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Genus Camelus Linnaeus, 1758 TYPE SPECIES. — Camelus bactrianus Linnaeus, 1758 by original designation., Published as part of Martini, Pietro & Geraads, Denis, 2018, Camelus thomasi Pomel, 1893 from the Pleistocene type-locality Tighennif (Algeria). Comparisons with modern Camelus, pp. 115-134 in Geodiversitas 40 (5) on page 118, DOI: 10.5252/geodiversitas2018v40a5, http://zenodo.org/record/5371960, {"references":["LINNAEUS C. 1758. - Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Salvii, Stockholm, 824 p. https: // doi. org / 10.5962 / bhl. title. 542"]}

Resumen: El registro óseo faunístico exhibe una preservación diferencial en los sitios arqueológicos a cielo abierto y aleros de la Puna Austral de Argentina. Dada esta variabilidad, y con el fin de establecer los principales factores que actuaron como elementos de control tafonómico sobre los huesos y piezas dentales de Camelidae (ca. 9000 años AP hasta Período Republicano), se realizaron estudios petrográficos y geoquímicos aplicando DRX. Las muestras presentan un comportamiento diagenético diferente, siendo los huesos más sensibles que los esmaltes dentales a las variaciones en la composición química. En los sitios aleros la diagénesis fue más intensa, donde la DRX ósea muestra picos de Ca5 (PO4)3(OH) con reemplazo mineral de KCl para el Holoceno temprano. En el Holoceno tardío, para el alero y para los sitios a cielo abierto se observan picos de PH3. El material óseo alterado térmicamente y las partes dentales no muestran reemplazos minerales. Abstract In order to assess the variability in preservation of the bone record from the ca. 9000 BP to recent date were conducted several X-ray Diffraction (XRD) and petrographical analysis on samples from several archaeological sites from the Southern Argentine Puna. The samples are composed of bone and teeths from open-air and overhang sites. Samples display variability in diagenetic pathways, being bones more sensitive than teeths to suffer chemical exchanges. In overhang sites diagenetic processes were more intense, displaying XRD high values of Ca5 (PO4)3(OH) and mineral exchange of KCl to the Early Holocene. In Late Holocene, in open-air and overhang sites were observed high values of PH3. Thermal altered bones and teeth fragments have not displayed mineral exchanges.

Zazula, Grant D., Macphee, Ross D. E., Hall, Elizabeth, Hewitson, Susan (2016): Osteological assessment of Pleistocene Camelops hesternus (Camelidae: Camelinae: Camelini) from Alaska and Yukon. American Museum Novitates 2016 (3866): 1-46, DOI: 10.1206/3866.1, URL: http://www.bioone.org/doi/10.1206/3866.1

Camelops hesternus Leidy, 1854 MANDIBLE AND LOWER DENTITION Referred specimens: F: AM 35172 (Gold Hill, Alaska), mandible fragment, juvenile, with unworn, partially erupted p4–m1 (fig. 3 A–C); F: AM 34631 (Cripple Creek, Alaska), right mandible fragment, with heavily worn m1–m3 (fig. 3 D–F); F: AM 35168 (Fairbanks Creek, Alaska), anterior mandible fragment with symphysis, diastema, and roots of right p4 (fig. 3, G, H); NMC 42549 (Sixtymile Loc. 3, Yukon), left, juvenile, mandible with dp3–dp4, m1, lacking symphysis and coronoid process (fig. 3 I–L); F: AM 144676 (Cripple Creek, Alaska), heavily worn left m3, roots broken off (fig. 4 A–C); F: AM 35154 (Cripple Creek, Alaska), heavily worn, right m3, roots broken off (fig. 4 D–F); F: AM 35169 (Gold Hill, Alaska), right, moderately worn m2?, root of posterior lobe sampled, anterior root broken off (fig. 4 G–I); F: AM 35170 (Dawson Cut, Alaska), relatively unworn, left m1 or m2, roots and anterior margin broken off (fig. 4 J–L); F: AM 35173 (Gold Hill, Alaska), left m2, anterior cusp and root missing (fig. 4 M–O). Description: The most complete mandible in this collection represents a juvenile (NMC 42549), described below. This and others specimens described here exhibit several features consistent with their allocation to Camelops hesternus (Webb, 1965), including: (1) sharp, labially concave diastemal crest (fig. 3H, L); (2) deep horizontal ramus featuring slight concavity below diastema and relatively straighter ventral border in lateral view compared to other BARCs (fig. 3 I–K); (3) large mental foramen situated low on the mandible, below posterior end of canine root (fig. 3G, J); (4) distinct angular “spur-shaped” process present on posterior margin of ascending ramus, below condyle but above level of horizontal ramus (fig. 3 I–K), and lacking inflection seen in other camelines (Harrison, 1985). The mandibular fragments representing mature adult individuals (F: AM 34631 and F: AM 35168) exhibit horizontal rami that are much broader transversely and more robust than in other BARCs (table 1). Individual mandibular dental loci can be readily differentiated from those of other BARCs. Key features consistent with descriptions and illustrations of Camelops hesternus (Webb, 1965; Dalquest, 1992) include: (1) lower molars large, highly hypsodont (fig. 4G, H), and relatively long mesiodistally compared to their transverse widths (fig. 4 A–F); (2) molar lophs separated by deep, narrow valleys on the labial side (fig. 4D, F), relatively flat on the lingual side (fig. 4E, F); (3) loph crests higher and sharper lingually than labially, and more strongly developed on anterior cusps than posterior (fig. 4B); (4) typical wedge shape when viewed in lateral profile, with the occlusal surface much wider than the base, best exemplified by the unworn m2 F: AM 35169 (fig. 4G, H); (5) long, thin, weakly U-shaped infundibula (lakes) on occlusal surfaces, opening lingually, with thicker enamel on lingual side than labial and lacking internal cementum (fig. 4C, F, I,L). Only one specimen, F: AM 35173 (partial left m2; fig. 4O) exhibits infundibular cementum; (6) F: AM 35169, an unworn m2, presents a weakly developed mesiolabial enamel fold or “llama buttress” (fig. 4I), a typical feature of Camelops hesternus lower molars (Webb, 1965, Dalquest, 1992). However, “llama buttresses” are missing from most of our sample of mandibular molars, presumably as a consequence of wear. The juvenile NMC 42549 possesses deciduous premolars dp3 and dp4 and an unerupted m1. Both premolars are notably molariform, though splayed roots can be detected radiographically (fig. 3K). The dp4 is strongly trilobate, resembling an m3, while dp3 is weakly bilobate, resembling a deformed m2 (fig. 3L). Due to marked interproximal wear (a common feature of cameline lower dentitions; see Meachen, 2003), the dentition of F: AM 34631 appears very compressed (fig. 3D). Enamel has been almost completely removed from the mesial and distal surfaces of m1 and m2, with the result that the mesial margin of the latter is inset into the distal margin of the former. The degree of mandibular molar wear on this individual suggests that it was an old, but not yet senile, individual (see Dalquest, 1992)., Published as part of Zazula, Grant D., Macphee, Ross D. E., Hall, Elizabeth & Hewitson, Susan, 2016, Osteological assessment of Pleistocene Camelops hesternus (Camelidae: Camelinae: Camelini) from Alaska and Yukon, pp. 1-46 in American Museum Novitates 2016 (3866) on pages 6-8, DOI: 10.1206/3866.1, http://zenodo.org/record/4585311, {"references":["Leidy, J. 1854. Description of a fossil apparently indicating an extinct species of the camel tribe. Proceedings of the Academy of Natural Sciences of Philadelphia 68: 131 - 150.","Webb, S. D. 1965. The osteology of Camelops. Bulletin of the Los Angeles County Museum 1: 1 - 54.","Harrison, J. A. 1985. Giant camels from the Cenozoic of North America. Smithsonian Contributions to Paleobiology 57: 1 - 29.","Dalquest, W. W. 1992. Problems in the nomenclature of North American Pleistocene camelids. Annales Zoologici Fennici 28: 291 - 299.","Meachen, J. A. 2003. A new species of Hemiauchenia (Camelidae; Lamini) from the Plio-Pleistocene of Florida. Master of Science thesis, University of Florida, Gainesville, 58 pp."]}

O'Leary, Maureen A. (2016): Comparative Basicranial Anatomy Of Extant Terrestrial And Semiaquatic Artiodactyla Maureen A. O'Leary. Bulletin of the American Museum of Natural History 2016 (409): 1, DOI: 10.1206/0003-0090-409.1.1, URL: http://www.bioone.org/doi/10.1206/0003-0090-409.1.1

