Tetrapod Zoology

I said the other day that I was going to have go all ‘text lite’ for a while. Here is my solution: a series of short posts, one per day, each of which features a different stuffed megamammal. Yes, welcome to day 1 of stuffed megamammal week. All of the stuffed megamammals you’re going to see were photographed in the excellent collection at the National Museum of Ireland, Dublin. So here we go.

This remarkable creature is a Hartebeest Alcelaphus buselaphus, and, specifically, it’s a female hartebeest belonging to the subspecies known as the Khama, Cape or Red hartebeest A. b. caama. Like so many subspecies, the Khama was originally regarded as a distinct species, and in fact still is by some. Those who do regard it as a species split it into two subspecies: A. caama caama (described in 1816 and extinct by 1940) and A. caama selbournei.

Though some controversy remains, it does now seem that the massive variation present among the many regional forms of hartebeest – about 70 taxa have been named, ranging across most of sub-Saharan Africa* – can all be accommodated within a single species, and indeed there are various hybrid zones where confusing intermediate morphologies are present (Ruxton & Schwarz 1929). Some authors – notably Jonathan Kingdon – regard Lichtenstein’s hartebeest or the Nkonzi as a subspecies of A. buselaphus, but most workers regard it as a separate species that even warrants it own ‘genus’: Sigmoceros lichstensteinii. In overall shape, all hartebeest are alike: they are long-legged, short-necked alcephaline antelopes with tall shoulders and very long, narrow faces.

* And, previously, north of the Sahara too. The ‘original’ hartebeest, the nominate form of the species, is the now extinct Bubal A. b. buselaphus, a semi-desert specialist and the smallest of all the hartebeest. Formerly widespread across northern Africa, it was hunted to extinction by about 1923 (or perhaps a little later).

Hartebeest might be similar in shape, but they differ in colour (varying from dark red with black patches to light golden brown) and in horn shape. In some forms, the horns are widely spaced, relatively short and with backswept tips, while in others the horns have long, virtually conjoined bases (or pedicels) and curve outwards and then inwards at their tips, thereby forming tall, lyre-like shapes. In fact Dutch settlers thought that the shape of the horns resembled that of a heart, hence the common name. The adjacent photo shows some hartebeest horns I photographed in an antiques shop last year. I think they are Khama horns.

Kingdon (1997) proposed that the forms with the short horn bases, like the Kongoni A. b. cokei, Korkay A. b. swaynei and Tora A. b. tora (all of which are from eastern Africa), are more primitive than the tall-pedicelled forms like the Khama (of the Kalahari) and Kanki A. b. major (of western Africa). This has been mostly confirmed by DNA evidence (Flagstad et al. 2001, 2004) which indicates that hartebeest originated in eastern Africa and colonised the rest of the continent from there. DNA and fossil evidence also indicates that the extant alcelaphines (hartebeest, wildebeest, topi and kin) became extinct across most of their Pleistocene range and survived into the Holocene in just a few refugia: for hartebeest, two of these were north of the equator and one was south of it (Arctander et al. 1999, Flagstad et al. 2001). It is well known that alcelaphines are a very young clade (perhaps less than five million years old), and that, within their short history, they diversified rapidly. Data shows that their diversification events match key climatic changes, and in fact their evolution inspired Elisabeth Vrba to come up with the ‘turnover pulse’ hypothesis.

And… so much more to say, but I gotta stop there.

Refs – -

Arctander, P., Johansen, C. & Coutellec-Vreto, M. A. 1999. Phylogeography of three closely related African bovids (tribe Alcelaphini). Molecular Biology and Evolution 16, 1724-1739.

Flagstad, Ø, Olsaker, I. & Røed, K. H. 2004. The use of heterologous primers for analysing microsatellite variation in Hartebeest Alcelaphus buselaphus. Hereditas 130, 337-340.

- ., Syvertsen, P. O., Stenseth, N. C. & Jakobsen, K. S. 2001. Environmental change and rates of evolution: the phylogeographic pattern within the hartebeest complex as related to climatic variation. Proceedings of the Royal Society of London B 268, 667-677.

Kingdon, J. 1997. The Kingdon Field Guide to African Mammals. Academic Press, San Diego.

Ruxton, A. E. & Schwarz, E. 1929. On hybrid hartebeests and on the distribution of the Alcelaphus buselaphus group. Proceedings of the Zoological Society of London 38, 567-583.

Comments

#1 Hai~Ren
April 6, 2009

Wait a minute, isn’t the hartebeest Alcelaphus?

