Camelidae

Camelid feet lack functional hooves, the toe bones being embedded in a broad, cutaneous pad.[1]

Camelids are large, strictly herbivorous animals with slender necks and long legs. They differ from ruminants in a number of ways.[2] Their dentition show traces of vestigial central incisors in the incisive bone, and the third incisors have 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.[1] They have 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.[2] They also have a unique type of antibodies, which lack 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.

Camelids do not have hooves; rather, they have two-toed feet with toenails and soft foot pads (Tylopoda is Greek for "padded foot"). Most of the weight of the animal rests on 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.[3] Many fossil camelids were unguligrade and probably hooved, in contrast to all living species.[4]

Camelids are behaviorally similar in many ways, including their walking gait, in which both legs on the same side are moved simultaneously. Consequently, camelids large enough for human beings to ride have a typical swaying motion.

Dromedary camels, bactrian camels, llamas, and alpacas are all induced ovulators.[5]

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.[6]

Comparative table of the seven extant species in the family Camelidae:

Species	Image	Natural range	Weight
Camelus
Bactrian camel (Camelus bactrianus)		Central and Inner Asia (entirely domesticated)	300 to 1,000 kg (660 to 2,200 lb)
Wild Bactrian camel (Camelus ferus)		China and Mongolia (entirely wild)	300 to 820 kg (660 to 1,800 lb)
Dromedary or Arabian camel (Camelus dromedarius)		South Asia and Middle East (entirely domesticated)	300 to 600 kg (660 to 1,320 lb)
Lama
Llama (Lama glama)		(domestic form of guanaco)	130 to 200 kg (290 to 440 lb)
Guanaco (Lama guanicoe)		South America	c. 90 to 120 kg (200 to 260 lb)
Vicugna
Alpaca (Vicugna pacos)		(domestic form of vicuña)	48 to 84 kg (106 to 185 lb)
Vicuña (Vicugna vicugna)		South American Andes	35 to 65 kg (77 to 143 lb)

A dymaxion map of the biogeographic distribution of Camelidae species:

  Tertiary distribution

  Present-day distributions

  Introduced (feral) distributions

The yellow dot is the origin of the family Camelidae and the black arrows are the historic migration routes that explain the present-day distribution.

Camelids are unusual in that their modern distribution is almost the inverse of their area of 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.[4][8]

The family diversified and prospered, but remained confined to the North American continent until only about two to 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. Camelids were domesticated by early Andean peoples[9] and remain in use today.

Fossil camelids show a wider variety than their modern counterparts. One North American genus, Titanotylopus, stood 3.5 m at the shoulder, compared with about 2 m for 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.[4]

Whether the wild Bactrian camel (Camelus ferus) is in fact a distinct species or a subspecies (Camelus bactrianus ferus) is still debated.[10][11] The divergence date is 0.7 million years ago, long before the start of domestication.[11]

A dromedary camel (C. dromedarius) in the Australian outback, near Silverton, New South Wales

South American vicuña (Vicugna vicugna)

Family Camelidae

• †Subfamily Poebrodontinae
• †Subfamily Poebrotheriinae
• †Subfamily Miolabinae
• †Subfamily Stenomylinae
• †Subfamily Floridatragulinae
• Subfamily Camelinae
  • Tribe Lamini
    • Genus: Lama
      • Llama, Lama glama
      • Guanaco, Lama guanicoe
    • Genus: Vicugna
      • Vicuña, Vicugna vicugna
      • Alpaca, Vicugna pacos (previously categorized in genus Lama)
    • Genus: Hemiauchenia
      • †Hemiauchenia macrocephala
      • †Hemiauchenia minima
      • †Hemiauchenia blancoensis
      • †Hemiauchenia vera
      • †Hemiauchenia paradoxa
    • Genus Palaeolama
      • †Palaeolama mirifica
  • Tribe Camelini
    • Genus: Camelus
      • Dromedary, Camelus dromedarius
      • Bactrian camel, Camelus bactrianus
      • Wild Bactrian camel, Camelus ferus
      • †Syrian camel, Camelus moreli
      • †Camelus sivalensis
    • Genus: Camelops
      • †Camelops hesternus
    • Genus: Paracamelus
      • †Paracamelus gigas

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• Pictures of camelid species