Love on the Mountain Tops

Caprines - members of the goat subfamily - are amongst the mammals most adapted to harsh environments, with the majority of species adapted to living in the cold, barren, and precipitous slopes of mountains. There are some exceptions; sheep (which are taxonomically a subtype of "goat") originally evolved to live in barren rocky hills rather than on true mountains, while some of the East Asian species inhabit forested slopes. 

There are, as with many animal groups, more species of caprine than one might at first think, and I covered them all individually about ten years ago. Looking through that series, it should be possible to appreciate that the group is also varied, not only inhabiting a range of environments but also living varied lifestyles, from those that are near-solitary to those that prefer large herds. This is also reflected in their mating habits which, are as one might expect, related to the size of the community in which they live. One would also expect that the habitat would have some effect on how the animals choose to live, and, in turn, on that mating behaviour.

No Boys Allowed

Even in mammals that are otherwise social, it is quite common for males and females to live apart for most of the year. The most common pattern involves females living in groups with their children, until the latter approach adulthood, while males either live alone, or in much smaller bands, outside of the mating season. In many hoofed herd animals, however, the males live in herds that aren't much smaller than those of the females - in other words, both sexes live in herds, but the two don't mix until it's time to do the necessary.

Clearly, these are creatures that benefit from the group protection that living in herds provides, and, while there's obviously a limit on how large a herd can be before there isn't enough food to support them all, it's not always so clear why the herds should be single-sex. Why, in short, do so many hoofed herd animals practice sexual segregation?

There are a number of theories, and, as is often the case, they aren't mutually exclusive. Nor is it likely that the reason - or the balance of reasons, if there's more than one - will be the same for every species. It's something we have to examine on a case-by-case basis.

Caprines of the Distant Past

Reconstruction of Myotragus balearicus
(Note the odd position of the eyes)

Having said something about all the living species of goat-like animal, it's time to wrap up with a review of where they all came from. Although fossil goats have not received the attention of sabre-tooth cats, let alone dinosaurs, we do have a number of remains, especially from the most recent epoch before today, the Pleistocene. Goats, after all, are reasonably sized animals, and both sexes have prominent horns, which can help to make them distinctive.

Indeed, goats and their kin seem to have done rather well during the Ice Ages of the Pleistocene. Because they are well-adapted to harsh and cold environments, vast ice sheets covering the north were less of a problem for them than for many other animals. They could not, of course, live on the ice sheets themselves, but then very little could. But with most of the rest of Europe, and large swathes of Asia and North America, covered in chilly tundra, that was as good a place for goats as mountainous plateaus are today.

It perhaps particularly suited muskoxen, which, unlike true goats, prefer lowlands rather than precipitous mountains. Even today they are found on the chilly tundra plains of Siberia and Canada and the coasts of Alaska and Greenland. During the Ice Ages, such terrain stretched further south, and (the world being a globe, and continental masses being where they are) that meant there was more of it. That muskoxen are exceptionally large by the standards of "goats" means that they also left some pretty big and impressive skeletons.

Almost a Caprine: The Chiru

Chiru

If there's no clear and universal definition of what a species is, there's no definition at all of what the larger groupings - families, tribes, genera, and whatnot - are. These days, there is a rule that all such groupings should also be "clades" - that is, all the animals in them are more closely related to one another than to anything outside the group. But that's really all there is; there's no way of saying whether a particular clade should be called a "family", an "order", or not given any named ranking at all.

For instance, this rule tells us that humans must belong to the great ape family, since chimps are closer to us than they are to gorillas, but there's nothing in it to say that gibbons can't also belong. We could, in other words, have an "ape family", and the only reason we don't is that we figure gibbons are sufficiently different from great apes that we ought to give them a family of their own. (In fact, "apes" as a whole are considered a superfamily).

Caprines: The World's Largest Goats

Takin (Sichuan subspecies)

All of the various kinds of caprine that I have described so far in this series have looked, more or less, either goat-like or sheep-like. There are just two species left, and those form something of an exception. While most caprines are fairly medium-sized as hoofed animals go, these are much larger, more muscular, animals.

It used to be thought, on this basis, that they were closely related to one another, representing an early branch in the evolution of goat-like animals that split away from their relatives well before the appearance of actual goats or sheep. From modern genetic evidence, that no longer looks the case. They are, as we suspected, caprines, (although, despite the title of this post, not literally goats) but within that group, they are not particular closely related to one another. Instead, their apparent similarities are a coincidence, a case of parallel evolution where two animals, both fairly goat-like to start with, faced selective pressures to become larger.

The first of these animals is the takin (Budorcas taxicolor). Takins are very distinctive animals, quite hard to mistake for anything else, once you get a good look at them. The most obvious point, as I've already implied, is the size. A fully grown male takin stands over four feet high at the shoulder, and weighs upwards of 300 kg (660 lbs). The females are noticeably smaller, but still larger than even the males of any other caprine species (with, of course, one exception that I'll get to in a moment).

