Bovines: Prehistoric Bison and Buffalo

Steppe bison, from a cave in Altamira, Spain

In the course of this series I have described 22 species of bovine. Whether that number accurately reflects the number of species alive today depends, in part, on what you consider to be a species - not least whether or not long-term domestication is a sufficiently dramatic thing to constitute the creation a new species. But that's the number I came up with, and it's worth noting that no less than seven of these species - almost a third - are currently considered "endangered", and at high risk of extinction. Indeed, the kouprey may well already be extinct, while the species to which domestic cattle belong is extinct in the wild.

(On that last note, incidentally, I received a note after posting on the subject reminding me of the Chillingham Cattle of Northumberland. These are sometimes claimed to be aurochs, or genuinely wild cattle, but this has no foundation, and, while they have been living wild for centuries, they are actually feral animals, likely descended from escaped medieval stock).

At any rate, there have been many other species of bovine that have gone extinct over the millions of years since their first appearance, in most cases for reasons that have nothing to do with humans. So, albeit with a focus on the genuinely cow-like ones, rather than the spiral-horned antelopes and their kin, today I'm going to look at some of them.

Bovines: Blue Bulls and the Smallest Bovine

Male nilgai

The bovine subfamily can be divided into two main evolutionary branches. One consists of the truly cow-like bovines, including such things as bison, buffalo, and yak, alongside the regular domestic animal. The other are the "spiral horned antelopes", African species that, despite being more closely related to cows than (say) gazelles, nonetheless have a generally antelope-like body form. But there are two living species that fit into neither of these branches. They are relatively closely related to one another, and so form a third, much smaller, evolutionary branch (usually given the taxonomic rank of "tribe"). Although generally considered "antelopes", both species are native to Asia, unlike their spiral-horned kin.

The larger, and likely better-known, of the two is the nilgai (Boselaphus tragocamelus). Standing 120 to 140 cm (4' to 4'7") at the shoulder, although they are considered antelopes, there is a somewhat cow-like appearance to them, albeit with a much narrower head. Indeed, the scientific name reflects their somewhat odd form, since it literally translates as "cow-deer goat-camel" (in fairness, the two halves of the name were coined separately, and not used together for over 60 years afterwards). They are found throughout India, and in some neighbouring regions of Pakistan and Nepal. They have also been introduced into South Africa, Italy, Mexico, and Texas; while the Italian population died out in the 1940s, and the South African ones remain confined to their ranches, some of those in North America managed to escape, and can still be found wild in those areas today.

Bovines: Bushbuck and More

Bushbuck

Of all the many kinds of antelope found in Africa, the one that is most widespread is the bushbuck (Tragelaphus scriptus). Bushbuck are essentially found throughout the whole of sub-Saharan Africa outside of the densest parts of the Congo jungles, the Kalahari Desert, and the drier parts of the veldt and the Horn of Africa. That's a huge range; everywhere from Senegal to Angola, and from Djibouti to the Cape.

Bushbuck are the smallest of the African bovines, sufficiently so that any resemblance to cows beyond the basics true of all antelopes (horns, cloven hooves, and so on) is quite absent. Specifically, they stand 60 to 100 cm (2' to 3'3") tall at the shoulder, with the males slightly taller, and much more heavily built, than the females. They do, however, much more closely resemble their larger immediate kin, such as bongos, having a somewhat similar coat colour, and the same twisted horns that make about a single turn along their length. Unlike in bongos, however, the horns are only found in the males - a feature shared with another close relative, the swamp-dwelling sitatunga.

Bovines: Bongos in the Jungle

Bongo

Broadly speaking, the more cow-like bovine species tend to be found wild in forested habitats, often with quite dense foliage. In contrast, the antelope-like bovines tend to prefer more open wooded country or scrubland. But these are both very crude generalisations. Plains bison and yak, for example, both live in open terrain, and, on the other side of the coin, there are bovine antelopes that prefer forest to savannah.

Among these is the bongo (Tragelaphus eurycerus), which can be found in three, entirely separate, regions of equatorial Africa. The two main populations live along the southern coast of West Africa, from Guinea to Benin, and across the heart of the Congo Basin and surrounding jungles, from Cameroon and Gabon in the west almost to the Ugandan border in the east. The vast majority of bongos live in these two areas, which are nonetheless separated by over 1,000 km (620 miles) of intervening land (much of which is Nigeria). The third population is found only in a couple of tiny mountain refuges in Kenya, where they cling to the forested slopes, migrating up and down them each year as the weather changes.

Bovines: Spiral-Horned Kudu

Greater kudu (male)

Most "bovine antelopes" - those more closely related to cows than, say, sheep - are collectively referred to as "spiral-horned antelopes", because of the shape of their horns. In many species, the horn makes a single 360° turn once it is fully grown, but others have several twists to the spiral. In elands, the twist is tight, marked clearly by a ridge that runs along one edge, but in no species is the spiral perhaps more evident than in the two species of kudu, where the horns are relatively narrow, with a loose, open curve.

