Bactrian & Dromedary Camels Camelus bactrianus, C. dromedarius, C. ferus March 2009; minor update April 2013

Kingdom: Animalia
    Phylum: Chordata
        Class: Mammalia
             Order: Artiodactyla (a.k.a. Cetartiodactyla*)
                  Family: Camelidae
                       Subfamily: Camelini
                            Genus: Camelus
                                 Species: Camelus bactrianus (Linnaeus 1758) - domestic Bactrian camel
                                 Species: Camelus dromedarius (Linnaeus 1758) - dromedary
                                 Species: Camelus ferus (Prejevalsky [Przewalski] 1879) - wild Bactrian camel
                       Subfamily: Lamini
                            Genus: Vicugna
                                 Species: Vicugna vicugna - vicuña
                                 Species: Vicugna pacos - alpaca
                            Genus: Lama
                                 Species: Lama glama - llama
                                 Species: Lama guanacoe - guanaco

*New anatomical and DNA evidence on the relationship between Artiodactyla (even-toed ungulates) and Cetacea (whales and dolphins) recently led to a merging of the two orders into a new group, Cetartiodactyla (Montgelard, 1997; reviewed in Kulemzina, 2009). As of October 2012, experts have not yet agreed on whether to define Cetartiodactyla as an official taxonomic order, replacing Artiodactyla and Cetacea. Some continue to list camels in the order Artiodactyla (Franklin, 2011) or use the term Cetartiodactyla without defining it as an order (IUCN, 2008).

• Domestic Bactrian camel
  • First classified by Linnaeus as Camelus bactrianus in 1758
    • Typically wild animal named first, but Linnaeus only knew of domestic camels
  • Common names
    • Bactrian camel
    • Two-humped camel
  • Word origins
    • Camel - Latin camelus from Greek kamelos, from Hebrew or Phoenician gamal; perhaps related to Arabic jamala "to bear" (Harper, 2012)
    • Bactrian - for Bactriana region of ancient Persian Empire (Franklin 2011; Bulliet, 1990)
      • Latin bactriana from Persian bakhtar "the west"
      • Camel caravans transported goods through this northwestern region of modern-day Afghanistan via major trade routes, including the Silk Road
• Dromedary
  • First classified by Linnaeus as Camelus dromedarius in 1758
  • Common names
    • Dromedary
    • Arabian camel
    • One-humped camel
  • Word origins
    • From Greek dromas "running" (Arabian camels were bred and trained for riding)
    • Early variant - drumbledairy (1560s) (Harper, 2012)
• Wild Bactrian camel
  • Taxonomic history
    • Discovered by Przewalski in 1878, named Camelus ferus
    • C. ferus confirmed in 2003 by International Commission on Zoological Nomenclature; recognized by the IUCN
    • DNA analysis confirmed that C. ferus should be considered a separate species from the domestic Bactrian camel (Silbermayr, 2009)
      • Previously, some considered it to be a sub-species, C. bactrianus ferus (Grubb, 2005)
  • Word origins
  • From Greek ferus "wild"