2. Llama Lama glama French: Lama / German: Lama / Spanish: Llama Taxonomy. Camelus glama Linnaeus, 1758, Peru, Andes. The Llama was selectively bred from the Guanaco (L. guanicoe) for use as a pack animal and producer of meat. It is regarded throughout the world as the premier symbol of South American fauna. With the possibility of more than one center, Llama domestication occurred 4000-4500 years ago in the South-Central Andes (northern Chile and north-western Argentina) and/or 4500-5500 ago in the Central Andes (Junin de los Andes, Argentina). It is often assumed and reported that the Lake Titacaca region was a core of Llama domestication, but supporting data are lacking from early archaeological sites. Osteological remains and DNA analysis document the origin of domestication to be within the range of the northern subspecies of Guanaco L. g. cacsilensis. From its points of domestication, archeological evidence reveals that breeding and herding of Llamas spread widely throughout the Andean region to intermountain valleys, cloud forest on the eastern slope of the Central Andes, southern coast of Peru, to the mountains of Ecuador. Llamas closely resemble their progenitor the Guanaco in almost all aspects of physiology, behavior, general morphology, and adaptability to a wide range of environments. There are no subspecies, but two distinct phenotypic breeds: Short-Woolled and Long-Woolled Llamas. Distribution. Llamas are found at 3800-5000 m above sea level in the Central Andes, from C Peru to W Bolivia and N Argentina. Llama distribution reached its apex during the expansion of the Inca Empire (1470-1532 ap), when pack trains were used to carry supplies for the royal armies to S Colombia and C Chile. Although originally indigenous and endemic to South America, Llamas have now been exported to countries around the world as a companion animal, featured in livestock shows, used for trekking and backpacking, cottage industry and home use ofits wool, and in North America increasingly utilized as a guard animal for protecting sheep and goats from canid predators. Descriptive notes. Head-body 180-229 cm, tail 18-22 cm, shoulder height 102-106 cm; weight 110-220 kg. Llamas are the largest of the four cameloids and tallest of all Neotropical animals. Classical, camelid-body shape with long slender necks, long legs, and small head compared to the body. Their pelage can be white, black, or brown with all intermediate shades occurring and a tendency for spots and irregular color patterns. Wild-type Llamas occur with Guanaco coloration. There are two distinct phenotypic breeds. SHORT-WOOLLED LLAMAS: Slim and long-bodied, with short coats and visible guard hairs, Short-Woolled Llamas are typically the breed utilized for carrying cargo and are the more common of the two. In the Altiplano regions of La Paz, Oruro, and Potosi Departments the proportion of Short-Woolled Llamas varies between 65% and 83%, while in the Peruvian highlands they represent 80% of the total Llama population. The fleece is low density, low weight (1-3 + 1-1 kg biennially), and relatively thicker fibers with high medullation of 77-88%. Medullation refers to the presence and degree of medulla at the center of the fiber, and high medullation is “undesirable” because the greater the medullation, the bigger the fiber in diameter and so less fine. LONG-WOOLLED LLAMAS: The less common (17-35%) of the two, this breed is compact, short-bodied, and the pelage has fewer guard hairs. Their wool is longer, covering the entire body, generally uniform, and soft to the touch. The fleece is heavier (2-8 + 1-1 kg), denser, and has finer fibers (26-28 microns) with medullation of 26-33%, and on the average wool coarser than the Alpaca (Vicugna pacos). Genetic studies have revealed that 40% of the Llama population shows signs of hybridization with Alpacas. Intentional hybridization has been especially common during the past 25 years both in South America and abroad with the aim of improving wool quality, fleece weight, and economic value. Unfortunately the outcome has been a major loss of pure genetic lines. Indigenous Quechua peoples in the Andes subdivide hybrids into “llamawari” (Llama-like) and “pacowari” (Alpaca-like) based upon physical appearance. Llama ears are banana-shaped (distinctively curved inward) and relatively long (14-16 cm). They are docile, intelligent, and can learn simple tasks after a few repetitions. Mature Llamas weigh an average of 140 kg with full body size reached by four years of age. There are no obvious differences between the sexes, but males tend to be slightly larger. The male prepuce is slightly bent down and directed posteriorly for urination. The female vulva is small, located immediately below the anus, with a nose-like structure pointing out from the base; the compact udderis in inguinal area with four small teats. Llamas are long-lived with a life span of 15-20 or more years. The Llama is woolly in appearance; individual fibers are often coarse, not homogeneous, and have a wide variation in diameter. Its fleece is the heaviest (1.8-3. 5 kg) of the four cameloids, but often of uneven quality. As with the other cameloids, Llama fleece lacks grease,is dry, highly hygroscopic, and naturally lanolin free. Through selective breeding and/or hybridization with Alpacas, some Llama bloodlines have finer-fibered fleeces. The typical Llama fleece is dominated by external guard hair covering (c.56% offleece with fiber diameter 50-70 microns) with an internal undercoat of smaller diameter fibers (c.44% with 25-30 microns). Llama wool is more variable in color and diameter than Alpaca wool. Due to its relative coarseness, [Llama wool has little textile value and is worth half the value of Alpaca wool. Llama woolis rougher to the touch, but with greater felting properties since the cuticle scales protrude more. However, because Llama wool is characteristically strong and warm, it is commonly used by indigenous families for making blankets, ponchos, carpets, rugs, shawls, rope, riding gear, sacks, and “costales"—bags tied to the back of Llamas and used for carrying cargo. Only c.40% of the Llama population is shorn annually because producers want heavy fleeces with long fibers to sell commercially. Some Llamas are not shorn for years because the fleece pads the back for carrying cargo. Annually there are c.1,122,667 kg of Llama wool produced in Peru (60%), Bolivia (34%), and relatively small amounts from Argentina and Chile. In Bolivia an estimated 70% is sold commercially and 30% used for home use. Although the textile industry prefers white, Llama fleeces are of different colors (47% solid, 27% mixed, 25% white). A major problem with Llama woolis its high medullation: without (20%), fragmented (37%), continuous (39%), and kemp/hair (4%). If the fleece wool is separated from different parts of the body and coarse fibers are removed, a favorable proportion of fine wool is obtained. There are no sustainable plans for genetic selection of animals with fine diameter of high value. Although the population of Llamas in Argentina is relatively low, fiber diameter is fine: 48% at 21 microns or less and only 16% at 25 microns and more. Habitat. In the Andean Altiplano where large numbers of Llamas are raised, the animals are a central part of the agro-pastoral system and the lifestyle of many people, since Llamas are heavily relied upon for carrying cargo and produce. In general these high-altitude grazing lands are low producing with annual production at 200-600 kg/ha for plains and mountainous zones and 600-2450 kg/ha for bofedales. The bofedal habitat is especially important for foraging Llamas during the dry season, yet fragile and susceptible to erosion if overgrazing is permitted. Food and Feeding. [lamas are considered by their indigenous herders to be extremely hardy because of their ability to prosper in desolate-Andean environments. They have similar feeding habits to Alpacas, but distinct enough to make joint husbandry compatible and possible. Herders view the land andits forage as a single valuable unit because it feeds their Llamas and Alpacas. Land ownership is not important, but traditional use and designated rights to graze particular areas is critical. Studies in the highlands of Peru and Chile on the botanical composition of the diets of Llamas feeding on wet (bofedales) and dry (gramadales) meadows found a high overlap with Alpaca and sheep feeding habits, but significantly differed from Alpacas in the summer (61%) and winter (74%) because the two camelids were managed by herders to minimize competition. Llamas had higher digestion coefficients than sheep of organic matter, crude protein, dry matter, and fiber fractions of bunchgrasses, important forage for Llamas. These feeding trials comparing the abilities of Llamas vs domestic sheep in digesting organic material ofvarious qualities revealed the coefficients of digestibility for low quality to be 51 vs. 41 (24% difference between the species), medium quality 60 vs. 52 (15%), high quality 73 vs. 75 (-=3%), high fiber 58 vs. 52 (12%), medium fiber 62 vs. 58 (7%), and low fiber 67 vs. 65 (3%). Thus Llamas were significantly more efficient than sheep when forage was low to medium quality and high in fiber. Maintenance energy requirements for a 108 kg Llama is 2% of its body weight, or 2-2 kg dry matter of forage per day. Breeding. The breeding season is from February to May. Males used for breeding are commonly familiar to the females, who when approached sit down in the copulatory position. Unfamiliar males usually have to chase the female and force her to recline. Research and extension agencies encourage herders to use one male with five females for three days, and then remove the male. The male is reintroduced 15 days later and the cycle is repeated until all females are bred. Gestation is 340-360 days. A single offspring is born, although exceptionally twins do occur. The female gives birth standing up, there is no licking of the neonate, and newborns can follow their mother within an hour. Llamas have a high potential for reproduction under good management: 85-95% annual reproductive rate on research and well-managed farms. However, in the Andes due to poor forage conditions frequently available to indigenous herders and the resulting poor condition of animals,fertility has been reported to be as low as 45-55%. In years of severe cold or droughts subadult mortality can be as high as 30-50%. Offspring regularly nurse up to the fifth month and are weaned by the herder at eight months, although if allowed to do so will continue to nurse irregularly until the female gives birth again the following year. Some females breed 8-10 days after parturition, but two to three weeks later is the norm. Mother Llamas are patient with suckling their young and some will accept nursing another female’s offspring. After weaning the young Llamas, some herders separate them from females until two years old, and then segregate them by sex. At three years of age a final selection is made for the best males to be sires with the balance to be used as pack animals or eventually meat. Females are bred at two and half years and not used as beasts of burden. Activity patterns. Daily activity patterns of Llamas are essentially the same as Alpacas. That is, after having spent the night in or next to a rudimentary stone corral adjacent to the family’s residence (often a single-room hut called “choza” made of stones with a thatched roof), the Llamas move out soon after sunrise to feed in the local highaltitude grasslands, moving and grazing often unattended by a herder, then return to the choza as darkness approaches. Activity budgets (percentage of time) of Llamas compared to sheep grazing on native Andean pasture dominated by favorable forage (Festuca dolichophylla), have shown that all other activities were similar, but that Llamas feed more (71% vs. 57%) and rested less (15% vs. 25%) than sheep. Movements, Home range and Social organization. [Llamas (as well as Alpacas) of South America exist within a society of indigenous herders, whose main societal features are seasonal migration, a scattered population without villages or urban centers for permanent residence, and whose social structure is centered around large families with strong ties. Rules and traditions exist within each community that often determines important aspects of herd management. For example, in some systems male [lamas are maintained apart from the female segment in distant community pastures, and then reunited for the breeding/rainy season from January through March. The female segmentis a mixed herd of reproducing and replacement females, young, and one-year-olds of both sexes. When the yearlings are 12-18 months old some herders make a preliminary selection for meat production or future reproduction. Another system used extensively is one in which breeding males are permanently kept with the mixed herds of females year-round. Invariably Llama herders also maintain a flock of sheep that offers an important source of food for the family. A problem, however, is that the sheep compete for forage with the Llamas, reduce the land’s carrying capacity, and increase the probability of overgrazing. Llama herd size averages 40-60 in the more heavily populated communities, compared to 120-180 Llamas in the Altiplano with fewer people. In contrast, average accompanying flocks of sheep average 40-70 animals in both areas. Status and Conservation. Bolivia has most (58%) of South America’s ¢.3-91 million Llamas, followed by Peru (37%), Argentina (4%), and Chile (1%); nearly all of which (99%) are found in indigenous communities. The total number of Llamas has increased 12% during the past decade. In Bolivia 370,000-500,000 families raise Llamas, in Peru 297,414, Argentina 2803, 80% of which have fewer than 90 animals. There are no known wild or feral populations. Llamas were intertwined with the rise and spread of the Inca Empire since its beginning in the Peruvian Andes in the early 1200s. With the help of Llamas, the Incas built sturdy walls, buildings, irrigation systems, and some 8700 km of roads throughout their empire. These roads eventually extended around 2500 km from Ecuador south to central Chile and parts of Argentina. Before the Spanish Conquest of Peru, Llamas numbered into the multimillions, but were severely decimated during the post-conquest period. While early Spanish chroniclers recorded the “virtual disappearance” of these animals within a hundred years, that was obviously an exaggeration. Indigenous and endemic to the South America, [Llamas have now been exported to countries around the world as a companion animal, featured in livestock shows, used for trekking and backpacking, and increasingly utilized as a guard animal for protecting sheep and goats from canid predators. In the USA there were 162,000 registered Llamas in 2010. Export of Llamas (as well as Alpacas) has increased international interest in these species, stimulating research in the medical, nutritional, reproductive, and disease disciplines. In the Andes, Llamas are viewed and described by native herders by masculine terminology. During ceremonies in which Llamas are being honored or sacrificed, they are referred to as “brothers” and when higher rank or importance is expressed, they are called “fathers.” Indigenous Llama terminology is based upon fleece color, patterns and patches, sex, reproductive status, age, size, shape, wool quality, and behavior. The combinations of these descriptive characters amount to over 20,000 words, forming a rich nomenclature used to identify and distinguish individual Llamas as well as Alpacas. Llamas have long been important beasts of burden for the Andean cultures and nations. Cargo-carrying Llamas have made it possible for peoples of the Andes to successfully inhabit this rugged-mountain environment. Extensive and ritualized prayers are said before departure of a Llama caravan, requesting permission and protection from local deities for the journey, safe passage, and that no incident will befall the Llamas or family members while traveling. Only men of the family travel with the Llama pack train, walking behind the animals. Castrated male Llamas are used primarily, and depending upon their maturity, size, and training, individuals are capable of carrying up to 25-30% of their body weight, or 25-35 kg, and traveling 20-30 km in a day for up to 20 days. The cargo is carried in a sack tied to the Llama’s back with a rope made from coarse Llama wool. Males are castrated at two years of age and training begins by accompanying the older animals on trips. Individual Llamas in the traveling caravan are not tied together, but follow the lead of two to three “Llama guides.” Such lead Llamas traditionally were adorned with colorful halters, frontal tapestries, and even a family or national flag atop its back. Animal leaders are given names to reflect their status, such as Road-Breaker, Condor Face, and Champion. Lead Llamas prevent junior animals from usurping the front position, and guide the way when fording rivers, crossing dangerous bridges, and when negotiating narrow paths along steep ravines. If the lead Llama stops, the entire caravan waits until it resumes traveling. At the end of the day, Llamas are unpacked and turned loose to graze and water. If available, they are kept in stone corrals for the night, otherwise they are grouped together with a rope running around them at neck level. Llama caravans use traditional trading circuits in the Altiplano, some descending into the Andean valleys, the Pacific coast to the west, or the Amazon jungle to the east. Goods are sold, purchased, and traded along the route as needed. From the coastal agricultural valleys fresh produce was obtained in exchange for corn, potatoes, wheat, and barley that were grown and carried down from the mountain agricultural valleys. In times past Llama trains traveled hundreds of kilometers to obtain highly prized salt from inland, natural-salt deposits and mines, some types of which were used for human consumption, others for animals. Today, the world of the Andean Llama herdsmen is rapidly changing with goods and produce now primarily transported by trucks and rail. Still, Llama caravans continue to be used and are especially important for transporting goods to remote regions of the Andes. Secondarily Llamas are used in South America for meat, fuel, and wool. Llama meatis often dried and stored as jerky, typically made by alternative freeze-drying during the Andean winter when nights are freezing and days sunny. With a dressing percentage of 44-48% yielding 25-30 kg of fresh meat, 12-15 kg ofjerky are produced. Jerky is not only a convenient form for long-term storage of meat, but is an excellent source of protein (55-60%). Llama hides are used for making shoes, ropes, and bags. Their fecal matter is valuable as fuel in regions where wood is scarce. It should be noted that despite the presence of many animals, the diet of Andean herdsmen is based upon agricultural produce. They eat little meat and drink no milk. Goat or cow cheese is consumed by some, and when meat is eaten,it is in the form of cameloid jerky or fresh sheep meat. Outside South America the Llama’s cargo-carrying ability has been discovered by hikers, hunters, and forest-work crews in North America, Europe, Australia, and New Zealand. Llamas are quiet, gentle, unobtrusive, and easy to manage. Their hardiness, surefootedness, and tr, Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Camelidae, pp. 206-246 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 236-238, DOI: 10.5281/zenodo.5719719

