.
They do not have hooves, rather they have two-toed feet with toenails and soft foot pads (Tylopoda is Latin for "padded foot"). The main weight of the animal is borne by these tough, leathery sole pads. The South American camelids, adapted to steep and rocky terrain, can move the pads on their toes to maintain grip. Many fossil camelids were unguligrade and probably hooved, in contrast to all living species.
Camelids are behaviorally similar in many ways, perhaps most notably in their walking gait during which both legs on the same side are moved simultaneously. The consequent swaying motion of camelids large enough for human beings to ride is legendary.
The three Afro-Asian camel species have developed extensive adaptations to their lives in harsh, near-waterless environments. Wild populations of the Bactrian camel are even able to drink brackish water, and some herds live in nuclear test areas.
Comparative table of the eight mammals in the family Camelidae (ordered by weight):
|Camel-like mammal||Image||Natural range||Weight|
|Bactrian camel||Central Asia|
|300 to 1,000 kg (660 to 2,200 lb)|
|South Asia and Middle East|
|300 to 600 kg (660 to 1,320 lb)|
|Wild camel||Central Asia|
|Llama||(domestic form of guanaco)||130 to 200 kg (290 to 440 lb)|
|Guanaco||South America||c. 90 kg (200 lb)|
|Huacaya Alpaca||(domestic form of vicuña)||48 to 84 kg (106 to 185 lb)|
|Suri Alpaca||(domestic form of vicuña)||48 to 84 kg (106 to 185 lb)|
|Vicuña||South American Andes||35 to 65 kg (77 to 143 lb)|
Camelids are unusual in that their modern distribution is almost the reverse of their origin. Camelids first appeared very early in the evolution of the even-toed ungulates, around 45 million years ago during the middle Eocene, in present-day North America. Among the earliest camelids was the rabbit-sized Protylopus, which still had four toes on each foot. By the late Eocene, around 35 million years ago, camelids such as Poebrotherium had lost the two lateral toes, and were about the size of a modern goat.
The family diversified and prospered, but remained confined to the North American continent until only about two or three million years ago, when representatives arrived in Asia, and (as part of the Great American Interchange that followed the formation of the Isthmus of Panama) South America. A high arctic camel from this time period has been documented in the far northern reaches of Canada.
The original camelids of North America remained common until the quite recent geological past, but then disappeared, possibly as a result of hunting or habitat alterations by the earliest human settlers, and possibly as a result of changing environmental conditions after the last ice age, or a combination of these factors. Three species groups survived: the dromedary of northern Africa and southwest Asia; the Bactrian camel of central Asia; and the South American group, which has now diverged into a range of forms that are closely related, but usually classified as four species: llamas, alpacas, guanacos, and vicuñas.
Fossil camelids show a wider variety than their modern counterparts. One North American genus, Titanotylopus, stood 3.5 m at the shoulder, compared with the about 2 m of the largest modern camelids. Other extinct camelids included small, gazelle-like animals, such as Stenomylus. Finally, a number of very tall, giraffe-like camelids were adapted to feeding on leaves from high trees, including such genera as Aepycamelus, and Oxydactylus.
Recent genetic studies have shown that the wild Bactrian camel (Camelus ferus) is in fact a distinct species, with an independent evolutionary history to its domestic relative. Dromedary camels, Bactrian camels, llamas, and alpacas all have 74 chromosomes while the wild camel has 77 chromosomes.
- †Subfamily Poebrodontinae
- †Subfamily Poebrotheriinae
- †Subfamily Miolabinae
- †Subfamily Stenomylinae
- †Subfamily Floridatragulinae
- Subfamily Camelinae
- Tribe Lamini
- Tribe Camelini
|Camelid ancestor||North America|
|Lamini||10.4 mya||1.4 mya||Guanaco||South America|
|Camelini||8 mya||Bactrian camel||Asia|
|Aepycamelus||Miocene||Tall, s-shaped neck, true padded camel feet|
|Camelops||Pliocene-Pleistocene||Large, with true camel feet, hump status uncertain|
|Floridatragulus||Early Miocene||A bizarre species of camel with a long snout|
|Eulamaops||Pleistocene||From South America|
|Hemiauchenia||Miocene-Pleistocene||A North and South American lamine genus|
|Megacamelus||Miocene-Pleistocene||The largest species of camelid|
|Megatylopus||Miocene-Early Pleistocene||Large camelid from North America|
|Oxydactylus||Early Miocene||The earliest member of the "giraffe camel" family|
|Palaeolama||Pleistocene||A North and South American lamine genus|
|Poebrotherium||Oligocene||This species of camel took the place of deer and antelope in the White River Badlands.|
|Procamelus||Miocene||Ancestor of extinct Titanolypus and modern Camelus|
|Protylopus||Late Eocene||Earliest member of the camelids|
|Stenomylus||Early Miocene||Small, gazelle-like camel that lived in large herds on the Great Plains|
|Titanotylopus||Miocene-Pleistocene||Tall, humped, true camel feet|
- Clutton-Brock, Juliet (1987). A Natural History of Domesticated Mammals. p. 208. ISBN 0-521-34697-5.