Awesome post. I’ve always been very fascinated with African antelopes, and the alcelaphines are especially interesting to me. I’m wondering just how divergent the nkonzi is from the other hartebeests to warrant such a wide separation at the genus level.

Despite their widespread range, hartebeest seem to be seriously overlooked in documentaries and books on African wildlife, which will focus heavily on blue wildebeest and maybe show a bit of topi here and there, but neglect to mention the black wildebeest, hartebeest, nkonzi, blesbok, bontebok, and hirola. So far, I’ve only ever seen fleeting glimpses of kongoni and khama in documentaries, but never any of the other hartebeest subspecies. Similarly, the only topi subspecies I’ve ever seen is the topi (Damaliscus lunatus jimela), but never any of the other subspecies like the tiang (Damaliscus lunatus korrigum) or tsessebe (Damaliscus lunatus lunatus).

Speaking of which, I wonder why the tsessebe’s relation to the topi mirrors that of the hartebeest, in that the southern African form is often considered a separate species. And what’s up with the Bangweulu tsessebe (Damaliscus superstes)?

By the way, I recall offhand reading a book in the late 90s which mentioned that the bubal hartebeest was known to inhabit parts of the Middle East in early historical times. I wonder if that has been verified or not. It’s a little odd that few desert-adapted antelope seem to be found in both North Africa and the Middle East; the only ones I can think of are the addax and dorcas gazelle, and these only extended as far as Israel and Jordan. What kept these species out of the Arabian deserts? Likewise, what kept the Arabian oryx and the various gazelles of the Middle East from invading the Sahara?
#2 Darren Naish
April 6, 2009

Thanks for that, I agree wholeheartedly: wildebeest and gemsbok and such get covered on TV at length, but tsessebes and so on are positively neglected (though a lot of cool stuff has been included in Big Cat Diary). Silly typo corrected. Re: Bubal, yes, some sources say that they were present in Palestine.
#3 Dartian
April 6, 2009

Hai~Ren:

hartebeest seem to be seriously overlooked in documentaries and books on African wildlife, which will focus heavily on blue wildebeest

You’ve discovered the infamous Serengeti bias… There seems to be almost a tradition of not straying off the beaten path among many wildlife documentary makers. Which means that, when filming in Africa, they will primarily go to Kenya – Tanzania – Uganda or, to a lesser extent, South Africa. (Zimbabwe used to be well-frequented too, until Mugabe went nuts.) When was the last time you’ve seen a mainstream nature documentary filmed in, say, Senegal, Chad, Cameroon, Nigeria, or Mozambique?

This inevitably affects the range of animal species that you’re likely to see portrayed. For example, Thomson’s and Grant’s gazelle are staple items in documentaries, but how often do you get to see Speke’s gazelle Gazella spekei, red-fronted gazelle Eudorcas rufifrons, or Soemmering’s gazelle Nanger soemmerringii? They all live in Africa, just not in those countries usually frequented by nature documentary makers.

As for why antelopes aren’t widely distributed outside of the African continent… Good question, to which I’m not sure there is an equally good answer. As so often is the case, however, the modern taxa used to be more widespread in the past (Miocene-Pliocene, in this case). ‘Antelopes’ of various kinds are common in the Eurasian Neogene fossil record. To what extent these were closely related to the modern African taxa is, of course, another matter. But I think I’ve seen some sources* claiming that alcelaphine (or at least alcelaphine-like) antelopes were part of the Indian Siwalik faunas.

* I don’t have the references handy, unfortunately.
#4 Dartian
April 6, 2009

Addendum to my previous comment:

alcelaphine (or at least alcelaphine-like) antelopes were part of the Indian Siwalik faunas

…and that would be Damalops, which was classified as a true alcelaphine by Vrba (1979).

Reference:

Vrba, E. 1979. Phylogenetic analysis and classification of fossil and recent Alcelaphini Mammalia: Bovidae. Biological Journal of the Linnean Society 11, 207-228.
#5 Jerzy
April 6, 2009

Cool post. Indeed, it is one overlooked group of antelopes.

I find topi to be very beautiful creatures.
#6 Hai~Ren
April 6, 2009

Dartian: You left out Botswana and Namibia as other countries that are heavily represented in wildlife documentaries. =) Oh, if I could have a penny for every documentary shot in the Okavango/Chobe/Etosha/Kalahari…

What intrigues me is that even within the Serengeti-Masai Mara, the hartebeest are hardly featured at all. Perhaps the much larger wildebeest herds play a more prominent role in the local ecology, but surely the hartebeest deserve at least a bit of airtime? We already get topi standing on termite mounds and getting ripped apart by spotted hyenas while lekking, surely there’s something about hartebeest that’s worth filming?