• Camelidae family: origins
  • Camelids originated in North America during Eocene, 40-45 million years ago (Cui et al., 2007)
    • Appeared in middle Eocene in North American mountains/plains/deserts (Franklin, 2011)
    • Earliest camelids similar to modern guanaco but rabbit-sized (30 cm at shoulder) (Franklin, 2011)
    • One of most common ungulates in N. America by late Eocene and Oligocene (24-36 million years ago) (Franklin, 2011)
• Camelid tribes: four emerged, two survived
  • Camelini and Lamini diverged about 11-25 million years ago
    • DNA evidence dates the split to 25 million years ago (Cui et al., 2007)
    • Fossil record dates the split to 11 million years ago (Webb 1974; Harrison 1979)
  • Only Camelini and Lamini tribes survived worldwide climactic changes at end of Miocene
    • Camelini (wild and domestic Bactrian camels and dromedaries)
      • Old World (Asia, Middle East, Arabia, North Africa)
    • Lamini (lamas, alpacas, vicuñas and guanacos)
      • New World (South America)
• Migration: two major waves (Webb, 1974)
  • About 3 million years ago (late Tertiary):
    • Camelini migrated across Bering land bridge into Asia
      • Across dry belt of Eurasia, into India
    • Lamini migrated to mountains of South America
  • Both tribes became extinct in North America
• Modern camelid species
  • Unclear evolutionary relationship between C. bactrianus, C. dromedarius, and C. ferus
  • DNA studies show wild Bactrian Camel not ancestor of two domesticated species as previously thought
    • C. ferus is separate lineage and not direct progenitor of C. bactrianus (Ji et al, 2009)
  • Bactrian camels and dromedaries diverged about 5-8 million years ago (Cui et al., 2007; Franklin, 2011)
    • No evidence to indicate that Bactrian camels were ancestor of dromedaries (Kinne et al, 2010
• Domestication
  • Bactrian camels
    • Domesticated about 4,000-6,000 years ago in steppes of eastern central Asia ( Peters & von Driesch, 1997; reviewed in Groeneveld et al., 2010)
  • Spread to Asia Minor ca. 4000 BC, Middle East ca. 2000 BC, China ca. 400 BC
  • Currently 16 Bactrian camel breeds (FAO, 2008)
• Dromedaries
  • Domesticated about 4,000-5,000 years ago (Peters, 1997; reviewed in Groeneveld et. al., 2010)
  • Currently 97 dromedary breeds (FAO, 2008)
• Wild Bactrian camels
  • Unclear how they avoided domestication
    • Possibly due to isolation in the Gashun Gobi (Lop Nur) area and Mongolia
    • Perhaps instinctive shyness played a role
• Hybrids (reviewed in Bulliet, 1990)
  • Bactrian camel-dromedary hybrid (bukht) - single long hump
    • First generation larger and stronger than either parent (hybrid vigor)
  • Camel-llama hybrid (cama) (Jones et al., 2008)
    • Female llama fertilized with camel sperm at Camel Reproduction Centre (Dubai)

• Early camels ranged from Asia to Eastern Europe.
• Some crossed into Africa, expanding across northern Africa and as far south as northern Tanzania.
• Wild Bactrian camel: 3 separated regions in China, 1 in Mongolia (click here for IUCN distribution map)
  • China
    • Gashun Gobi, in Gansu
    • Taklamakan Desert, in Xinjiang (this population may now be extinct)
    • Arjin Shan Lop Nur Nature Reserve' (est. 2000) and adjacent northern slopes of Arjin Shan mountains
    • Great Gobi Reserve A (est. 1982) and adjacent areas (Reading et al., 1999; Mix et al., 2002; Wang et al., 2002)
  • Mongolia
    • Trans Altai Govi Desert (Mix et al., 2002; Yadamsuren et al., 2004; Yadamsuren & Dovchindorj, 2005)
      • Foothills of the Edren Range to Shiveet Ulaan
      • Hükh Tömörtei Range to the state border
• Domestic animals have been introduced throughout the arid and semiarid regions of the world.
  • Domestic Bactrian camel
    • Cold desert regions of Iran, Afghanistan, Pakistan, Kazakhstan, Mongolia, and China.
  • Dromedary
    • Hot arid deserts of Africa and Middle East Most in Somalia and Sudan.
  • A large feral population of dromedaries exists in Australia
  • Archeological finds indicate that both the dromedary and domestic Bactrian camel were imported into northern provinces of the Roman Empire for military and civilian use (Pigière & Henrotay, 2012).

Past and current distribution of wild Bactrian camels in Mongolia and China (Schaller, 1998). Click map to enlarge. Click here for IUCN distribution map	Normal distribution and points of introduction of domestic camels (Wilson, 1998). Click map to enlarge.

Habitat of wild Bactrian camel:

• Desolate mountain and desert areas
  • Rocky mountain massifs
  • Deserts: flat pavement-like expanses, sand dunes
  • Oases fringed with poplars (Populus)
• Sparse vegetation
  • Plants include: Haloxylon, Caragana, Salsola
• Large temperature range
  • Up to 60-70°C (140-160°F) in summer
  • Down to -30°C (-22°F) in winter
• May inhabit areas with no fresh water
• Adapted to drinking saltwater slush, which domestic camels will not touch.

