LLAMA, sometimes spelt Lama, a word by which the Peruvians designated one of a small group of closely allied animals, which, before the Spanish conquest of America, were the only domesticated hoofed mammals of the country, being kept, not only for their value as beasts of burden, but also for their flesh, hides, and wool,in fact, supplying in the domestic economy of the people the place of the horse, the ox, the goat, and the sheep of the Old World. The word is now sometimes restricted to one particular species or variety of the group, and sometimes used in a generic sense to cover the whole. Although they were often compared by early writers to sheep, and spoken of as such, their affinity to the camel was very soon perceived, and they were included, in the genus Camelus in the Systema Naturae of Linnaeus. They were, however, separated by Cuvier in 1800 under the name of Lama, changed by Illiger in 1811 to Auchenia (in allusion to the great length of neck, _______), a term afterwards adopted by Cuvier, and almost universally accepted by systematic zoologists, although there has been of late a disposition to revive the earlier name.
The animals of the genus Auchenia or Lama are, with the two species of true camels (to which the generic term Camelus is now restricted), the sole existing representatives of a very distinct section of the "artiodactyle" or even toed ungulates, called Tylopoda, or "boss-footed," from the peculiar bosses or cushions placed on the under surface of their feet, and on which they tread. This section thus consists of a single family, the Camelidae, the other section of the same great division being the Suina or pigs, the Tragulina or chevrotains, and the Pecora, or true ruminants, to each of which the Tylopoda have more or less affinity, standing in some respects in a central position between them, borrowing as it were some characters from each, but in others showing great special modifications not found in any of the other sections.
Until within the last few years the existence of two genera having so very much in common as the camels and the llamas, and yet so completely isolated geographically, had not received any satisfactory explanation, for the old idea that they in some way "represented" each other in the two hemispheres of the world was a mere fancy without philosophical basis. The discoveries made mostly within the past ten years of a vast and previously unsuspected extinct fauna of the American continent of the Tertiary period, as interpreted by the able palaeontologists Leidy, Cope, and Marsh, has thrown a flood of light upon the, early history of this family, and upon its relations to other mammals. It is now known that llamas at one time were not confined to the part of the continent south of the Isthmus of Panama, as at the present day, for their remains have been abundantly found in the Pleistocene deposits of the region of the Rocky Mountains, and in Central America, some attaining a much larger size than those now existing. There have also been found in the same regions many camel-like animals exhibiting different generic modifications, and, what is more interesting, a gradual series of changes, coinciding with the antiquity of the deposits in which they are found, have been traced from the thoroughly differentiated species of the modern epoch down through the Pliocene to the early Miocene beds, where, their characters having become by degrees more generalized, they have lost all that especially distinguishes them as Camelidae, and are merged into forms common to the anecstral type of all the other sections of the Artiodactyles. Hitherto none of these annectant forms have been found in any of the fossiliferous strata of the Old World; it may therefore be fairly surmised (according to the evidence at present before us) that America was the original home of the Tylopoda, and that the true camels have passed over into the Old World, probably by way of the north of Asia, where we have every reason to believe there was formerly a free communication between the continents, and then, gradually driven southward, perhaps by changes of climate, having become isolated, have undergone some further special modifications; while those members of the family that remained in their original birthplace have become, through causes not clearly understood, restricted solely to the southern or most distant part of the continent. There are few groups of mammals of which the palpeontological history has been so satisfactorily demonstrated as the one of which we are treating.1
The special characters which the llamas and camels have in common, and the combination of which distinguishes them from the rest of the Artiodactyles, are as follows. The premaxillae have the full number of incisor teeth in the young state, and the outermost is persistent through life, an isolated laniariform tooth. The canines are present in both jaws, and those of the mandible are differentiated from the long, procumbent, and spatulate incisors, being suberect and pointed. The crowns of the true molars belong to the crescentic or "selenodont" type, and are very long or "hypsodont"; but one or more of the anterior premolars is usually detached from the series, and of simple pointed form. The hinder part of the body is much contracted, and the femur long and vertically placed, so that the knee-joint is lower in position, and the thigh altogether more detached from the abdomen than in most quadrupedal mammals. The limbs are long, but with only two digits (the third and fourth) developed on each, no traces of any of the others being present. The trapezoid and marum of the carpus, and the cuboid and navicular of the tarsus are distinct. The two metapodal bones of each limb are confluent for the greater part of their length, though separated for a considerable distance at the lower end. Their distal articular surfaces, instead of being pulley-like, with deep ridges and grooves, as in other Artiodactyles, are simple, rounded, and smooth, The proximal phalanges are expanded at their distal ends, and the wide, depressed middle phalanges are embedded in a broad cutaneous pad, forming the sole of the foot, on which the animal rests in walking instead of on the hoofs. The ungual phalanges are very small and nodular, not flattened on their inner or opposed surfaces, and not completely encased in hoofs, but bearing nails on their upper surface only. The cervical region is long and flexuous, and the vertebrae of which it is composed are remarkable for the position of the canal for the transmission of the vertebral artery, which does not perforate the transverse process, but passes obliquely through the anterior part of the pedicle of the arch (a condition only found in two other genera of mammals, Macrauchenia and Myrmecophaga). There are no horns or antlers. Though these animals ruminate, the stomach differs considerably in the details of its construction from that of the Pecora. The interior of the rumen or paunch has no villi on its surface, and there is no distinct psalterium or maniplies. Both first and second compartments are remarkable for the presence of a number of pouches or cells in their walls, with muscular septa, and a sphincter-like arrangement of their orifices, by which they can be shut off from the rest of the cavity, and into which the fluid portion only of the contents of the stomach is allowed to enter.1 The placenta is diffuse as in the Suina and Tragulina, not cotyledonary as in the Pecora. Finally, they differ not only from other ungulates, but from all other mammals, in the fact that the red corpuscles of the blood, instead of being circular in outline, are oval as in the inferior vertebrated classes.