Family CAMELIDAE (CAMELS) • Medium-sized to large mammals with forequarters larger than hindquarters, long, small head, slender muzzle with split upper lip, thin neck, padded feet with two toes and nails rather than hooves, and woolly pelage. • 130-410cm. • Palearctic, Afrotropical, and Neotropical Regions. • Desert, semi-arid to arid plains, grassland, steppe, and shrubland; from sea level to 4800 m. • 3 genera, 6 species (including 3 domestic), 9 taxa. • 1 subspecies Critically Endangered, 1 subspecies Endangered; none Extinct since 1600., Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Camelidae, pp. 206-246 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on page 206, DOI: 10.5281/zenodo.5719719

Don E. Wilson, Russell A. Mittermeier (2011): Camelidae. In: Handbook of the Mammals of the World – Volume 2 Hoofed Mammals. Barcelona: Lynx Edicions: 206-246, ISBN: 978-84-96553-77-4, DOI: http://doi.org/10.5281/zenodo.5719719

1. Guanaco Lama guanicoe French: Guanaco / German: Guanako / Spanish: Guanaco Taxonomy. Camelus guanicoe Muller, 1776, Patagonia, Argentina. The Guanaco is a direct descendent of Hemiauchenia, a genus of camelid that migrated from North to South America three million years ago. Two million yearold fossils of L. guanicoe can be found today in Argentine Pleistocene deposits and others in strata dated 73,000 -97,000 years ago in Bolivia. Phylogenetically, L. guanicoe is monophyletic. Historically, four sub-species of Guanacos were recognized, albeit based upon incomplete information on skull measurements, coat coloration, distribution, and body size. However, recent molecular studies using mtDNA cytochrome-b sequences, recognize only two subspecies, placing the Peruvian and northern Chilean populations in subspecies cacsilensis and assigning the remainder of the clade to subspecies guanicoe. A significant biogeographic revision of the two valid subspecies is needed, especially an analysis and classification of populations on a regional and ecosystem basis. Subspecies and Distribution. L.g.guanicoeMiller,1776—Bolivia,Chile,WArgentina(fromJujuytoSPatagonia),TierradelFuego,andNavarinoI. L. g. cacsilensis Lonnberg, 1913 — N Peru to N Chile between 8° S and 22° S. Introduced from Argentina to Staats I (Falkland Is) during the late 1930s. Descriptive notes. Head-body 190-215 cm,tail 23-27 cm, shoulder height 90-130 cm; weight 90-140 kg. Measurements vary because of their wide distribution and differences between subspecies. Guanacos are extremely striking, with their contrasting colors, large, alert brown eyes, streamlined form, and energetic pace. Described by Darwin as “an elegant animal, with a long, slender neck and fine legs,” itis one of South America’s largest terrestrial mammals, reaching its maximum size in southern Chile and smallest in northern Peru. There is no obvious sexual dimorphism in size, color, or structure, except for the presence of large canines in the male. The color of the woolly pelage is similar for all Guanacos, varying from light brown with ocher yellow tones in the north to dark reddish brown in the south. The chest, front of neck, belly, and internal portion of the legs are more or less pure white, the head gray to black. Guanaco wool is prized for its softness and warmth, and second only to that of the Vicuna (Vicugna vicugna). The pelts from “chulengos” (newborns and less than one-year-old juveniles) are particularly soft. Like their domestic descendant the Llama (L. glama), the Guanaco is double-coated, with coarse guard hair (3-5% offleece) and a soft undercoat. The undercoatfibers range 12-17 (average 14-16) microns in diameter. Fiber (strand) length is 35 mm, the same as the Vicuna. Average fleece weights vary from 322 g to 350 g. Fiber diameter increases about two microns from one to seven year of age. Value of shorn, unclean wool is US $ 100-200/kg and US $ 400/kg for cleaned and dehaired wool. Guanacos have “thermal windows” in the front and rear flanks (underarms) that are nearly without wool. Both wild and captive Guanacos may live as long as 28 years. Habitat. The Guanaco inhabits environments from sea level up to 4500 m or more, characterized by highly seasonal weather, with snow cover or dry winters, cold to freezing temperatures, moderate to high winds, and low precipitation. These combine to produce high evapo-transpiration and dry conditions that lead to low primary productivity. At a broad scale, Guanacos inhabit four of the ten major habitats found in South America: desert and xeric shrublands, montane and lowland grasslands, savannas and shrublands, and temperate forests, which botanically include the categories of Puna, pre-Puna, Andean steppe, Chacoan grasslands and shrublands, Espinal, and the southern Pampa. They inhabit flatlands, hilly foothills, and mountainous environments. In the arid habitats of southern Chile, isolated mesic subsystems were the preferred plant community accounting for 85% of female sightings and 60% ofterritorial males. These “vegas” (meadows) were not only preferred for their productive, higher-quality succulent forage, but for females a major influence for their selection was the avoidance ofsites and habitats favored by Pumas (Puma concolor). Food and Feeding. The Guanaco is a non-specialized, opportunistic, intermediate, mixed-feeding herbivore, foraging on a wide variety of plants. It is basically a grazer, but also browses. When the availability of the herbaceous strata decreases or becomes unavailable, especially during winter, Guanacos feed mainly on the shrub or tree strata. This flexibility as a generalist to change diet according to availability or preference extends to eating epiphytes, lichens, fungi, cacti, succulent plants, fruit, flowers, and leaves. Their summer diet in a mixed habitat of southern Chile averaged 62% grasses (mainly Festuca), 15% browse (Nothofagusspp.), and 11% forbs, which were particularly important in the spring. In another study on Tierra del Fuego their diet was made up of 90% grasses and forbs. In austral coastal forests of southern Chile, Guanaco browsing significantly diminishes the rate of regeneration of the commercially important southern beech tree (Nothofagus pumilio) and was a limiting factor on initial growth of seedlings and saplings (89% were browsed), although the Guanaco’s diet was less than 10% shrubs and trees. At Torres del Paine National Park, at the western edge of the Patagonia, vegas were highly utilized in summer (86% ofall feeding observations) and preferred by all feeding Guanacos (n = 1659) in family groups, whereas all other vegetation types were avoided (shrub = 3%, upland = 10%). Adult females in family groups showed the greatest feeding preference for vegas, followed by chulengos, yearlings, and adult males. In Argentine Patagonia, Guanaco densities are low and negatively related to domestic sheep numbers. Guanacos and sheep largely overlap in their forage preferences, with over 80% oftheir diets being identical. Histological analysis of fecal samples revealed that Guanacos and sheep were intermediate herbivores feeding on a wide range of grasses and forbs, capable of changing their diets seasonally, and their food niches greatly overlapped particularly in summer when food resources were more scarce than in spring. Breeding. Nearly all females breed at two years of age and have their first offspring as three-year olds. At Torres del Paine, Chile males began breeding after obtaining a feeding territory when they were 2—4 years of age. Essentially all breeding took place within feeding territories during the summer. A high percentage (88%) of males established a territory for three or fewer years (average 2-3), although some males held territories for as long as eight years. Few solo males became family group males (19%), that is, 81% remained non-reproductive during their territorial tenure. Those males leaving male groups to become territorial, only 35% directly became family group territorial males while 65% became solo territorial males. Like other camelids, Guanacos are induced ovulators. The territorial mating period was from early Decemberto early January (91% of 88 observed copulations), so males defended their feeding territories for nine weeks before and eight weeks after the mating season. Mature female Guanacos give birth to a single offspring each year after a gestation of about 11-7 months (345-360 days). Only three sets of twins were documented in over two decades offield studies at Torres del Paine during which several thousand newborns were observed; in all cases no more than one survived past the first week. Half of all births occur in the last two weeks of spring. The timing of parturition varies with latitude. At Torres del Paine the birth season occurred from late November to early January. Parturition occurs during the day in Patagonia with 78% of births between 10:00 h and 14:00 h, when the young are able to dry during favorable midday temperature conditions. Birth weight averages 13 kg (7-15 kg) and shows marked density dependence, with lower birth weights at higher population densities. Low weight at birth is related to high rates of mortality. Newborns are very precocious. They can stand as early as 5-76 minutes after birth and can run within hours. Young are weaned at 6-8 months and are expelled from family groups by the territorial male when they are 11-15 months old. Dispersing yearling males join male groups and yearling females join family groups or female groups. Monitoring of 409 radio-collared chulengos at Torres del Paine over a seven-year period revealed an average first-year survival rate of 38% (31-55%). Puma predation was the primary cause of mortality of young Guanacos, especially in the first two weeks of life. Relative to their availabilities, chulengos were preyed upon about four times as much as adults. With every centimeter increase in winter snowfall, the risk of chulengo mortality increased by almost 6% because of greater vulnerability to predation. Out of 731 Guanaco skulls collected at Torres del Paine from 1979 to 1988, 33% showed clear evidence of having been killed by Pumas, and that was considered an underestimate. Observations of farmed Guanacos revealed that allosuckling (nursing of non-filial offspring) comprised 6% of