WHATEVER HAPPENED TO THE HUEQUE? SEEKING THE LOST LLAMA OF CHILE

Vidaurre's engraving from 1776 depicting four of the five South American camelids once recognised – llama (top left), hueque aka Chilihueque (top right), vicuna (bottom left), guanaco (bottom right) (public domain)

Llamas are very familiar animals to science and the general public alike. As will now be revealed in this ShukerNature blog article, however, they may well have once shared their native Andean homelands with a highly unfamiliar, all-but-forgotten close relative whose zoological identity and disappearance have remained unexplained for well over 300 years.

ASIAN CAMELS AND SOUTH AMERICAN LLAMAS – THE LAST OF THEIR DYNASTY

Back in prehistoric times, the camelids were a very diverse taxonomic family of artiodactyl (even-toed) ungulates, distributed widely across the globe and represented by small, large, humped, humpless, short, tall, and sometimes very tall forms (such as North America's prairie-inhabiting giraffe camel Aepycamelus, aka Alticamelus).

Early restoration of Aepycamelus the giraffe camel by Heinrich Harder in 1920 (public domain)

Today, however, this once-mighty and extremely diverse dynasty is reduced to just six representatives – the two species of humped camel (one-humped dromedary Camelus dromedarius and two-humped Bactrian C. bactrianus) native to Asia (plus feral populations variously established in parts of Europe and Australia); and the four humpless species native to South America. These latter four species are the vicuna, alpaca, llama, and guanaco.

Following the discovery and conquest of South America by the Spanish during the 1500s, its humpless camelids attracted great interest from Western naturalists, with the llama in particular featuring in a number of bestiaries (such as Edward Topsell's famous work The History of Four-Footed Beasts and Serpents, 1658), in which it was generally referred to as the allocamelus.

The llama or allocamelus as depicted in Edward Topsell's bestiary The History of Four-Footed Beasts and Serpents (1658) (public domain)

It even entered European heraldry, where it was sometimes dubbed the ass-camel, and was duly represented with the head of an ass and the body of a short-legged, convex-backed camel.

The New World's camelid quartet also incited much confusion as to how they were related to one another. For whereas the vicuna and guanaco are wild species, the llama and alpaca are entirely domesticated.

Llama (© Dr Karl Shuker)

And to make matters even worse, the term 'llama' eventually established itself colloquially as a term not just specific to its own particular single species but also as a general term covering all four South American species.

Vicunas (© Haplochromis/Wikipedia CC BY-SA 3.0 licence)

Eventually, the consensus was that the small goat-like vicuna was not just a valid species but one so distinct from the others that it deserved its own genus, and was duly dubbed Vicugna vicugna. The remaining trio were housed together in a second genus, Lama, and were formally christened Lama guanicoe (=huanacos) (the guanaco), L. glama (the llama), and L. pacos (the alpaca), with both the llama and the alpaca being deemed to be domesticated descendants of the guanaco.

Guanaco (public domain)

DNA studies published in 2001, however, revealed that the alpaca is in fact most closely related to the vicuna, and is believed to have descended from this latter species, not from the guanaco after all. Consequently, the alpaca is now housed with the vicuna in the genus Vicugna, as Vicugna pacos.

Alpacas (public domain)

In addition, there are two non-taxonomic breeds or varieties of alpaca – the rare Suri alpaca, sporting a long, shiny, very soft, slightly-curled fleece, which is very expensive; and the more common Huacaya alpaca, sporting a shorter, fluffier fleece, which is far less expensive.

CAMELID HYBRIDS – THE HUARIZO AND THE CAMA

Yet even though they are now split into separate genera, the alpaca and the llama are sufficiently closely related genetically to yield viable crossbred offspring– a hybrid resulting from interbreeding between a male llama and a female alpaca is known as a huarizo. Much smaller than llamas, it is greatly valued for its very lengthy fleece and gentle disposition, but is usually sterile. Remarkably, moreover, there have even been cases of successful intergeneric hybridisation in captivity between male dromedaries and female llamas, the resulting camel x llama crossbreed being referred to as a cama.

Adult cama (© unknown to me – all information would be welcomed; photographs included on a strictly non-commercial Fair Use basis only)

This surprising feat was first achieved in 1998, via artificial insemination, at the Camel Reproduction Centre in Dubai, the aim being to create an animal with the size, patience, and stamina of a camel but with a fleece at least as good as (if not better than) a llama's. The first cama, a male, born in 1998, was named Rama; a second cama, a female, was born in 2002, and was named Kamilah. In each case, it looked like a large version of its llama mother in overall appearance, and lacked its camel father's hump, but it did possess his small ears and short tail.

A HISTORY OF THE HUEQUE, CHILE'S MISSING MINI-LLAMA

Hybrids notwithstanding, what is not widely known nowadays is that in addition to the vicuna, alpaca, guanaco, and llama, not so very long ago there may also have been a fifth New World camelid species, or at least a well-defined variety of one of the still-existing quartet. Formerly found in Chile, this seemingly-lost and certainly long-forgotten llama was known as the hueque, or the chilihueque in full.

Between the 16^th and 17^th Centuries, Spanish-speaking travellers who visited the central and south-central valleys of Chile reported the presence in territories owned by the Araucanians (i.e. the Mapuches peoples known here as the Moluche) of a small, distinctive type of llama not seen anywhere else. Moreover, the travellers learnt that its existence here pre-dated the Hispanic conquest, and it may well have been adopted by the Moluche from the Inca culture. This intriguing creature was the hueque, which was generally bred not as a beast of burden (like the llama is in other South American countries) but for its meat and in particular for its fleece, which was extremely soft, luxuriant, and so long that it dragged on the ground as the animal walked.