- Franklin, William (1984). Macdonald, D., ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 512–515. ISBN 0-87196-871-1.
- Savage, RJG, & Long, MR (1986). Mammal Evolution: an illustrated guide. New York: Facts on File. pp. 216–221. ISBN 0-8160-1194-X.
- Chen, B.X., Yuen, Z.X. and Pan, G.W. (1985). "Semen-induced ovulation in the bactrian camel (Camelus bactrianus).". J. Reprod. Fert. 74 (2): 335–339.
- Wild Bactrian Camels Critically Endangered, Group Says National Geographic, 3 December 2002
- Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. pp. 274–277. ISBN 1-84028-152-9.
|This article may be expanded with text translated from the corresponding article in the Spanish Wikipedia. (December 2012)|
Lamini (members are called lamoids) is a tribe of the subfamily Camelinae. It contains two genera and four species, all exclusively from South America: llamas, alpacas, vicuñas, and guanacos. The former two are domesticated species, while the latter two are only found in the wild. All lack sexual dimorphism. The four species can interbreed and produce fertile offspring.
Alpaca (Vicugna pacos)
The alpaca, a domestic camelid, weights between 50 and 65 kg (110 and 143 lb), while its height at the shoulder is 94–104 cm (37–41 in). It is slightly larger than the vicuña. Normally, the alpaca is found in the Andes in Peru and Bolivia, though it also inhabits northern Chile and northwestern Argentina. There are about 3.5 million alpacas in the world. In the 1980s, alpacas started being exported to other countries for farming purposes: they can be found in the United States, Australia, and New Zealand, though the vast majority still reside in South America.
Guanaco (Lama guanicoe)
The guanaco is a wild camelid, standing at 100–120 cm (39–47 in) at the shoulder and 150–160 cm (59–63 in) at the head. It can weigh up to 140 kilograms (310 lb). Its pelage is longer than that of the vicuña but shorter than that of the alpaca; it is considered to be of excellent quality and has a light brown, reddish, or brown-yellow color. The diameter of its fleece's fibers varies between 16 and 18 micrometers.
90% of the world's guanacos are in Argentina, distributed from the islands of Beagle Channel and the southern extremity of the Patagonia to the Puna grassland in northeastern Argentina. Guancos can also be found in Bolivia, Chile, Paraguay, and Peru.
Llama (Lama glama)
The llama is the largest of the extant laminoids and weighs 130–150 kilograms (290–330 lb) with a height of 109–119 cm (43–47 in) at the shoulder. Llamas are not a natural species; rather, they were domesticated by the Peruvians and Bolivians of the highlands. Commercial trade led to the llama's current abundance in Columbia, Ecuador, Peru, Bolivia, Chile, Paraguay, and northeast Argentina. There are bands of llamas in the United States, Europe, Japan, and New Zealand.
The color and length of the llama's wool is variable, depending on the race. The diameter of llama wool's fiber varies between 20 and 80 micrometers, depending on whether the llamas were raised for its wool or as a pack animal.
Vicuña (Vicugna vicugna)
The vicuña is the smallest camelid, with a shoulder height of 75–100 cm (30–39 in) and a weight of 40–60 kg (88–132 lb). Its coat is mainly beige in color and is said to make "the best wool in the world", with the average fiber diameter between 11 and 14 micrometers. Like rodents, the vicuña has continuously-growing incisors. It lives only in areas of high altitude – 3,200 m (10,500 ft) or greater – in the highlands of Argentina, Bolivia, Chile, Peru, and Ecuador.
- Fowler, Murray E. (1998). Medicine and surgery of South American camelids: llama, alpaca, vicuña, guanaco (2nd ed.). Ames, Iowa: Blackwell. ISBN 0813803977.
- Hogan, C. Michael (2008). Strömberg, N., ed. "Guanaco: Lama guanicoe". GlobalTwitcher.
- Stahl, Peter W. (4 April 2008). "Animal Domestication in South America". In Silverman, Helaine; Isbell, William. Handbook of South American Archaeology. Springer. pp. 121–130. ISBN 9780387752280.
- Castillo-Ruiz, Alexandra. "Lama pacos: alpaca". Animal Diversity Web. University of Michigan. Retrieved 10 December 2012.
- Quiggle, Charlotte (Fall 2000). "Alpaca: An Ancient Luxury". Interweave Knits: 74–76.
- "Guanaco: Lama guanicoe". World Association of Zoos and Aquariums. Retrieved 10 December 2012.
- "San Diego Zoo's Animal Bytes: Guanaco". Zoological Society of San Diego. Retrieved 10 December 2012.
- Baldi, B.; Lichtenstein, G.; González, B.; Funes, M.; Cuéller, E.; Villalba, L.; Hoces, D.; Puig, S. (2008). "Lama guanicoe". IUCN Redlist. International Union for Conservation of Nature and Natural Resources. Retrieved 11 December 2012.
- Furlong, Charles Wellington (October 1912 – March 1913). "Hunting the Guanaco". The Outing Magazine 61 (1): 5.
- "Mammal Guide: Vicuña". Animal Planet. Discovery Communications, LLC. Retrieved 11 December 2012.
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