all suckling events by 62% of calves and was allowed by 52% of dams. Dams whose calves performed allosuckling exhibited poorer body condition, suggesting juvenile Guanacos allosuckled to compensate for nutritional deficiencies. Preliminary research on the cytochrome-b gene sequence has found no evidence of hybridization between Guanacos and Vicunas. Activity patterns. Observations of both wild and captive Guanacos have shown peaks of eating, bedding, and ruminating in the morning and again later in the afternoon. Wild Guanacos spent a greater proportion of their time moving and less time resting, probably the result of the need to forage more and maintain intraspecific social interactions. At Torres del Paine during the summer, Guanacos in family groups in vega habitat fed 54% of time, rested 45%, and were involved in other behaviors 1% of the time (3084 focal observations). There was no difference in the activity-time budgets of 23 marked solo territorial males compared to family-group territorial males based on social group type, total number of females, total number of Guanacos present, or age of the ter ritorial males. The pattern suggested thatterritorial male behaviors were related to resource defense rather than to any direct ability to attract potential mates. Males, in all categories, spent most of their time foraging (65% of overall time budget). However, based upon habitat type there was a significant difference in time spent in aggressive and in miscellaneous activities (defecation, alertness to observer, scratching). Most aggressive encounters and miscellaneous activity occurred on hilltops of areas dominated by mata barrosa shrubs (Mulinum spinosum). Vigilance patterns were assessed in Chubut, Argentina for Guanacos occupying a tall shrubland covering 40-60% of the area, where 40% of the mortality was from Puma predation. Family group territorial males devoted more time to scanning their surroundings and less time feeding than did females, and both sexes benefited from grouping by reducing the time invested in vigilance and increasing foraging time. Males reduced the time invested in vigilance as the number of females in the group increased, while the presence of chulengos increased territorial male vigilance. However, in closed habitats collective vigilance increased with the number of adults but decreased with the number of chulengos. Although male and female Guanacos differed in their time allocation, results supported the hypothesis that both sexes received significant anti-predator benefits from group living. Adjustments in Guanaco body posture can modify the exposure of body surface area H—22%. Guanacos can decrease and increase body heat loss through radiation and convection by “closing” and “opening” their thermal windows in the axillary and flank regions. Researchers report that when ambient temperatures were 0-10°C, animals used postural adjustments to decrease the area of the thermal windows by 5-7% oftotal surface area. At temperatures greater than 10°C they increased the area of the thermal windows up to 22% to regulate heat loss through radiation and convection. When the temperature was below 0—1°C Guanacos bedded and huddled together, often with their hindquarters into the wind. Movements, Home range and Social organization. Intensively studied Guanaco populations have been either migratory or sedentary. In the San Guillermo Biosphere Reserve (Argentina), both occur but most are sedentary, while on the island of Tierra del Fuego (Chile) both occur, but most are migratory. In Argentina populations have been reported to be migratory over short distances, altitudinal-facultative migratory, and strictly sedentary. On Tierra del Fuego sedentary populations were incompletely so, because during the winter when territories were snow covered, many or all of the family group members abandoned the site, leaving the territorial male by himself or only with a few members. The following spring the male regained his group membership. Although some local populations now appear to be sedentary at Torres del Paine, in the 1970s and 1980s the population was completely migratory. In those decades essentially all animals abandoned the summer range and moved in late autumn 8-18 km to where the snow was less deep and browse species were more abundant. The following spring the animals returned to their summer range. Daily movements of family groups on Tierra del Fuego were highly predictable: days were spent in open meadows feeding and nights were spent resting in the adjacent closed Nothofagus forest. In the open habitat of Torres del Paine, the animals spent days on territories and nights on adjacent slopes or hill tops. The social organization of Guanacos is similar to that of Vicunas, except that territorial, resource-defense polygyny is seasonal instead of yearround and there is fluid movement of females between male territories. The social units of Guanacos for the migratory population at Torres del Paine were family groups, male groups, solo males, mixed groups, and female groups. Typical family groups were composed of one adult territorial male, seven females, and four juveniles less than 15 months old. Group size and composition frequently changed. Family groups occupied feeding territories 7-13 ha in size, which were defended by the resident male. With minor adjustments to the center of activity from year-to-year,territory locations were the same, spatially discrete, and non-overlapping. There was no indication of defended sleeping territories as found with Vicunas. Although family groups were “open” in the sense that females couldjoin and leave at will, the territorial male determined whether or not females were accepted into or rejected from the group. In a study of marked territorial males at Torres del Paine, most (73%, n = 60) returned to the same territorial location from year-to-year. Those males (27%) that shifted territorial locations showed no pattern in changes between solo territorial males and family-group territorial males. Male groups were composed of non-breeding, non-territorial, immature and mature males. Group size was highly variable (3-60) and averaged about 25. Male groups lived apart from family groups, in male-group zones. Solo males were mature males with an established territory, but commonly without females; the mean “group size” for solo males was three. Mixed groups formed in winter and included Guanacos of both sexes and all age classes. They averaged 60 animals with as many as 500. Female groups were gatherings of females of all ages and occasionally included a small number of immature males. These groups came together temporarily immediately before and after the winter migratory season. Female group size was highly variable, and could number 10-90 animals. At high population densities, the size of Guanaco territories can decrease significantly. Large female groups and their chulengos can shift daily from territory to territory. The annual cycle of migratory Guanacos at Torres del Paine was divisible into four general socioecological periods: summer territorial, autumn transitional, winter aggregational, and spring transitional. The summer period was the longest, from mid-October to the end of March. This was reproductive season, during which males defended territories and when birth and mating occurred. The summer territorial social units were 35% family groups, 15% male groups, 42% solo male groups, and 8% female groups. Most of the animals (65%) were in family groups; 21% were in male groups, 7% were solo males, and 3% were in female groups. The autumn-transitional period was short, lasting from early April to late May, and Guanacos were mainly in family groups and male groups. During this period the territorial system broke down as the Guanacos began migrating to their winter range. The winter aggregational period extended from early June through late August. In this period, the social units were primarily mixed groups (39%) and female groups (41%) with most of the animals (80%) in mixed groups. The spring-transitional period started in late August and ended in mid-October. At this time all social units were found with equal proportion of Guanacos in family groups and male groups. Differences in the weather from year-to-year, especially at the beginning and end of winter, caused slight variations in the timing of animal movements, formation of social groups, and migration. Snow storms and snow cover were especially important in triggering sudden movements west to the winter grounds. Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. Northern subspecies cacsilensis is recognized as Endangered on The 2006 IUCN Red List with about 4000 remaining.The Guanaco is the most widely distributed native artiodactyl in South America. It originally ranged from the Andean areas of northern Peru, south to Bolivia and adjacent parts of Paraguay and down to Tierra del Fuego, covering most of Argentina and Chile. Based upon the carrying capacity of the territory Guanacos originally occupied, the pre-Hispanic Guanaco population in southern South America has been estimated to be 30-50 million. Indiscriminate hunting and competition with sheep, particularly during the past century, caused a steep decline in numbers. In the Argentine Patagonia the introduced sheep reached 22 million head within 50 years in the late 1800s and early 1900s. Today, the best estimate of the total Guanaco population is 536,000-840,000 animals. Although still widely distributed, the Guanaco’s current distribution is less than 40% of its original range, with remaining populations often isolated and fragmented. The killing of young chulengos for their soft skins has had a serious impact on Guanaco populations, especially in Patagonia. Some 444,000 skins were exported from Argentina between 1972 and 1979. The number dropped to 10,000 annually by 1984, reflecting decreased availability. Conservation classification and laws regarding the remaining Guanaco populations vary from nation to nation, based primarily upon total numbers and without regard to subspecies. Percenta, Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Camelidae, pp. 206-246 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 235-236, DOI: 10.5281/zenodo.5719719