• Unlike their llama/vicuna/guanaco relatives, all camel species have:
  • At least one hump on their backs
  • Long curved necks
  • Broad, large feet
  • Tufted tails
  • Four teats rather than two
  • Annual molt 
• Domestic camels have been selectively bred over many years
  • Hybridization between species has occurred in ancestry of both domestic species

Wild Bactrian	Bactrian	Dromedary
Southern Silk Road, China Image credit: John Hill via Wikimedia Commons, some rights reserved	Frankfurt Zoo, Germany Image credit: Elinor D via Wikimedia Commons, some rights reserved	Tierpark Berlin, Germany Image credit: Agadez via Wikimedia Commons, some rights reserved
2 humps: small, pointed, conical 1.7 m (5.6 ft) at shoulder 450 kg (992 lb) Frame: small and lithe relative to domestic Bactrian; laterally compressed (Mongolian name ‘havtagai’ means ‘flat’) Legs: slender, no callosities on knees Feet: narrow Hair: tan or grayish on body; long and dark brown on upper legs, neck, top of humps, tail	2 humps: large, irregularly shaped, may become flaccid and flop to one side in adult 1.8 m (6.0 ft) at shoulder (shorter than dromedaries) 600-1,000 kg (1,320-2,200 lb) Legs: short and stout Feet: broad, 2-toed, cushioned by fat Hair: long and dark; annual molt	1 hump: more elastic than Bactrian, shrinks with age instead of flopping to side 2.0 m (6.5 ft) at shoulder 400-600 kg (880-1,325 lb) Frame: lighter than Bactrian Legs: longer and more slender than Bactrian Hair: short, light-colored

Feet

• Large broad ‘elastic’ pad – hooves are 2 fingernail-like toenails on front of pad.
  • Unique among mammals
  • Other ungulates walk on tips of hoof-covered toes.
• Wild Bactrian’s hoof is claw-like.

• Composed of fibrous tissue and fat.
  • When fat is metabolized, it acts as a source of energy.
  • Concentration of body fat in humps is advantage in hot climate
    • Minimizes its presence throughout the rest of body
    • This reduces heat-trapping that occurs with insulating layers of fat.
• Available nutrition determines size and shape - humps nearly disappear with starvation.
• Not used for water storage.

• Slit-like - can be closed to protect against blowing sand.
• Special nasal cavities moisten air on way in, trap moisture going out.

• Long lashes protect against blowing sand.
• Bony arch over eye acts as sun shield.
  

• Small and rounded
• Hairs protect ears from blowing sand.

• Prehensile lips, upper split in 2 halves.
• Incisors and canine teeth grow though out life.

• Adaptations for extremely dry environment
  • Red blood cells are oval (round in other mammals) Can flow quicker in a dehydrated state.
  • Urine is more concentrated than other animals (less water loss).
  • Dung is dry.
  • The kidneys and intestines are good at retaining water.
  • Can tolerate loss of water equal to over 30% of body weight (Franklin, 2011)
    • Most mammals die if they lose 15%
  • Wild Bactrian camel (not domestic) is able to drink saltwater slush when fresh water is unavailable.
• Adaptations for extreme heat
  • Able to endure temperature extremes, from -40°F in January to over 100°F in summer.
  • Have only 25% the number of sweat glands as found in cattle
  • Only sweat when body temperatures reach 41-42°C (105.8-107.6°F)
  • Long legs keep bodies further from the hot ground to reduce overheating.
• Adaptations for blowing sand
  • Long eyelashes and ear hairs form a barrier against blowing sand.
  • Slit-like nostrils can close.

• Domestic camels are maintained in a semi-wild state.
  • Obtain food from wild vegetation, water from humans.
  • Unguarded camels will return to a familiar well.
• Wild camels extremely shy
  • Acute eyesight allows them to spot danger 2-3 miles away and flee.

• Browse or graze 6-8 hours and chew cud another 6-8 hours each day.
  • Constant work required to maintain health on low-nutrient desert plant life.
• Previously, migrated seasonally to green pastures.
  • Movements now curtailed by human settlements.

• Form small groups of ~6 (2-15 animals).
• Gather at oases and small springs.

• Dromedaries may blow out (spit) cud when excited.
• Bactrian camels do not deliberately spit (may if abused).
• Little aggressive behavior except among males during breeding season.
• Adults may push with lowered head and neck.
• Canine teeth used as weapons.
• Extreme fights can result in death of both combatants.