The more widespread and common of the two species is the greater kudu (Tragelaphus strepsiceros). These are particularly large antelopes, with the males standing up to 157 cm (5' 2") in height at the shoulder, and weighing up to around 270 kg (600 lbs). Indeed, they are the second tallest species of antelope, after their relatives, the elands. Females are quite a bit smaller, at around two-thirds of the weight, and standing no more than 132 cm (4' 3"). As with most other spiral-horned antelopes, the coat is a brownish colour, with narrow vertical white stripes on the flanks.

The horns, are, however, their most distinctive feature - the second half of their scientific name means "twisted horn". These are typically about 100cm (3' 3") long in fully grown males, and would be even longer were you to measure them along the curve, rather than in a straight line from the base to the tips. Although the occasional horned female has been reported, this seems to be a rare aberration. Even in males, the horns grow slowly enough that it is possible to age younger individuals by how well they've developed; it takes two years to complete the first full turn, and the final adult form, with two to two-and-a-half turns, is reached by around four and a half years.

Bovines: The Largest Living Antelopes

Common eland (male)

Over the last few months I have looked at the various species of cow-like bovine, including such creatures as yak, bison, and water buffalo. But we've known, since at least the 1950s, that there are also a number of other animals that are more closely related to cattle than they are to other kinds of bovid. Although the choice of terminology is ultimately arbitrary, these are commonly considered to be members of the subfamily Bovinae, and thus can also be described as "bovines".

They are not, however, physically very cow-like, and so are instead described by the more generic term "antelope" - which really just means "any bovid that's not obviously some kind of cow, sheep, or goat". The great majority of antelopes are therefore not "bovines" in any sense, belonging to their own distinct subfamilies. The majority of those that are are commonly referred to as spiral-horned antelopes. While all bovines are said to have some degree of spiral growth pattern to their horns, it's only in these antelopes that it's really obvious, with multiple turns of the spiral clearly visible to the observer.

Bovines: Mini-Buffaloes

Anoa (probably lowland species)

One of the themes that we see played out from time to time in evolution is that of insular dwarfism. Here, a population of some large animal is trapped on an island or (less commonly) some other isolated locale from which they cannot easily escape. Faced with a limited food and other resources, over sufficiently large periods of time, they become smaller, making the best of whatever there is. (There is also a related phenomenon of insular gigantism, whereby really small creatures become larger, likely because they no longer need to hide from predators that happen to absent on their island).

Today, there are no tiny elephants or rhinos, but bovines - where the males of some species can weigh upwards of a ton - are a different matter. At least two, and probably three, species of living bovine are examples of exactly this principle in action.

The reason that I'm vague about the numbers here is that there is still some dispute about the nature of the anoa. There are usually regarded as being two species: the lowland anoa (Bubalus depressicornis), and the mountain anoa (Bubalus quarlesi). But not everyone agrees that the two are distinct. The first formal suggestion that they might be separate species was made by Dutch scientist Pieter Ouwens, better known for being the first scientist to describe the komodo dragon. That was back in 1910, and at least as late as the 1960s, there was still general agreement that the two forms were physically distinct, and that they likely favoured different terrain.

Bovines: Will the Real Buffaloes Please Stand Up?

African buffalo

Outside of North America, the term "buffalo" refers not to the bison, but to two (or more) species of bovine native to Africa and Asia. In the wild, by far the more common of the two is the African buffalo (Syncerus caffer), which has the distinction of being the only non-domesticated species of truly cow-like bovine that isn't even slightly endangered.

African buffalo are very numerous, and found across wide swathes of Africa south of the Sahara. This includes a fair number of different habitats, and African buffalo vary not just in their habits and preferences, but in their physical appearance, across that range. They were first described scientifically by Swedish naturalist Anders Sparrman in 1779, based on an animal found at Algoa Bay, near present-day Port Elizabeth in South Africa. Given the association with that general part of the world, the alternative name of "Cape buffalo" is often used for the animal, although that seems a little restrictive for something so widespread to me.

At any rate, over the next century and a half, scientists racked up descriptions of an impressive number of subspecies - by 1913, at least 21 were recognised. In the hundred years since, that number has steadily declined as it has become clear just how tiny the differences between some of them were. Even now, the question isn't entirely settled. There is agreement that there at least three subspecies, but there might be more, and it has been also been argued that some of them are genetically distinct enough to be full species in their own right.

Bovines: Almost-Cows of Southeast Asia

Banteng female

So far, in my survey of the Bovinae subfamily, I've mostly looked at species that most Westerners are, if not personally familiar with, at least well aware of: domestic cattle, yak, and bison. The exception, perhaps, is the gaur, an unusually large (or, at least, tall) species of bovine native to Southeast Asia. The gaur, however, is not alone, for there are two other very closely related species that are native to the same general area.