3. Vicuna Vicugna vicugna French: Vigogne / German: Vikunja / Spanish: Vicuna Other common names: Argentine Vicuna (vicugna), Peruvian Vicufa (mensalis) Taxonomy. Camelus vicugna Molina, 1782, Chile, “abondano nella parte della Cordigliera spettante alle Provincie de Coquimbo, e di Copiap6” (Cordilleras of Coquimbo and Copiapo in northern Chile). The separation of the Vicuna and the Guanaco (Lama guanicoe) occurred 2-3 million years ago. Palaeontological evidence suggests the genus Vicugna evolved from Hemiauchenia, a North American immigrant to South America, in the lowlands east of the Andes some two million years ago, with widespread distribution as recently as 10,000 -13,000 years ago in lowland grasslands of Bolivia, Paraguay, Patagonia, and Tierra del Fuego. Then 9000-12,000 years ago, during the last Pleistocene glacial advance and the subsequent establishment of the Holocene climate, Vicugna moved from its lowland distribution to its present day, high-elevation habitat. Today, two subspecies are recognized, distinguished on the bases of genetics, habitat, and morphology. Northern subspecies mensalis is closely related to the domestic Alpaca (V. pacos). Subspecies and Distribution. V.v.vicugnaMolina,1782—WBolivia,NWArgentina,andNEChilefrom18°Sto29°S. V. v. mensalis Thomas, 1917 — SE Peru, W Bolivia, and NE Chile from 9° S to 19° S. Ecuador has a small population (c.3000) introduced from Peru, Chile, and Bolivia in the 1980s. Descriptive notes. Head-body 125-190 cm,tail 15-25 cm, shoulder height 85-90 cm; weight 38-45 kg. The body color of the soft, woolly coat is pale cinnamon to reddishbrown, with insides of the legs and underside white. There is a bib of coarse white hairs 20-30 cm long on the chest at the base of the neck in the northern subspecies, which is short to almost unnoticeable in the southern subspecies. The body is slender with a long neck. The head is small and wedge-shaped; the ears are slender and pointed. The Vicuna has unique rodent-like incisors. Males and females weigh approximately the same, look alike, and are sexually indistinguishable in the field. Southern subspecies vicugna, sometimes called the “ Argentine Vicuna,” is ¢.15% larger (c. 45 kg vs. 38 kg), length of molars longer, taller at the withers, substantially shorter chest-bib hair, lighter colored, larger white underside countershading, and in general exhibits higher levels of genetic diversity than the northern subspecies mensalis, sometimes referred to as “Peruvian Vicuna.” Vicuna wool (often called fiber) is among the finest in the world at 12-5 + 1-5 microns (Cashmere goat fibers measure 14-19 microns; Chiru, Pantholops hodgsonii fiber is 9-12 microns). The shorn fleeces of 30,391 Vicunas in Peru averaged 220 g /animal. With its silky texture, Vicuna woolis highly prized, retailing for US $ 250/0z in the USA and in 2004 selling at wholesale for US $ 566,/kg from certified liveshorn animals. World prices for Vicuna wool ranged from US $ 350/kg to US $ 900/kg over the past decade, promoting reference to it as the “Gold of the Andes.” The adaptations responsible for the animal’s outstanding physical endurance at high altitudes include lightweight insulating fleece, which protects against cold and the sun’s ultraviolet light, and high blood-oxygen affinity (highest of all mammals investigated). Full saturation of the blood with oxygen occurs at the lower partial pressure of oxygen that is found at high altitudes. The Vicuna’s heart muscle capillary densities are exceptionally high for a mammal of its body size. Its ability to load and unload oxygen is improved by a relatively high oxygen transfer conductance because ofits small red blood cells, and it has low blood viscosity due to a relative low percentage of red blood cells. Both are advantageous for maximum cardiac output. The Vicuna’slife span is 20 years, with a maximum age of 24 years and nine months recorded in captivity. Habitat. Vicunas are restricted to the Puna and Altoandina biogeographic provinces of the Andes.The Vicuna is the highest-altitude ungulate in all of South America, living in a unique montane zone from 3200 m to 4800 m above sea level called the Puna, a high-altitude, equatorial grassland that is above the tree line but below the snow line. Summer precipitation is typically in the form of rain or hail, rarely snow. It is a dry and cold environment with summer nights hovering close to freezing. Winter nights plummet well below freezing, 10-20°C below the daytime highs. Two distinct habitats with different levels of precipitation can be found within this dry-Andean ecosystem: the high-elevation moist or semi-humid high-Andean Puna and the lower elevation dry or semi-arid Puna habitat. The dry Puna is an extremely dry belt called the Andean Dry Diagonal, a north-west/south-east transition zone between two major hemispheric wind belts centered at the junction of north-west Argentina, south-west Bolivia, and north-east Chile. Within the Dry Diagonalthere is essentially no precipitation, no lakes, and no glacier formation. The distribution of subspecies vicugna is within the Dry Diagonal. Subspecies mensalis is found to the north of it. From a landscape perspective the Puna is characterized by peaks and pronounced slopes, the typical elevated plain that defines the region (Puna or Altiplano), and the intermediate piedmont fringes of smooth slopes. Vicunas use the habitats within the Puna and piedmont zones. The most common habitats in the Puna are xerophytic shrub steppes that are often mixed with an understory of sparse short grasses and forbs (typically not an important habitat for Vicunas); bunchgrass steppes (variable importance); open rocky areas with sparse vegetation (not important); short grass and forb areas on lower slopes, gentle slopes, and plains (important); and wetlands of short plants with high ground cover associated with high ground water, surface water, streams, and lagoons, regionally called bofedales, mojadales, and vegas (important). Bofedales (around 4500 m elevation in Peru) are perennially green sedge communities typically dominated by Eleocharis albracteata and Carex ecuadorica with vegetative cover of ¢.22% grasses, 42% sedges and reeds, and 33% forbs, and a crude protein content of 12%. Short-grass forb areas on lowerslopes (around 3190 m in Peru) are dominated by the grass species Festuca dolichophylla and Muhlenbergia fastigiata with a vegetative cover of 66% grass, 13% sedges and reeds, and 6% forbs, and a crude protein content of 10%. Food and Feeding. As with other ungulate herbivores, Vicuna habitat and forage selection is based not only upon availability, but heavily influenced by climate (wet vs. dry years), population density (high vs. low), and the social status and ranking of individual animals (high vs. low). Thus for studies done at different locations, landscapes, and animal densities,it is not surprising to see variable results. For example in Peru at the Pampa Galeras National Reserve, Vicunas were observed to be strict grazers on grasses and forbs, whereas in Argentina at the Laguna Blanca National Park, Vicunas showed some diet plasticity by being grazers on grasses 16-19% ofthe time and browsers on shrubs 59-72% of the time with two grasses (Panicum chlorolewcum and Distichlis spp-) representing nearly 50% of the diet. Feeding preference in productive habitats dominated by grasses has been documented in several areas of the Vicuna’s distribution. Of the two main habitats (grass steppe and vegas) at the Ulla-Ulla Vicuna Reserve in Bolivia, Vicunas preferred vegas. In the San Guillermo Biosphere Reserve of Argentina Vicunas were more abundant on the grassland plains. In Pampa Galeras National Reserve Vicunas preferred to forage in grass steppe communities characterized by Festuca nigesens or by Calamagrostis vicunarum, the latter being the dominant species in the vegetative type known as Excrement Influenced Vegetation (EIV). EIV is the result of long-term defecation-urination by Vicunas on dung piles; the resulting fertilization and accumulation of organic matter accelerates soil and plant succession over a long period of time. The third sere is an isolated “putting-green,” an island of short, abundant plants in rocky terrain downhill from a single or series of dung piles. The top soil is deeper and the plants grow thickly, close to the ground. EIV covered 18% of the total surface area of Pampa Galeras National Reserve, andits third sere had the highest productivity and was the most preferred forage vegetation. The general pattern of Vicuna habitat selection for grass-dominated communities, especially those of high nutritional value, is consistent with the results of diet studies that found Vicunas forage mainly on grasses (Poaceae) and grass-like sedges (Ciperaceae). When Vicunas coexisted with domestic livestock (sheep, goats, donkeys) in Argentina, they were spatially segregated, had a high overlap in diets, and used preferred habitatless than expected. The native Vicuna and exotic domestic animals coexisted because Vicunas were able to utilize suboptimal habitats while herders kept livestock in the richest habitats. A sympatric population of Vicunas and Guanacos in the San Guillermo Biosphere Reserve, Argentina, revealed that the diet of both species was similar, but Guanacos utilized more tall grasses and Vicuna fed more on forbs and short grasses. Breeding. Vicunas are strong seasonal breeders. Males reach sexual maturity at 3-5 years of age, and become reproductive after obtaining a feeding territory and recruiting females. Most females first mate as two-year-olds. Breeding is almost an unnoticeable event except for a short chase before mating, because the prolonged copulation takes place with the mating pair in sternal recumbency. Females show no unusual behavior to indicate they are in estrus and males do not persistently smell the female’s hindquarters as is common in other ungulates. There is no prolonged preoccupation or tending that ungulate males traditionally show for females in heat, because a territorial male has exclusive and immediate access to females within his territory. In Pampa Galeras National Reserve mating was observed from the end of February to the end of May, but peaked in April (3 matings/100 hours of observation), followed by May (1-8) and March (1-3). Almost all matings were in the morning and within the family group’s feeding territory. Gestation averages c.11-3 months (330-350 days). Females within a few weeks of parturition are visibly pregnant. At the start of the birth season at Pampa Galeras over a three year period, a high percentage of females two years of age or older were pregnant: 85%, 95%, and 85%. The non-pregnant females were mostly two-year-olds because only 20% of one-year-olds mated. Rectal palpation by experienced veterinarians of captive adult Vicunas in Puno (Peru) revealed 99% of females pregnant. Most (90%) births occurred from the last week of February through the first week of April with 75% of the births in March. February is the reported peak birth season for subspecies vicugna in Argentina and Chile. Nearly all (96%) births occurred in the morning and within the feeding territory. Females give birth to one offspring, with twins not reported or observed. Morning births are likely a behavioral adaptation to the stormy weather typical of Andean afternoons. In summer and early autumn, daily storms regularly start around noon. Young born during the afternoon storms of rain and hail would not be able to dry their soft-insulating wool before entering the nearfreezing nights. Wet newborns would have little chance for survival. A female aboutto give birth separated herself 15-25 m from the group. During the usual one-hour labor the female looked back to inspect her hindquarters, laid down and stood intermittently, changed positions, and walked a few steps every few minutes. Parturition occurred while the female stood. The labor and birthing behaviors were subtle and not easily noticed, as females commonly continued to feed during the process— surely to avoid attracting potential predators in this wide-open habitat. The mother immediately smelled and inspected the head and muzzle of the newborn, but licking did not occur (thus no assistance with the newborn’s thermoregulation) nor was the afterbirth eaten (the latter two behaviors common in other ungulates). Other members of the group frequently came over to smell and inspect the newborn. Newborns weighed 4-6 kg, were on their feet and able to walk in a wobbly way 15-20 minutes after birth, and began nursing when they were 30-45 minutes old. Vicuna young are “followers” as opposed to “hiders” and stay close to their mothers, especially in the first few months. Most young are weaned during the dry season in July and August, when they are 4-6 months old. During the first four months after birth 10-30% of newborns died. The cause was undetermined, but predation by Culpeos (Pseudalopex culpaeus) and Pumas (Puma concolor) was suspected. Reproductive success as measured in August by the ratio of young to adult females ranged 35-70 young per 100 females in different regions of the Pampa Galeras National Reserve. In Chile, density dependency seemed to be operating: the more females in a family group the lower the number of young per female. Very little research has been conducted on the physiology of Vicuna reproduction, but a noteworthy study on reproduction in captive male Vicunas in the Puna of northern Chile found higher levels of plasma testosterone, large testessize, greater size of seminiferous tubules, and large diameters of Leydig cell nuclei beginning in February, a month before the summer breeding season started. In the winter month of August spermatogenesis was in the regressive phase. Both findings suggested photoperiod as the mechanism for when males are sexually effective. Insight can be gained from studies on the Vicuna’s domesticated descendent, the Alpaca: an experimental study on Alpacas demonstrated that the continuous association of females and males exerted an inhibiting effect on male sexual activity. After continuous association of two weeks, there was no mating activity by the male even though receptive females were present. However, when mature males were separated from females and reunited at two week intervals, mating occurred. These observations shed light on why nonpregnant Vicunas (and other cameloids) are not bred during non-breeding seasons of the year. Activity patterns. During a year-round, 3-5yearfield study in Pampa Galeras National Reserve, home to a population of the northern subspecies mensalis, Vicuna family groups showed a predictable daily activity pattern in the large open valleys they occupied. Having spent the night on the upperflat ridges in small sleeping territories, within an hour after sunrise they moved as a group down to their daytime feeding territories. They spent the day on these lower slopes or flat plains, feeding at a constant rate. The sleeping territories, on higher ground, were warmer at night than feeding territories and were away from the streambeds where nocturnal predators hunted. The adult male Vicuna in the family group actively defended both territories by challenging and chasing away trespassing Vicunas. If human disturbance did not cause an early retreat back up the slope, the group returned to its sleeping territory in midto late afternoon. In the winter dry season, when small springs on the slopes dried up, groups left their feeding territory and moved to the center of the valley floor to drink from streams. Solo males holding a territory showed similar patterns. Male groups moved randomly and unpredictably through the area. In a separate study at Pampa Galeras National Reserve conducted during the March—-May breeding season, females spent significantly more time grazing than males (54 vs. 45 minutes/hour) and lactating females fed more than non-lactating females (56 vs. 53 minutes/hour). In general females fed 91% of the time; territorial males fed 75% of the time. In Salinas y Aguada Blanca National Reserve above Arequipa, Peru, there was a positive correlation between frequency of alert behavior per adult and the number of offspring in the family group, and at the Laguna de Pozuelos Biosphere Reserve in Argentina, solitary Vicunas spent more time standing and being vigilant and less time foraging than did members of groups. In Abra Pampa (Argentina), territorial males spent more time being vigilant and less time feeding as the size of their family group increased, but not more time in territorial defense (walking, running). At the Laguna Blanca Reserve in Argentina, which is extremely arid, daily activity patterns during the dry season were similar to the patterns of family groups at Pampa Galeras in the same season, with morning feeding on slopes, midday movement to a river on the flat, and a return to the slope and more feeding in the afternoon. At Pampa Galeras there was no significant difference between the amount of time males spent in territorial behavior and the number of female group members or the total group size. Studies at Laguna Blanca on the other subspecies, however, found a substantial decrease in amount of time males spent foraging when the number of females in their group increased. Movements, Home range and Social organization. All Vicuna populations of both subspecies studied to date were sedentary and non-migratory. However, as described above, daily activity patterns and movements are greatly influenced by the need to drink water. Several studies have shown that water distribution and availability can have a major effect on local movement of Vicunas. Despite being well adapted to living in arid conditions, the Vicuna is an obligate drinker and needs to drink often, especially in the dry season when it seeks water daily. Visits to small streams often include water bathing accompanied by dust bathing in dust bowls. The home range of a family group encompasses the area where the group is typically found, thatis,its sleeping territory, feeding territory, and the space in between, as well as those areas it commonly moves through during the dry season on its way to drinking water. The two territories are defended by the male and exclusively occupied by the family group, but the balance of the home range is not;it overlaps and is shared with other groups. These are neutral zones, especially the corridors that groups move through on their way to water. The social units ofVicuna are family groups, male groups, and solo males. A characteristic family group is composed of one adult male, 3-4 females, and two juveniles. Although the permanent territorial family group is the classic Vicuna social unit occupying good habitat containing semi-permanent water, other groups found within Vicuna populations may include marginal territorial family groups occupying secondary habitat types without water, and temporary mobile family groups that lack a territory. Permanentterritorial family groups occupy a year-round feeding territory averaging 18-4 ha (2-56 ha), and sleeping territory averaging 2-6 ha. This is highly unusual for an ungulate. Territorial males defend their sites passively (standing guard near borders) and actively (walking towards, chasing, fighting, biting, etc.) multiple times daily. The territorial system is based upon resource-defense polygyny in which the territorial male defends food resources esse, Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Camelidae, pp. 206-246 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 238-240, DOI: 10.5281/zenodo.5719719