Caprines: Goats of the Shadowed Cliffs

Himalayan goral

It is, of course, the case that all animals are continuously evolving, even if some of them don't change much visibly over protracted periods of time. So we can't really point to a modern species and say that this is the kind of thing that everything else in its group evolved from. But some have, at least on visible examination, changed far less from their ancestors than others. They have found some suitable niche in which to live, and, because they're environment hasn't changed much, they've been able to stay more or less the same. It's not that they haven't changed, of course, but just that the changes are relatively subtle.

In the case of the goats, the species that have probably changed the least from the ancestral form are the serows and gorals of eastern Asia. Both kinds of animal have retained a "resource defender" lifestyle, where they find patches of lush food in the otherwise rocky wastes of precipitous cliffs and mountainsides, and defend them from all comers.

Caprines: Goats Among the Volcanoes

Japanese serow

While most goat-like animals today are grazers, subsisting on relatively large quantities of low-quality forage, it's thought that their ancestors were originally browsing animals, defending small patches of high quality food in otherwise inhospitable and craggy terrain. In the west, the most familiar surviving "resource defenders" of this kind are the chamois of Europe. While the exact evolutionary relationships of chamois aren't entirely clear, they are part of a broad cluster of species that is otherwise crammed with grazing animals, such as "true" goats and sheep.

Not so their counterparts in Asia, which form their own distinct evolutionary line, one that never appears to have given rise to true grazing animals. It is these species, not the chamois, that probably most resemble the ancestral goats from which all others evolved.

Caprines: Browsing Goats of the Western Mountains

Alpine chamois

The caprines as a whole can be divided into two groups, depending on which of two different lifestyles they happen to follow. The best known of these, since it is the lifestyle of sheep and "true" goats, is that of the grazing caprines.

Grazers feed fairly indiscriminately, munching down lots of grass and similar plants. In the case of the caprines, they survive in marginal habitats by eating pretty much anything that's available. They range across large areas in search of food, and adopt safety in numbers by packing themselves together in herds. Today, the great majority of caprine species adopt this lifestyle, and it's often what we think of when we think of goats and sheep.

But it appears that, in the evolutionary history of goat-like animals, it's a relatively recent innovation, one that was given a significant boost by the arrival of the Ice Ages. Before that, goats had adopted a rather different lifestyle, and there are still a minority of species - perhaps no more than five - that still live this way.

These apparently "primitive" goats were once grouped together in their own tribe, and given impressive sounding technical names like "rupicaprines" or "naemorhadins", but it's now less clear that they're really related. Instead, they represent at least two evolutionary lines within the goat-like animals, each a relic of the distant past when all goats were like this. Around them, some of their relatives switched to grazing, and, in the long run, proved the more successful.

Caprines: Half-Goats and False Sheep

Himalayan tahrs

Sheep and goats are fairly closely related animals. In a sense, sheep are just goats that don't like climbing. However, even once we acknowledge that ibex, say, are really just another kind of goat, it turns out that there are a number of species that are also closely related to sheep and goats, without really being either.

Quite how they're related, and which ones are on the goat side of the tree, and which on the sheep side, has been a matter of some debate, and depends largely on whether or not you think that physical appearance is more important than genetic similarity. Even if you look only at the genetics, it seems that they're all close enough to one another that it makes a fair difference which genes you happen to consider most important.

Still, we can say that some look more like goats, and some look more like sheep. On the goat-like side are the tahrs - not to be confused with turs, which really are goats. There are three species of tahr, all of which look fairly similar to one another, and have traditionally been placed together in the genus Hemitragus - a word that literally means "half-goat". In the traditional, appearance-based, scheme it's thought that these are the closest animals to true goats, without actually being them. Genetic analysis, however, shows that this is just an illusion, and the three animals aren't especially close relatives. Two of them, therefore, have been removed from the genus, and given new scientific names to reflect their distinct nature.

Caprines: Lands of the Ibex

Alpine ibex

The Ice Ages were, on the whole, not a bad time for goats. True, they had to leave the vertiginous cliffs of their mountain homes when they became too cold and barren to support life - and, in many cases, were swathed in vast glaciers. But, as the world grew colder, the vegetation also moved down the mountain slopes, so that, down in the lowlands, goats found plenty of food they had been used to. Indeed, they were better suited to it than most other animals, which had to move to southern climes, rather than merely heading downhill.

Since there are rather more lowland areas than there are mountains, goats could spread much further than they could during warmer times. When the Ice Ages ended, and the hot weather returned, they simply headed back up the mountains. But not, necessarily, the same mountains that they had previously come down from.

As a result, we now have quite a range of goat species across Asia, and, to some extent, Africa. After all, between (and after) the Ice Ages, each population was isolated from those in other ranges, and could develop on its own. Taking, at least for today, our definition of 'goat' to be "any species from the genus Capra", there are probably at least seven, and maybe eight or nine, wild species of goat. The wild goat itself is one, and the markhor, with its bizarre corkscrew horns, is another. Most of the others are collectively known as "ibexes".