The better known of these is likely the banteng (Bos javanicus), an animal that was once found from eastern India to southern China, and across the whole of the Southeast Asian peninsula; there are even distinct subspecies on Borneo and Java. Today, the wild animal is extinct in India and Bangladesh, and found only in a few limited patches elsewhere. It has been formally listed as an endangered species since 1996, but the species as a whole is in much better shape than that would suggest... because this is another species that has been successfully domesticated.

The domesticated animals are known as Bali cattle, and they are in wide use across Indonesia and Southeast Asia. Over the years, a number have escaped from captivity, with the result that at least some feral populations are now found on a number of Indonesian islands that did not (so far as we can tell) ever host the genuinely wild form. More dramatically, British troops took some of them to the Northern Territory of Australia in 1849, but released them all just one year later when crop failure forced them to abandon their new settlement. The resulting feral animals are still there, and, while there are several thousand of them, all in Garig Gunak Barlu National Park near Darwin, since they are descended from just 20 imported animals, it's perhaps unsurprising that they are now highly inbred.

Bovines: Bison bison bison, the Bison

American bison

The only bovine native to the Americas is the bison (Bison bison). Indeed, it is one of only five species of bovid that fit that description - the other four all being members of the goat subfamily. To give at least some variety, however, there are two generally recognised subspecies of American bison, one of which, following the standard rules for naming subspecies, necessarily goes by the wonderfully tautonymous trinomial Bison bison bison.

This, in fact, is the plains bison, which was once found across pretty much the whole of what is now the contiguous United States, leaving aside only the Pacific and Atlantic coasts and the arid deserts of the Southwest; they also lived in far northern Mexico and up as far as central Alberta. The other subspecies, commonly known as the wood bison (Bison bison athabascae) is native to north-western Canada and to Alaska. Physically, it's slightly larger than its southern relative, which means that, in the absence of American rhinos, elephants, and so on, it is, in fact, the largest living, land-dwelling animal of any kind native to the Americas.

Because bison are, indeed, pretty big. A fully grown male can be anything up to 195 cm (6' 6") tall at the shoulder, and can weigh as much as a tonne (2,200 lbs). Females are, admittedly, quite a bit smaller, although at a maximum of about 180 cm (6') and 550 kg (1,200 lbs) they're still pretty big - and closer in size to the males than, say, those of yak. The horns are also about the same size in both sexes, although (relative to the rest of the animal) fairly small in comparison to those of most other bovines.

Bovines: The Other Domesticated Species

Domestic yak

The large physical size and herd mentality of bovines makes them ideal subjects for domestication, especially as beasts of burden. So much so in fact, that not only have true cattle been domesticated at least twice, but no less than four other species of bovine have also been domesticated on a wide scale since ancient times. (I am ignoring here the fact that bison, for example, have been farmed for their meat in recent decades, and am only considering "traditionally" domesticated animals). Unlike true cattle, the wild ancestors of these animals remain alive - all four of them in Asia.

One of the more familiar of these "other" domesticated bovines is the yak (Bos grunniens), which is descended from... well, the wild yak (Bos mutus). Today, wild yak are found only on the Tibetan Plateau, although in medieval times they may have lived much further north and west. One small population is known to cross the border into Ladakh in far northern India at certain times of the year, but they are otherwise extinct outside of China, having vanished from Nepal as recently as the 1990s.

Bovines: How Cows Were Domesticated

Taurine bull

Cattle (Bos taurus) are among the most familiar of herbivorous domestic animals. According to recent estimates, there are well over a billion of them on the planet today, found almost anywhere that humans practice agriculture. In the west, we are familiar with them as a source of beef or milk, but, in many parts of the world, they are still used primarily as draft animals ("oxen"). Indeed, they are important enough that Christopher Columbus first brought them to America in 1493, on only his second voyage to the New World.

But one unusual thing about cattle is that there is no living wild form of the animal. There are still wolves and wildcats, wild goats, wild sheep, and wild boars... but there are no wild cattle. They're not unique in this respect, to be sure, and there are a few populations of feral cattle around the world (that is animals living in the wild whose ancestors were domesticated), but it's a fairly exclusive club nonetheless. And, obviously, it wasn't always the case.

Bovids and Bovines - What Makes a Cow?

The cattle family, Bovidae, is the largest and most diverse of the various families of big herbivorous mammals, with only the deer family coming close to it in number of species. The majority of its members are antelopes of one kind or another, and it also includes goats and sheep, as well, of course, as the cattle themselves. Given the number of species within it, and the fact that a bison, say, has a number of obvious differences from a gazelle, there have been various attempts down the years to group its members into subfamilies or other smaller groupings.

However, opinions have differed as to quite how this should be done. How many significant sub-groups are there, which of them are deserving of the rank of "subfamily", and which animals go with which? Even with modern techniques for analysing genetics and coming up with computer-generated metrics of relatedness, the answers to these questions are not entirely clear. Partly that's because the first two questions at least are essentially arbitrary, and partly because we haven't really got enough samples from some of the more obscure species to really nail the family tree down. There could be anything up to sixteen subfamilies, or just two, or somewhere in between.