Don E. Wilson, Russell A. Mittermeier (2011): Camelidae. In: Handbook of the Mammals of the World – Volume 2 Hoofed Mammals. Barcelona: Lynx Edicions: 206-246, ISBN: 978-84-96553-77-4, DOI: http://doi.org/10.5281/zenodo.5719719

6. Dromedary Camel Camelus dromedarius French: Dromadaire / German: Dromedar / Spanish: Dromedario Other common names: Camel, Arabian Camel, One-Humped Camel, Single-Humped Camel, Ship of the Desert Taxonomy. Camelus dromedarius Linnaeus, 1758, “Habitat in Africae desertis arenosis siticulosis.” Restricted to “deserts of Libya and Arabia” by Thomas in 1911. This species is monotypic. Distribution. A species found in the arid and semi-arid regions of N Africa to the Middle East, and parts of C Asia. A sizeabledfreeranging/feral population in C&W Australia. The Dromedary overlaps with the domestic Bactrian Camel (C. bactrianus) in Turkey, Afghanistan, Iran, India, Kazakhstan, and Turkmenistan. Dromedaries are a domestic species with c¢.50 breeds selected and used for pulling carts, plowing, lifting water at wells, carrying packs, milk production, smooth riding, and racing. The breeds include those in Saudi Arabia (Mojaheem, Maghateer, Wadah, and Awarik), India (Bikaneri, Jaisalmeri, Kachchhi, and Mewari), Pakistan (Marecha, Dhatti, Larri, Kohi, Campbelpuri, and Sakrai), and Turkmenistan (Arvana). The evidence for domestication comes from archaeological sites dating ¢.4000-5000 years ago in the S Arabian Peninsula with the wild form becoming extinct ¢.2000-5000 years ago. No non-introduced wild populations exist. In Asia Dromedaries occur from Turkey to W India and N to Kazahkstan. All camels in Africa are Dromedaries, 80-85% in the Sahel and NE portion of the continent (Somalia, Sudan, Ethiopia, and Kenya), with the S distribution limited by humidity and trypanosomiasis. Dromedaries in S Africa show no evidence of loss of genetic diversity within 16 populations and very low differentiation among populations. In Kenyan Dromedaries two separate genetic entities have been identified: the Somali and a group including the Gabbra, Rendille, and Turkana populations. In India two distinct genetic clusters have been described for Dromedaries: the Mewari breed being differentiated from the Bikaneri, Kutchi, and the Jaisalmeri breeds. From the 17" to the early 20" century unsuccessful attempts were made to introduce Dromedaries to the Caribbean, Bolivia, Peru, Colombia, Brazil, Namibia, and south-western USA. Successful introductions of camels were made to the Canary Islands in 1405 and some 10,000 to Australia from 1840 to 1907. Camels were important for exploring and developing the Outback of C&W Australia, where they were used for riding; drafting; transporting supplies, railway, and telegraph materials; and as a source of meat and wool. Most (6600) introduced Dromedaries came from India. Three breeds were originally introduced: camels for riding from Rajasthan, India, camels for heavy work from the Kandahar region of Afghanistan, and camels for riding and carrying moderate cargo loads from Sind, Pakistan. The camels in Australia today are a blend of these original imports. By the 1920s, there were an estimated 20,000 domesticated camels in Australia, but by 1930, with the arrival of rail and motor transportation, camels were no longer needed and many were released to the wild. Well suited to the Australian deserts, the camels bred prolifically, spreading across arid and semi-arid areas of the Northern Territory, Western Australia, South Australia, and into parts of Queensland, and today they occupy 37% of the continent. Descriptive notes. Head-body 220-340 cm, tail 45-55 cm, shoulder height 180-200 cm; weight 400-600 kg. Males and females of near equal size, but in some breeds the females are ¢.10% smaller than males. Body is often sandy colored, but can range from nearly all white to black or even two-colored piebald. Body shape characterized by a long-curved neck, long and thin legs, and deep narrow chest. The hindquarters are less developed than the weight-bearing front legs. Large eyes are protected by prominent supraorbital ridges. Facial features include thick eyebrows, long eyelashes, and transparent eyelids that allow partial vision when the eyes close in sandstorms. Thick fine hair in winter for warmth sheds in summer. Hair is longer on throat, shoulder, and hump. The single hump, on the middle of the back (c. 20 cm higher than shoulder)is a reservoir offatty cells bound by fibroustissues, used in times of food and water scarcity. Hump size varies, depending upon an individual's nutritional status. In a state of starvation, the hump can be almost non-existent. The head is small relative to body size. Slit-like nostrils, surrounded by sphincter muscles, can close to keep out dust and sand. Split upperlip with two independently moving halves and a pendulous lowerlip allow for prehensile-like grasping of forage. Upper middle and inner incisors are replaced by a tough dental pad that opposes the lower incisors. Canines, especially the upper, are massive and pointed. Skin is tightly attached to underlying tissues and modified into horny pads at the sternum, elbows, carpals,stifles, and tarsals: these protect the body when a camelis lying down on hot or rough ground. No facial glands, but males have well-developed occipital glands 5-6 cm below the nuchal crest on either side of the neck midline. The glands increase in size with age, and during the rut they secrete a pungent coffee-colored fluid. Small oval erythrocytes may enhance blood circulation and oxygen carrying capacity. Dromedaries are digitgrade each with two dorsal nails and padded feet well adapted for sandy substrates; the front feet are larger than the hindfeet. The mammary gland has four quarters and teats. Adult dental formula: 11/3,C1/1,P 3/2, M 3/3 (x2) = 32 with permanent lower incisors appearing at 2:5—6-5 years and all teeth emerged by eight years. A triangular bone ¢.3 x 2: 5 cm is lodged in the tendinous fibers in the center of the diaphragm, preventing compression of the interior vena cava and distributing muscular pull over a larger surface. Lungs are not lobed. The stomach is complex, with three compartments. When foraging on green and moist plants Dromedaries do not require drinking water. If wateris available in summer, they will drink regularly at dawn. In extreme drought, they need access to waterholes. The Dromedary’s ability to endure severe heat and dryness does not depend upon water storage; instead, numerous mechanisms minimize water loss. In well-watered animals body temperature fluctuates only c.2°C. When necessary, water conservation is aided by heat storage (hyperthermia): camels do not sweat until body temperature exceeds 41-42°C, thus avoiding water loss through perspiration. The body temperature of camels deprived of water can fluctuate as much as 6°C by heating up during the day to 41°C and then cooling at night to 35°C. Dehydrated Dromedaries have a depressed rate of breathing, minimizing water loss through respiration. Paired, fluid-producing sacs connecting the nasal cavities and a pair of lateral nasal glands and sacs serve to moisten incoming dry air. Dromedaries can tolerate water loss greater than 30% of their body mass, whereas 15% lossis lethal for most mammals. Such water loss is from intraand intercellular fluids, and not from plasma, allowing for relatively constant circulation of blood and maintaining the ability to cool. Water loss is ¢.50% greater in shorn compared to unshorn camels. Dromedaries often go without water in the Sahara Desert for seven or eight months, beginning in October, existing only on water content of plants. At temperatures between 30°C and 50°C they can go without water for 10-15 days, and even in the hottest weather need water only every 4-7 days. They can quickly rehydrate by drinking large quantities of water (10-20 1/minute and up to 130 1/minute), consuming up to 30% of their body weight within minutes. Dromedaries can drink salt water in even greater concentrations than seawater. They can consume water containing 19,000 ppm (parts-per-million) in dissolved salt without a decline in condition, compared to sheep, which can consume water at 10,000 ppm and cattle 5000 ppm. Dehydrated camels excrete less fecal water, greatly reduce urine volume, highly increase urine concentration, and recycle urea from the kidneys to the rumen for protein synthesis and water recirculation. Their erythrocytes have high osmotic resistance and can swell to 240% of their initial size without hemolysis during rehydration. Accumulation of fat in the hump instead of subcutaneously facilitates heat dissipation. The gallbladderis absent. The dulla, a pink, tongue-like bladder that hangs out the side of mouth of rutting-agitated males,is actually an inflation ofthe soft palate and unique to Dromedaries. Dulla inflation is typically accompanied by large amounts of saliva foam and gurgling vocalization. Habitat. In Africa Dromedaries occupy the Sahara Desert, known forits long, hot-dry season and a short rainy season. In Australia, Dromedaries favor bushy semi-arid lands and sand plains because of the availability of year-round forage, and avoid heavily vegetated and hard rocky areas. Food and Feeding. Dromedaries are capable of surviving on poor-quality forage under arid conditions, aided by their ability to select high-quality plant species, increase digestion of low-quality forage, cover large distances while foraging, and diversify the nature of their diet by being both browsers (shrubs and trees) and grazers (forbs and grasses). In the Sahara browse and forbs make up 70% oftheir diet in winter and 90% in summer. Over 300 forage plants have been reported, with Acacia, Atriplex, and Salsola common in their diet. In Syria shrubs dominated the diet during the dry season, but camels switched mainly to herbaceous species with the onset of the wet season. In Australian deserts food intake by volume is 53% browse, 42% forbs, and 5% grasses. Dromedaries browse on trees and tall shrubs up to 3-5 m by grasping with their lips and either breaking off branches or stripping leaves. Under extreme cases of limited forage, the Dromedary can not only decrease its food intake, but also reduce its metabolic rate. Compared to sheep and cattle, Dromedaries