• Displays
  • Rutting male stretches to hold head higher than a rival.
• Vocalization
  • Rutting cry: “blo-blo-blo"
• Olfaction/Scent Marking
  • Male rubs back of head against shoulders (occipital gland)
  • Secretions tend to excite males

Locomotion

• Normal gait is a “pace”
  • One foreleg and one hind leg on same side of body move forward together.
• Horses, dogs and bears can also “pace” but more often “trot” (foreleg and diagonally opposite hind leg move together).
• If necessary, can travel 150 km (93 mi) in 15-20 hours.
• Maximum speeds
  • Up to 65 km/hr (40 mi/hr) in short bursts
  • Up to 40 km/hr (25 mi/hr) sustained speed
• Domestic camels have been known to swim (observed in Australia).

• Integral part of nomad culture
  • Domesticated by nomads 3,000-4,000 years ago
  • Still important part of nomad life in the Sahara
  • Provide meat, milk, hides, wool
    • Females can produce milk for several years after birth
      • Bactrian: 5 liters/day
      • Dromedary: 20 liters/day
  • Males used for transport and riding
    • Easily carry 200 kg (440 lb)
  • Can pull more than 1 ton with a wheeled cart
  • Convex back allows camels to carry more than a horse.
    • Load is limited by the weight it can rise with.
• Used in combat
  • Used because of their ability to scare horses
    • Camel smell spooks horses
• U.S. Army camel corps stationed in California in the 1800s
  

Food

• Ruminants (cud-chewing mammals) usually have 4 stomachs (occasionally 3).
  • Plant material partially broken down in the 1st two stomachs, then regurgitated as cud and chewed again.
  • Microbes (bacteria, protozoa, and fungi) in other stomach chambers assist in digestion.
  • Other ruminants: antelopes, sheep, goats, cattle, giraffes and bison and deer.
• Camels graze (like sheep) and browse (like antelopes).
• Move while feeding no matter how rich or poor the vegetation (do not degrade desert flora).
• Mainly grasses, leaves and twigs of shrubs and trees - all plants of the desert.
  • Green shoots of saxaul (Haloxylon).
  • Stems and foliage of Salsola, Ephedra, Zygophyllum.
  • Species in Mongolia include:Caragana, Haloxylon, Reaumuria, Salsola.
• At oases: poplar, willows and reeds
• Camels recognize poisonous plants growing in the area and will not eat them.
  • May eat poisonous plants in new, unfamiliar area,
• Can eat sharp, thorny plants other animals can’t eat (e.g., saltbush).

• Able to go without water for a long time (generally thought to be 4-5 days). (Wilson, 1998)
  • When dehydrated, body temperature can lower to 34°C to as high as 41°C.
  • Produce small amounts of concentrated thick urine.
  • Kept in same conditions, cattle lose water 3 times faster than camels
• If the camel is kept near a water source or river, it may drink daily.
• In cold weather, and when green feed is available, may not drink water for months
• A thirsty camel in a hot dry season can drink up to 200 liters of water in one day. (Gauthier-Pilters, 1981)

     Male:

• Experiences rut in cooler / wetter seasons (greatest availability of food)
  • Peak rut in the Gobi: late January/February
• Develops a dulaa (sack-like extension of the soft palate).
  • Triggered by rising testosterone levels
  • Used to produce a mating call: male blows through the dulaa to push it outside the mouth (looks like a red balloon) and uses saliva to create a low gurgle.
  • Size of dulaa indicates male's virility to female
• Marks territory with secretions from poll glands (on back of neck, between ears).
  • Rub necks on any solid object.
• Urinates on own tail and swishes it over its back
  • Urine contains female-attracting pheromones.
• Exhibits aggression toward other males.
• Becomes extremely active - collects a harem with as many as 10-20 females (Bannikov, 1976)

     Female:

• Undergoes estrous cycle in breeding season: receptive 3-4 days, non-receptive following 10 days.
• Bleats to indicate receptive
• Approaches male, presenting hind quarters.
  • Urinates constantly.
  • Flips tail up and down – short quick movements.
• Dulaa also present in females but never extruded.
  

• Only ungulate that copulates in a “sitting” position.
• Reproduction rate: 2 year interval (minimum).

• ~1 year (12-14 months).
• No reports of twins.
• Most young born at end of March / April.

• Female becomes restless, and separates from the other animals.
• Calf size dependent on the size of its parents. ~35 (25 kg – 52 kg) males larger than females.
• Front legs of calf appear first - then head.
• Calf’s drop to the ground breaks umbilical cord.
• Only ungulates that do not lick and clean baby. Sniff extensively and help the calf find milk.
• Mother is protective – won’t accept orphaned young (these must be hand-raised)

• Precocious young – stand and walk shortly after birth.