Having said that, Chilean Jesuit priest and naturalist Father Juan Ignacio Molina noted in his 2-volume magnum opus The Geographical, Natural and Civil History of Chili (1782) that when the Dutch sea captain Admiral Joris van Spilbergen had landed on Chile's small Mocha Island in 1614, he had observed hueques being used to pull small carts by Mapuches living there. Confusingly, however, further on in his book Molina contradicted himself by stating that what van Spilbergen had seen hueques being used for on Mocha Island was pulling ploughs.

Father Juan Ignacio Molina (public domain)

Writing in 1550 after having conquered southern Chile, the Spanish conquistador Pedro de Valdivia stated that the hueque was very abundant in this region, and that not only were the inhabitants dressed exuberantly in the most elegant woollen clothes but even their houses were stocked full of wool. Also present here was a second camelid, known as the luan, but it was the fleece of the hueque that was always used to manufacture the most prized, sought-after woollen garments. Indeed, the Spaniards were so impressed by this animal's superior fleece that they dubbed it 'the sheep of the land',

The luan was generally identified as the guanaco, but what exactly was the hueque? There was no doubt that it too was some type of llama (using this term in its general sense here), but its precise nature incited much controversy among early naturalists. Two conflicting schools of thought eventually arose. One asserted that it was a local (semi-)domesticated variety of the guanaco distinct from the llama, the other claimed that it was one and the same as the llama and that it had been introduced here from further north, but neither option garnered a significant majority of support. This remains true today.

Guanaco photographed in Chile's Torres del Paine National Park (public domain)

The few illustrations of the hueque that were produced while it still existed generally depicted it as being similar to the llama but somewhat smaller in overall size, with a slightly shorter neck and legs, but sporting a thicker fleece. However, these were based merely upon verbal accounts received from others, rather than upon first-hand observations made by the engravers themselves, so they may not be wholly accurate representations of this lost form. Written accounts of the hueque by Molina and others claimed that it occurred in several different colours – white, brown, black, and grey. Molina also stated that it was approximately 6 ft long, and stood about 4 ft tall.

Perhaps the most natural-looking representation of an alleged hueque, depicted alongside a llama, is the one reproduced below:

An engraving from a book by Amédée-François Frézier, published in 1716, depicting a llama (on the left) and an alleged hueque (on the right) (public domain)

It appeared as Plate 22 in A Voyage to the South-Sea and Along the Coasts of Chili and Peru in the Years 1712, 1713 and 1714, which was written by French explorer Amédée-François Frézier and published in 1716. Unfortunately, however, whereas the alleged hueque was portrayed in side view, the llama was merely depicted standing face-on, thereby preventing direct morphological comparisons of these two camelid types to be readily made.

WHY DID THE HUEQUE DIE OUT, AND WHAT EXACTLY WAS IT?

The reason why the hueque had vanished by the end of the 17^th Century, possibly even earlier, remains unclear. However, its extinction coincided with a major influx of domestic cattle into this region of Chile, brought here from elsewhere for their meat, milk, leather, and as sturdy beasts of burden, as well as European sheep introduced for their wool and meat. Consequently, it has been surmised that not only did they render the hueque superfluous, but these non-native livestock beasts may also have carried with them diseases hitherto unknown here and to which the hueque had no resistance, thus wiping it out.

As for the hueque's identity, that is still unresolved too – or is it? Although, as noted earlier, attempts have been made by various researchers to link the hueque to either the guanaco or the llama, I personally favour a third candidate – the alpaca. So too did English writer and alpaca authority William Walton when describing the alpaca of Peru in his book An Historical and Descriptive Account of the Peruvian Sheep, Called Carneros de la Tierra (1811), though his contribution to the debate concerning the hueque's identity had long been forgotten until I encountered his book recently.

A guanaco with a Peruvian warrior, from Walton's above-cited book (public domain)

Of the four still-extant South American camelids, it is unquestionably the alpaca that offers the closest correspondence to the hueque. After all, both the hueque and the alpaca were/are bred predominantly for their fleece; both of them yielded/yield wool so profuse and luxuriant that it could/can reach the ground (especially in Suri alpacas); and both of them were/are smaller and more compact than the larger, longer-necked, longer-limbed llama and guanaco.

Could it be, therefore, that a variety of alpaca was either raised within or introduced into central and southern Chile from northern Chile or Peru (where the alpaca occurs naturally), and it was this alpaca form that was in reality the mysterious, now-vanished hueque? If nothing else, it is interesting to note that in an engraving from Gómez de Vidaurre's Compendio della Storia Geografica, Naturale e Civile del Regno del Chile (1776), depicting four South American camelids and opening this present ShukerNature blog article, the hueque is included, but the alpaca is absent. Is this strange omission of such a well known relative in favour of the much more obscure hueque an indication that these two forms were actually one and the same creature?

Llama pattern on a Chilean alpaca-wool jumper that was owned by my mother Mary Shuker (© Dr Karl Shuker)

After all, if the hueque were actually a separate, distinct species in its own right and was once abundant in southern Chile, plentiful remains of this creature would surely have existed and would have been readily delineated by scientific scrutiny from those of the four known South American camelids. Yet no formal scrutiny and osteological differentiation seems to have been documented, thereby indicating that the hueque was indeed conspecific with one of the pre-existing quartet of species.

Consequently, I conclude that the hueque was most likely to have been a breed or variety of alpaca. Sadly, however, we may well be more than 300 years too late to ever know for sure.

And finally, on a much lighter note, straight from a famous if fictitious animal linguist's circus of exotic creatures, here is the rarest llama-inspired cryptid of all:

Courtesy of Doctor John Dolittle, my very own pushmi-pullyu (© Dr Karl Shuker)