require less energy for maintenance; their protein requirements are at least 30% lower than cattle, sheep, or goats. Feeding trials have revealed that Dromedaries utilized fed energy for maintenance with an efficiency of 73% comparable to sheep, and for growth with an efficiency of 61% better than sheep and cattle. The relationship of food intake to body size is low. They can live on only 2 kg of dry matter for limited periods, and 8-12 kg are sufficient for a working Dromedary carrying 130-227 kg load for six hours a day at a speed of 5 km /h for a 24day trip. When forage conditions are lush, camels tend to overeat for their immediate needs and store the excess energy in their humps. Dromedaries require six to eight times more salt than other animals, with 30% oftheir diet from halophytes (plants that tolerate and even require salty conditions). High salt intake is imperative for alimentary absorption of water by camels, with salt deficiencies leading to cramps and cutaneous necrosis. Although consumption of grain can cause indigestion in animals unaccustomed to it, working Dromedaries require 2 kg of grain per day. Breeding. The breeding season is variable depending upon latitude and climate patterns. Dromedaries typically breed in winter, except near the Equator, where there can be two mating seasons or even year-round mating. In the Arabian camel, sexual receptivity is triggered by rainfall and subsequent availability of forage. There is follicular activity in the female year-round, but it peaks in winter and spring. Mating induces ovulation, which occurs 30-40 hours afterwards; estrus ceases three days later. An unmated female's cycle averages 28 days;follicles mature within six days, are maintained for 13 days, and regress over eight days. The percentage of females that conceived: 50% after a single copulation, 30% after two, and 20% after three or more, during the first two days of estrus. Left and right ovaries are equally active and alternate in follicle production. Egg migration is common, 50% of left-horn implants form corpora lutea in the right ovary, explaining the long oviductal transport time of six days. Simultaneous ovulation from both ovaries occurs 14% of time, but twin pregnancies only 0-4%. The left horn of the uterus is larger than the right and carries 99% of pregnancies. Normal pregnancy produces one offspring, with twins being extremely unusual. The scrotum is high in the perineal region with testes larger during rut. Penis is covered with a triangular sheath opening pointing posteriorally and directed between the hindlegs. A complete separation of the penis from preputial adhesions prevents erections at 6-10 months before sexual maturity. Female lies down in sternal recumbency during copulations averaging 8-120 minutes involving 3-5 ejaculatory pulses by the male, each stimulated by intracervical pressure on his highly mobile urethral process. Mating females normally ruminate; the male may salivate, inflate his dulla, or gurgle during mating. Gestation averages 377-390 days (range of 360-411 days), regardless of whether the calf is male or female. The typical calving interval is 2-3 years (two in Australia), with estrus occurring 4-5—-10 months after parturition. The mean birth massis 37-3 kg (26-4-52-3) with no difference between sexes. Annual calving rates are low (35-40%) because of high (18-20%) embryonic death, abortions, and stillborns. In free-ranging herds, young remain with their mothers for first two years. Males begin rutting at three years but are not fully active sexually until they are 6-8 years old. They continue to breed until 18-20 years of age. Females are sexually mature at three years and typically first mate at 4-5 years and reproduce until they are 20-25, and some until the age of 30. Puberty is delayed by inadequate body weight caused by insufficient food. Birthing duration is typically 30 minutes, with the female in a sitting position. The mother noses and nibbles, but does not lick her newborn. The main birth season in Australia is June to November, during the rut, but newborns have been observed year-round. Before parturition cows segregate themselves (without their previous twoyear-old calf) from their original group and give birth in seclusion in dense vegetation. Isolation thought to be both an anti-predator behavior and perhaps more significantly, avoidance of infanticide by rutting males. After remaining alone for up to three weeks, when the new calf is fully mobile, the female joins other recent new mothers, forming a new cow group. These core groups remain stable until the calves are weaned at 15-18 months, the length of time depending on environmental conditions. Life span of wild/feral animals is 20-35 years. Domestic Dromedaries live substantially longer, with reported maximum longevity 40-49 years. The first cloned camelid was a Dromedary Camel achieved in 2010 by use of somatic cell nuclear transfer. Activity patterns. Wild/feral populations exist only in Australia, where they show a daily pattern of feeding in the morning and afternoon hours and increased resting in the middle of the day. Midday resting is highest in winter. Elsewhere Dromedaries are domestic and intensely managed and regulated by traditional pastoral communities, often in conjunction with other livestock. In the Sahara, when they are allowed to roam without herders, they form stable groups of 2-20 animals. Dromedaries graze for 8-12 hours per day and then ruminate for an equal amount of time. When forage is especially poor they spread out over large areas and break up into units of 1-2 individuals. Guarded herds feed by day (lying down during the hottest hours) and rest by night, but unguarded, their activity pattern is reversed. Movements, Home range and Social organization. Dromedaries are extremely mobile and capable of using large areas to fulfill their nutritional needs. Depending upon environmental and social parameters, wild/feral populations of Australia may be nomadic, migratory, or move within a home range. They commonly travel 30 km /day even when food is plentiful. In summer, when plants are dry, they comfortably walk up to 60 km to waterholes every second or third day; in winter they drink water only irregularly, some once per month, others less often. In free-ranging Australian populations, social groups are core cow groups, breeding groups, male groups, and solitary males. Core cow groups occur only in summer (October to March/April) outside the breeding season. These groups of about 24 animals consist of females and their calves of similar age; the groups are stable for up to 1-5-2 years until the young are weaned. Summer cow groups are open to all other individuals (cows with and without calves, including younger and weaker adult males). Individuals join for irregular periods of time. Breeding groups are seen in winter (April/May to September). They are composed of one mature male and several cows with their calves; the male defends the females against other males in a classical harem arrangement. Soon after taking over a cow group the rutting male aggressively chases away weaned two-year-old males; these young males join male groups. The rutting male herds cows for 3-5 months, leaves voluntarily, and does not return to the same cow group the following year; thus the rutting male is never the father of the calves in his group. Male or bachelor groups of up to 30 non-breeding males of all ages are present year-round. These are loose groupings that regularly split up as individuals leave the group and join other males. Solitary males tend to be old males. A rutting male shows ritualized postures and patterns, including vigorous biting when fighting with and defending his breeding group against other males. It should be noted that aggressive spitting, as observed in Bactrians and the South American cameloids has not been observed in Australian Dromedaries. However, Dromedaries may vomit when severely frightened or overly excited. Night time hypothermia in a rutting male may increase the duration and success of his daytime fighting before the male overheats. No classical territoriality has been observed in Australia, but short-term home ranges of 50-150 km* and an annual range, commonly of 5000 km?, shows a tendency forsite attachment to home ranges. Dromedaries show amazing plasticity of social organization with extremes in environmental conditions. During two years of extremely high rainfall in the Australian Outback when food productivity was extraordinarily high, animals coalesced into large herds of up to 200. During the rutting season the herd was subdivided into several breeding groups, each with one herding male, all roaming around together. The subgroup holders tolerated each other to a certain point within the big herd and even showed some cooperation in defending their cows when intruding bachelors came too close. However, during two years of virtually no rainfall (although water was always available for drinking) when food became acutely sparse, normal cow groups split up, even to the extreme of only one mother with her calf. In such harsh droughts conspecifics became each other’s strongest competitors. A social system similar to that seen in Australia occurs in Africa at Equatorial latitudes, except the non-breeding season is in winter. Mixed herds (males and females of all age classes), some as large as 500 camels, are more common. In Algeria domestic herds were much less rigid: 46, Published as part of Don E. Wilson & Russell A. Mittermeier, 2011, Camelidae, pp. 206-246 in Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona :Lynx Edicions on pages 245-246, DOI: 10.5281/zenodo.5719719

Vicugna Lesson, 1842. Nouv. Tabi. Regn. Anim. Mammifères, p. 167. TYPE SPECIES: Camelus vicugna Molina, 1782., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189, {"references":["Molina, G. I. 1782. Saggio sulla storia naturale del Chili. Stamperia di S. Tommaso d'Aquino, Bologna."]}

Camelus Linnaeus, 1758. Syst. Nat., 10th ed., 1:65. TYPE SPECIES: Camelus bactrianus Linnaeus, 1758., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 381, DOI: 10.5281/zenodo.7359189, {"references":["Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth ed. Vol. 1. Laurentii Salvii, Stockholm, 824 pp."]}