• Starts to eat grass around 2 - 3 months old.
• Weaning at 1-2 years.

• 40-50 years.

• Camel’s hostile environment is also unfavorable to many diseases.
• Wild adults are hardy, but infant mortality high.
  • Passive immunity not transmitted via the placenta - must be acquired after birth.
  • Colostrum carries antibodies to diseases the mother has been exposed to.
• Domestic camel problems
  • Surra (trypanosomiasis)
    • Caused by very small parasites (trypanosomes) that live in the blood.
  • Camel Pox (skin lesions)
    • Highly contagious, usually cures itself in dry climates.
  • Wide range of parasitic diseases (internal and external).
  • Respiratory disease complex
    • Several viruses, bacteria, fungi and parasitic worms involved.
  • Many reproductive disorders.
    • High abortion rates.
    • Males: cryptorchidism.
    • Females: cystic ovaries.
      

• A small population of introduced dromedaries and Bactrian camels survived in the Southwest United States until the 1900s.
  • Imported from Turkey.
  • Part of the US Camel Corps experiment, used as draft animals in mines.
  • Escaped or were released after the project was terminated.
• Camel milk is a staple food of desert nomad tribes - richer in fat and protein than cow milk.
  • Cannot be made into butter in the traditional churning method. – must be soured first, churned, and a clarifying agent added.
  • The milk can readily be made into yogurt.
  • Many healthful properties - used as a medicinal product in India.
  • Bedouin tribes believe it has curative powers.
• The Wild Camel Protection Foundation has established a captive breeding program in Mongolia.
  • Only 15 wild Bactrian camels are currently in captivity in China and Mongolia.
  • Females can produce 1 young every two years.
  • Artificial insemination may be necessary with such small numbers.

• ISIS captive population
• FAO data – more than 22 million worldwide, per WRI EarthWatch database
  • Majority are dromedaries (more than 15 million)
    • Most in Somalia (7 million) and Sudan (4 million)
    • Ethiopia and Kenya also have substantial numbers
  • 700,000 feral animals in Australia are becoming a problem
• IUCN data on wild Bactrian camels (2004)
  • About 600 in China
  • About 350 in Mongolia (down from about 650 in 1985

• 'Great Gobi Reserve A' established in Mongolia in 1982 and in 2000 - These are the herds completely isolated and unable to hybridize with domestic Bactrians
• 'Arjin Shan Lop Nur Nature Reserve' was established in China in 2000. Opening of a second Nature Reserve in China is also needed.
• The Wild Camel Protection Foundation has established a captive Wild Bactrian Camel breeding program in Mongolia.
  • Only fifteen wild Bactrian camels are currently in captivity in China and Mongolia.
  • Females can produce 1 young every two years.
  • Artificial insemination may be necessary with such small numbers.
• IUCN Status:
  • 1960's: Rare
  • 1970's - 1994: Vulnerable
  • 1996 - 2000: Endangered (Criteria: A1cd, C2a)
  • 2002, 2007: Critically Endangered

• In Mongolia, 25 to 30 Wild Bactrian Camels are being killed annually when they migrate across the international border into China (subsistence/sport hunting by locals).
• In China in the new Arjin Shan Lop Nur Nature Sanctuary, up to 20 Wild Bactrian Camels are killed annually by miners and hunters for subsistence use.
• Wolves are their principal non-human predator.
• Drought has reduced number of oases, making wolves more successful predators.
• For 45 years, this area of the Gashun Gobi was the nuclear test site area of China.
  • Wild Bactrian Camel survived and is apparently breeding naturally.
• Other threats include:
  • Highly toxic illegal mining
  • Habitat used for gas pipe line
• Domestic Bactrian camels and goats have also been introduced to the designated areas and hence compete for grazing and water.
  • Species could be wiped out if their natural habitats in China and Mongolia are destroyed.
  • Threat of hybridization
• High infant mortality due to harsh conditions.

• Animal Info: Wild Bactrian camel fact sheet
• FAO Animal Health Care Manual: Chapter 7: Camels, llamas and alpacas
• IUCN Red List: Wild Bactrian camel fact sheet
• USDA: Information Resources on Old World Camels: Arabian and Bactrian 1941-2004
  
• Wild Camel Protection Foundation
• ZipCodeZoo.com: Camelus fact sheet

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