The following characters apply especially to the llamas. Dentition of adults:incisors 1/3, canines 1/4, premolars 2/2, molars 3/3 ; total 32. In the upper jaw there is a compressed, sharp, pointed laniariform incisor near the hinder edge of the premaxilla, followed in the inale at least by a moderate-sized, pointed, curved true canine in the anterior part of the maxilla. The isolated canine-like premolar which follows in the camels is not present. The teeth of the molar series which are in contact with each other consist of two very small premolars (the first almost rudimentary) and three broad molars, constructed generally like those of Camelus. In the lower jaw, the three incisors are long, spatulate, and procumbent; the onter ones are the smallest. Next to these is a curved, suberect canine, followed after an interval by an isolated minute and often deciduous simple conical premolar ; then a contiguous series of one premolar and three molars, which differ from those of Camelus in having a small accessory column at the anterior outer edge. The skull generally resembles that of Camelus, the relatively larger brain-cavity and orbits and less developed cranial ridges being due to its smaller size. The nasal bones are shorter and broader, and are joined by the premaxillae. Vertebrae:cervical 7, dorsal 12, lumbar 7, sacral 4, caudal 15 to 20. Ears rather long and pointed. No dorsal hump. Feet narrow, the toes being more separated than in the camels, each having a distinct plantar pad. Tail short. Hairy covering long and woolly. Size smaller and general form lighter than in the camels. At present and within historic times they are entirely confined to the western side and southermost parts of South America, though fossil remains have been found in the caves of Brazil, in the pampas of the Argentine republic, and, as before mentioned, in Central and North America.
In essential structural characters, as well as in general appearance and habits, all the animals of this genus very closely resemble each other, so that the question as to whether they should be considered as belonging to one, two, or more species has been one which has led to a large amount of controversy among naturalists. The question has been much complicated by the circumstance of the great majority of individuals which have come under observation being either in a completely or partially domesticated state, and descended from ancestors which from time immemorial have been in like condition, one which always tends to produce a certain amount of variation from the original type. It has, however, lost much of its importance since the doctrine of the distinct origin of species has been generally abandoned. The four forms commonly distinguished by the inhabitants of South America are recognized by some naturalists as distinct species, and have had specific designations attached to them, though usually with expressions of doubt, and with great difficulties in defining their distinctive characteristics. These are(1) the llama, Auchenia glama (Linn.), or Lama peruana (Tiedemann) ; (2) the alpaca, A. pacos (Linn.) ; (3) the guanaco or huanaco, A. huanacus (Molina) ; and (4) the vicugna, A. vicugna (Molina), or A. vicunna, (Cuv.). The first and second are only known in the domestic state, and are variable in size and colour, being often white, black, or piebald. The third and fourth are wild, and lightbrown colour, passing into white below. They certainly differ from each other, the vicugna being smaller, more slender in its proportions, and having a shorter head than the guanaco. It may, therefore, according to the usual view of species, be considered distinct. It lives in herds on the bleak and elevated parts of the mountain range bordering the region of perpetual snow, amidst rocks and precipices, occurring in various suitable localities throughout Peru, in the southern part of Ecuador, and as far south as the middle of Bolivia. Its manners very much resemble those of the chamois of the European Alps; and it is as vigilant, wild, and timid. The wool is extremely delicate and soft, and highly valued for the purposes of weaving, but the quantity which each animal produces is not great.