Camelus dromedarius Linnaeus, 1758. Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: "Africa," deserts of Libya and Arabia (domesticated stock). DISTRIBUTION: Extinct in the wild; first domesticated about 4,000 yr BP from wild populations which had become restricted to the S Arabian Peninsula; domesticated from Senegal and Mauritania to Somalia and Kenya, throughout N Africa, the Middle East, Arabia, and Iran to NW India; feral populations in Australia. COMMENTS: Produces fertile hybrids with bactrianus (see comments therein). Bohlken (1961) considered dromedarius a synonym of bactrianus. Reviewed by Köhler-Rollefson (1991, Mammalian Species, 375). Biology reviewed by Gauthier-Pilters and Innis Dagg (1981). For history of domestication, see R. T. Wilson (1984)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 381, DOI: 10.5281/zenodo.7359189, {"references":["Bohlken, H. 1961. Haustier und zoologische Systematik. Zeitschrift fur Tierzuchtung und Zuchtungbiologie, 75: 107 - 113.","Kohler-Rollefson, I. U. 1991. Camelus dromedarius. Mammalian Species, 375: 1 - 8.","Gauthier-Pilters, H., and A. Innis Dagg. 1981. The camel: Its evolution, ecology, behavior and relationships to man. University of Chicago Press, Chicago, IL, 208 pp."]}

Lama pacos (Linnaeus, 1758). Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: Peru (domesticated stock). DISTRIBUTION: Domesticated in S Peru, W Bolivia. SYNONYMS: lujanensis. COMMENTS: Often regarded as a synonym for glama; see Corbet and Hill (1991:126). Probably originated from hybrids between Lama glama and Vicugna vicugna; see Hemmer (1990)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189, {"references":["Corbet, G. B., and J. E. Hill. 1991. A world list of mammalian species. Third ed. British Museum (Natural History) Publications, London, 243 pp.","Hemmer, H. 1990. Domestication. The decline of environmental appreciation. Cambridge University Press, Cambridge, England, 208 pp."]}

Peter Grubb (1993): Order Artiodactyla. In: Don E. Wilson, DeeAnn M. Reeder (Eds): Mammal Species of the World (2nd Edition). Washington and London: Smithsonian Institution Press: 377-414, ISBN: 1-56098-217-9, DOI: http://doi.org/10.5281/zenodo.7359189

Vicugna vicugna (Molina, 1782). Sagg. Stor. Nat. Chile, p. 313. TYPE LOCALITY: Chile, cordilleras of Coquimbo and Copiapo. DISTRIBUTION: S Peru, W Bolivia, NW Argentina, N Chile. STATUS: CITES - Appendix I, some Chilean and Peruvian populations Appendix II; U.S. ESA - Endangered; IUCN - Vulnerable. SYNONYMS: elfridae Krumbiegel?;/ro„fos«, gracilis, mensalis, minuta, pristina, provicugna., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189

Lama guanicoe (Müller, 1776). Linné's Vollstand. Natursyst. Suppl., p. 50. TYPE LOCALITY: Chile, Andes of Patagonia. DISTRIBUTION: Cordilleras of the Andes: S Peru, Bolivia, Argentina, and Chile; Patagonia and Tierra del Fuego (Chile and Argentina); Navarino Isl (Chile). STATUS: CITES - Appendix II. SYNONYMS: cacsilensis, huanacus, loennbergi, mesolithica, molinaei, voglii. COMMENTS: Has previously been included with the Llama, L. glama, of which it may be the wild ancestor, see Lydekker (1915). See also Hemmer (1990)., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 382, DOI: 10.5281/zenodo.7359189, {"references":["Hemmer, H. 1990. Domestication. The decline of environmental appreciation. Cambridge University Press, Cambridge, England, 208 pp."]}

Lama glama (Linnaeus, 1758). Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: Peru, Andes (domesticated stock). DISTRIBUTION: Domesticated in S Peru, W Bolivia, NW Argentina. SYNONYMS: ameghiniana, araucanus, castelnaudi, chilihueque, cordubensis, crequii, ensenadensis, intermedia, lama, moromoro, peruana., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 381, DOI: 10.5281/zenodo.7359189

Camelus bactrianus Linnaeus, 1758. Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: "Bactria" (= Uzbekistan, Bokhara) (domesticated stock). DISTRIBUTION: Exists in the wild only in SW Mongolia, Kansu, Tsinghai, and Sinkiang (China); domesticated in Iran, Afghanistan, and Pakistan, north to Kazakhstan, Mongolia, and China. STATUS: U.S. ESA - Endangered; IUCN - Vulnerable. SYNONYMS: ferus. COMMENTS: Includes ferus Przewalski, 1883, based on wild specimen; bactrianus Linnaeus, 1758, has priority. Produces fertile hybrids with dromedarius. Though their distributions merge, breeding is regulated (as all individuals are domesticated in the zone of contact). Corbet (1978c:197), citing A.P. Gray (1954), stated that male hybrids are sterile., Published as part of Peter Grubb, 1993, Order Artiodactyla, pp. 377-414 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 381, DOI: 10.5281/zenodo.7359189, {"references":["Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classis, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Tenth ed. Vol. 1. Laurentii Salvii, Stockholm, 824 pp.","Corbet, G. B. 1978 c. The mammals of the Palaearctic region: A taxonomic review. British Museum (Natural History), London, 314 pp.","Gray, A. P. 1954. Mammalian hybrids: A checklist with bibliography. Commonwealth Agricultural Bureau, Farnham Royal, United Kingdom, 262 pp."]}

Family Camelidae REVIEWED BY: I. U. Kohler (IUK); R. G. Van Gelder (RGVG); B. R. Stein (BRS). ISIS NUMBER: 5301419004000000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 317, DOI: 10.5281/zenodo.7352976

James H. Honacki, Kenneth E. Kinman, James W. Koeppl (1982): Order Artiodactyla. In: James H. Honacki, Kenneth E. Kinman, James W. Koeppl (Eds): Mammal Species of the World (1st Edition). Lawrence, Kansas, USA: Alien Press, Inc. & The Association of Systematics Collections: 315-343, ISBN: 0-89327-235-3, DOI: http://doi.org/10.5281/zenodo.7352976

Lama guanicoe (Muller, 1776). Linne's Vollstand. Natursyst. Suppl., p. 50. TYPE LOCALITY: Chile, Andes. DISTRIBUTION: Cordilleras of the Andes, in S. Peru, Bolivia, Argentina, and Chile; Patagonia, Tierra del Fuego; Navarino Isl. PROTECTED STATUS: CITES - Appendix II., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976

Lama G. Cuvier, 1800. Lecon's Anat. Comp., I, tab. 1. REVIEWED BY: P. Grubb (PG). COMMENT: Should probably include Vicugna (PG). ISIS NUMBER: 5301419004002000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976

Vicugna vicugna (Molina, 1782). Sagg. Stor. Nat. Chile, p. 313. TYPE LOCALITY: Peru, to S. Ecuador and C. Bolivia. (= Chilean Andes, from Coquimo to Copiapo.) DISTRIBUTION: S. Peru; W. Bolivia; N.W. Argentina; N. Chile. COMMENT: Should probably be included in Lama (PG). PROTECTED STATUS: CITES - Appendix I and U.S. ESA - Endangered. ISIS NUMBER: 5301419004002002001 as Lama vicugna., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976

Vicugna Lesson, 1842. Nouv. Tabl. Regn. Anim. Mammal., p. 167., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976

Lama glama (Linnaeus, 1758). Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: Peru, Andes (domesticated stock). DISTRIBUTION: Domesticated in S. Peru; W. Bolivia; N. W. Argentina. COMMENT: Should probably include guanicoe (PG). ISIS NUMBER: 5301419004002001001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976

Lama pacos (Linnaeus, 1758). Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: Peru (domesticated stock). DISTRIBUTION: Domesticated in S. Peru; W. Bolivia. COMMENT: Often regarded as a synonym for glama; see Corbet and Hill, 1980:118., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 318, DOI: 10.5281/zenodo.7352976, {"references":["Corbet, G. B., and J. E. Hill. 1980. A world list of mammalian species. British Museum (Natural History), London, 226 pp."]}

Camelus Linnaeus, 1758. Syst. Nat., 10th ed., 1:65. ISIS NUMBER: 5301419004001000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 317, DOI: 10.5281/zenodo.7352976

Camelus dromedarius Linnaeus, 1758. Syst. Nat., 10th ed., 1:65. TYPE LOCALITY: "Africa," deserts of Libya and Arabia, (domesticated stock). DISTRIBUTION: Extinct in the wild and unknown as fossil; domesticated in North Africa, Arabia, Mediterranean, Balkans, and Middle East; introduced into Australia. COMMENT: Produces fertile hybrids with bactrianus; though their distributions merge, breeding is regulated (as all individuals are domesticated in the zone of contact) (IUK). Corbet, 1978:197, citing Gray, 1954, stated that male hybrids are sterile. Bohldeen, 1961, Z. Tierzucht. Zuchtungsbiol., 76:107- 113, considered dromedarius a synonym of bactrianus. ISIS NUMBER: 5301419004001002001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Artiodactyla, pp. 315-343 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on pages 317-318, DOI: 10.5281/zenodo.7352976, {"references":["Gray, A. P. 1954. Mammalian hybrids: A checklist with bibliography. Commonwealth Agricultural Bureau, Farnham Royal, United Kingdom, 262 pp."]}

Camelus pacos [spec. nov.] C. tophis nullis, corpore lanato. Syst. nat. 13. n. 4. Camelus peruvianus laniger Pacos dictus. Raj. quadr. 147. Habitat in America meridionali. Totus sanguineus, subtus albus; Pedes didactyli. Priori minor, magis lanatus, nec oneribus portandis par., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 66, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Camelus bactrianus [spec. nov.] C. tophis dorsi duobus. Syst. nat. 13. n. 2. Dromedarius. Jonst. quadr. 1. 44. f. 1. Camelus. Gesn. quadr. 151. Aldr. bis. 907. Habitat in Africa. Cursu antecedente celerior, pro magnatibus, rarior., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 65, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Camelus [gen. nov.] Cornua nulla. Dentes Primores inferiores VI, spathiformes. Laniarii distantes: superiores III, inferiores II. Labium superius fissum., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 65, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206

Camelus dromedarius [spec. nov.] C. topho dorsi unico. Syst. nat. 13. n. 1. Camelus unico in dorso gibbo. Raj. quadr. 143. Camelus. Jonst. quadr. t. 41. Gesn. quadr. 159. Aldr. bis. 908. Habitat in Africae desertis arenosis siticulosis. Callos in pedibus anterioribus 4, in posterioribus 2, in sterno 1. gerit. Ventriculus secundus cellulosus pro aqua pura diutius asservanda per siticulosa deserta. Onera portat, lente festinat, lassus jacet super pectus., Published as part of Linnaeus, Carolus, 1758, Systema Naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Stockholm :Laurentius Salvius on page 65, DOI: 10.5962/bhl.title.542, http://zenodo.org/record/3922206