Caprines: At Last, The Goats

Wild goat

In English-speaking countries, and, for that matter, South America, sheep are a far more common sight on farms than goats are. (To be fair, in much of the US, even sheep aren't as common as they are in Britain, let alone Australia and New Zealand). However, while the worldwide population of sheep is, indeed, higher than that of domestic goats, it's not by as big a margin as you might think. In particular, goats are a vital mainstay of the agricultural economy in India, and throughout much of Africa.

The reason for this is their extreme hardiness. Adapted to living in habitats even more marginal than those of sheep - which, at least in their wild form, are themselves more resilient than, say, cows or pigs - they're ideal for raising in the sort of scrubby environments that you get in much of Africa and the Middle East. Even so, after approximately twelve thousand of years of selective breeding, the domestic goat has come a long way from its wild ancestors. They're often larger, with smaller horns, or none at all, and some have long, floppy ears. There are a number of different breeds, from milk-producers like Saanens and Toggenburgs, to the muscular Boer goats, bred for their meat, to the long-haired angora and cashmere goats that produce fine wool and mohair.

Caprines: Go West, Young Sheep!

Bighorn rams

As we've seen, the domestic sheep is descended from one of what may be quite a large number of related species native, broadly speaking, to southern Asia. Back in the Pliocene, however, long before those species separated from one another, one stock of ancestral sheep headed north, into Siberia. When the Ice Ages arrived, and sea levels dropped, these sheep were among several animals that headed... well, from their perspective, they were going east, but where they ended up was western North America.

Sheep, in general, inhabit high hills and rugged terrain in the shadow of mountains. They don't live on the cliffs and the impassable heights themselves, but they do like precipitous rocks nearby, so that they can flee into them at the first sign of predators. Given those requirements, western North America is an absolutely ideal place for sheep to be, and the descendants of those first colonists spread far and wide. However, while there's much argument about the exact number of wild Asian species of sheep, it's pretty much agreed that, in America, there are only two.

Probably the more familiar of these is the bighorn sheep (Ovis canadensis). It's also easily the more widespread of the two, and inhabits a surprisingly broad range of habitats, from the mountains of southern British Columbia and the hills of North Dakota down to Baja California and the Sonora Desert. That obviously includes both hot, scrubby, deserts and cold, damp, pine forests, as well as much in between. Indeed, aside from the insistence on rugged terrain that prevents them from reaching as far east as, say, Kansas or Nebraska, bighorn sheep don't seem to have much in the way of requirements.

Caprines: Telling the Sheep from the Goats

Transcaspian urials
(Note that the rams of this species/subspecies have
particularly argali-like horns)

To most people, especially in the West, the most familiar member of the goat subfamily is probably... the sheep. Worldwide, domestic sheep are more common than goats, although not necessarily by as much as you might think. Goats are more popular than sheep as farm animals in places like India and Africa, but overall, sheep have the edge in numbers. That's probably no great surprise to those of you in most parts of Europe or America, and even less so if you're in Australia or New Zealand.

But perhaps I should back up there. Did I really just say that sheep are a kind of goat?

Well, yes I did. Kind of. Sheep are members of the goat subfamily, but really, they're goats only in the sense that ferrets are a kind of weasel. But why is it that way round at all?

The goat subfamily, or Caprinae, was given its name by the great Victorian zoologist John Edward Gray, quite early in his career, in 1821. From a scientific standpoint, there was absolutely no reason why he couldn't have named it the sheep subfamily, but instead, he chose goats as the best example of the group. I don't know his actual reasoning, but it certainly makes sense: far more members of the group resemble goats than resemble sheep. And so, from this, admittedly arbitrary, standpoint sheep are an odd kind of goat, and not the other way round.

Caprines: Goats, Horns, and Antlers

Over the last year, I completed a survey of the weasel family, the largest, and most diverse, family of carnivoran mammals. When it comes to the big herbivores, the largest and most diverse family is the Bovidae - the cattle family. This includes a wide range of animals, from the true bovines (that is, bison, yak, and so forth) to slender gazelles. In fact, there are well over a hundred species in the cattle family, and covering them in the same way that I covered the weasels would take me a couple of years or so.

So, instead, I am going to look at just one subfamily within the group. After the gazelles, it's the second largest cattle subfamily, with the bovines down in third place. These, as you'll have guessed from the title of the post, are the goats.

Or, more accurately, "caprines". Just as many members of the weasel family aren't literally weasels, many members of the goat subfamily aren't goats in the usual sense of the word. If you've never really looked at mammalian taxonomy, there will probably be at least a couple of species I'm going to cover over the coming months that will surprise you, and likely a few you've never heard of. One collective term for all of these animals is "goat-antelopes", but it's hardly common usage, and I don't actually find it very useful - not least because most of them don't look much like the regular image of an antelope, either. So I'll stick with the technical term of "caprines", or sometimes just call them goats, and you'll have to accept that I mean that in an unusually broad sense.