The guanaco has an extensive geographical range, from the high lands of the Andean region of Ecuador and Peru to the open plains of Patagonia, and even
The wooded islands of Tierra del Fuego. It constitutes the principal food of the Patagonian Indians, and its skin is invaluable to them, as furnishing the material out of which their long robes are constructed. It is about the size of a European red deer, and is an elegant animal, being possessed of a long, slender, gracefully curved neck and fine legs. Dr Cunningham,2 speaking from observation on wild animals, says:
"It is not easy to describe its general appearance, which combines me of the characters of a camel, a deer, and a goat. The body, deep at the breast but very small at the loins, is covered with long, soft, very fine hair, which on the upper parts is of a kind of fawncolour, and beneath varies from a very pale yellow to the most beautiful snow-white. The head is provided with large ears, in general carried well back, and is covered with short greyish hair, which is darkest on the forehead. Occasionally the face is nearly black. As a rule, it lives in flocks of from half a dozen to several hundreds, but solitary individuals are now and then to be met with. They are very difficult to approach sufficiently near to admit of an easy shot, as they are extremely wary, but, on being disturbed, canter off at a pace which soon puts a safe distance between them and the sportsman, even though he should be mounted. Despite their timidity, however, they are possessed of great curiosity, and will sometimes advance within a comparatively short distance of an unknown object, at which they will gaze fixedly till they take alarm, when they effect a speedy retreat. Their cry is very peculiar, being something between the belling of a deer and the neigh of a horse. It would be difficult to overestimate their numbers upon the Patagonian plains; for in whatever direction we walked we always came upon numbers of portions of their skeletons and detached bones."
Darwin, who has given a most interesting account of the habits of the guanaco in his Naturalists Voyage, says that they readily take to the water, and were seen several times at Port Valdes swimming from island to island.
The llama is only known as a domestic animal, and is chiefly met with in the southern part of Peru. Burmeister, the latest and a very competent writer on the subject,1 says that he is perfectly satisfied that it is the descandant of the wild guanaco, an opinion opposed to that of Tschudi. It generally attains a larger size than the guanaco, and is usually white or spotted with brown or black, and sometimes altogether black. The earliest and often quoted account of this animal by Augustin de Zarate, treasurer-general of Peru in 1544, will bear repeating as an excellent summary of the general character and uses to which it was put by the Peruvians at the time of the Spanish conquest. He speaks of the llama as a sheep, observing, however, that it is camel-like in shape, though destitute of a hump:-
"In places where there is no snow, the natives want water, and to supply this they fill the skins of sheep with water and make other living sheep carry them, for, it must be remarked, these sheep of Peru are large enough to serve as beasts of burden. They can carry about one hundred pounds or more, and the Spaniards used to ride them, and they would go four or five leagues a day. When they are weary they lie down upon the ground, and as there are no means of making them get up, either by beating or assisting them, the load must of necessity be taken off. When there is a man on one of them, if the beast is tired and urged to go on, he turns his head round, and discharges his saliva, which has an unpleasant odour, into the riders face. These animals are of great use and profit to their masters, for their wool is very good and fine, particularly that of the species called pacas, which have very long fleeces ; and the expense of their food is trifling, as a handful of maize suffices them, and they can go four or five days without water. Their flesh is as good as that of the fat sheep of Castile. There are now public shambles for the sale of their flesh in all parts of Peru, which was not the case when the Spaniards came first; for when one Indian had killed a sheep his neigbbours came and took what they wanted, and then another Indian killed a sheep in his turn."
The disagreeable habit here noticed of spitting in the face of persons whose presence is obnoxious is common to all the group, as may be daily witnessed in specimens in confinement in the menageries of Europe. One of the principal labours, to which the llamas were subjected at the time of the Spanish conquest was that of bringing down ore from the mines in the mountains. Gregory de Bolivar estimated that in his day as many as three hundred thousand were employed in the transport of the produce of the mines of Potosi alone, but since the introduction of horses, mules, and donkeys the importance of the llama as a beast of burden has greatly diminished.
The alpaca is believed by most naturalists to be a variety of the vicugna; others have, however, identified it with the guanaco, and some consider it as a distinct species. It is usually found in a domesticated or semi-domesticated state, being kept in large flocks which graze on the level heights of the Andes of southern Peru and northern Bolivia at an elevation of from 14,000 to 16,000 feet above the sea-level, throughout the year. It is not used as a beast of burden like the llama, but is valued only for its wool, of which the Indian blankets and ponchas are made. Its colour is usually dark brown or black. The characteristics of its wool, and the history of its introduction into British manufacturing industry, are described in the article ALPACA. (W. H. F.)
FOOTNOTE (page 738)
1 See especially E. D. Cope, in Wheelers Report of the Survey West of the 100th Meridian, iv. pt. 2, pp. 325-46, 1877.
FOOTNOTES (page 739)
(1) The stomach of the camel inhabiting the Arabian desert is commonly looked upon as a striking example of specialized structure, adapted or modified in direct accordance with a highly specialized mode of life ; it is therefore, very remarkable to find an organ exactly similar, except in some unessential details, in the llamas of the Peruvian Andes and the guanacos of the Pampas. No hypothesis except that of a common origin will satisfactorily account for this, and, granting that this view is correct, it becomes extremely interesting to find for how long a time two genera may be isolated and yet retain such close similarities in parts which in other groups appear readily subject to adaptive modifications.
(2) Natural History of the Strait of Magellan, 1871.
FOOTNOTE (page 740)
1 Description Physique de la République Argentine, vol, iii. p. 458, 1879.
The above article was written by Rev. W. Howard Frere.