Same goes for other antelope species in the Serengeti; where are all the roan antelope, fringe-eared oryx, eland, bushbuck, bohor reedbuck and waterbuck?

I think the following species overwhelmingly dominate all footage of African antelope:

1) Blue wildebeest
2) Impala
3) Thomson’s gazelle
4) Greater kudu
5) Gemsbok

It’s as if the rest of the antelope species don’t exist at all, except to make token appearances in the scenery or to get chased by some carnivore.
#7 Hai~Ren
April 6, 2009

P.S Just realised that the hartebeest is Alcelaphus busephalus
#8 Darren Naish
April 6, 2009

Three different spellings of the same name: I’ve done well so far
#9 Hai~Ren
April 6, 2009

Ack. Just realised my misspelling, I meant buselaphus. I hate these tongue-twisting names. I’d grown up associating the name with Alexander the Great’s horse…
#10 Dartian
April 6, 2009

I’ve done well so far

If it’s any consolation, I didn’t notice a single one of all these seemingly endless misspellings until they were pointed out…
#11 Metalraptor
April 6, 2009

Out of curiostiy, I was wondering if the National Museum in Ireland has any prehistoric animals in it, other than Megaloceros of course. I once heard from someone that the Irish Museum was a “museum within a museum”, looking just like a Victorian museum from “back in the day”. They may have remodeled since then, though.
#12 tai haku
April 6, 2009

@ Hai-Ren – the best place to see different antelope species on tv in my experience has sadly been on hunting shows in the US. Those guys will correctly identify and provide details on a number of species [sadly for nature fans they do tend to then shoot them].
#13 Brendan
April 6, 2009

Doesn’t this have 6 subspecies?

brendan
http://www.wildramblings.com
#14 Mark Lees
April 6, 2009

Since many of the wildlife documentaries are set in the Masai Mara it is surprising we don’t get to see topi and hartebeest more often, since both occur there, and hartebeest is common.

Indeed I saw quite large numbers of hartebeest in the Mara – the subspecies cokei, sometimes referred to as the Kongoni. You mention their having “very long, narrow faces”, and when I first saw them , I thought their expression looked permanently sad, morose even. They also looked like the horns were an after thought. Topi were much less common.
#15 David Marjanović
April 6, 2009

Hunting shows? On TV?

Strange country.
#16 Dr Vector
April 6, 2009

I was going to have go all ‘text lite’ for a while. Here is my solution: a series of short posts [sarcastic emphasis added]

Ah, good to see that’s working out for you. You may need to recalibrate your short-o-meter, however.

Seriously, great idea, fascinating post. And if they’re all as “short” as this one, by the end of the week you’ll have another book written. Bastard.
#17 Allen Hazen
April 6, 2009

Like Dr. Vector, I’m bemused by your idea of a “text lite” post! This one was fascinating.

Etymologies of these names? “Elaphus” is Greek for deer? “Alc” maybe from the Latin for elk (“elk” interpreted loosely, as “top of the line deer”?)? “Bus”???? (maybe from Greek “bous”, cow, but the “s” in a combining form is strange)? Europe isn’t great antelope country (give or take the chamois), and the classical European languages don’t seem to have provided for for them: zoological names for antelope seem to be cobbled together from words for more or less analogous European ungulates.
#18 Hai~Ren
April 7, 2009

I was suddenly thinking about the extinct giant alcelaphine Megalotragus, and found this picture via Google. I wonder how accurate it is in depicting the shape of the horns.
#19 Darren Naish
April 7, 2009

That picture is by Eric Alibert and is from Jean-Christophe Balouet’s Extinct Species of the World: I have a photo of the Megalotragus skull with me now (photographed alongside the other alcelaphines Damalops and Beatragus) and, no, it isn’t accurate. The horncores of Megalotragus curve outwards and then inwards at their apices, but they don’t sweep as far outwards as the horns do in the reconstruction. In fact, they’re not all that different from extant hartebeest horns in this respect. Furthermore, that colour scheme is extremely unlikely for an alcelaphine.

Incidentally, some authors regard Megalotragus van Hoepen, 1932 as synonymous with Rhynotragus Reck, 1925.
#20 Hai~Ren
April 7, 2009

Quite right, Darren; in fact, the colour scheme reminds me a lot of sable antelope, which aren’t that closely related anyway. Some sort of brown or grey would probably be more appropriate.

Thanks for sending me the picture as well. Indeed, the difference in horn shapes between the 3 species is quite apparent, and does show that Megalotragus horns were quite similar to those of Alcelaphus.
#21 Willem van der Merwe
January 26, 2010

Hi Darren! I’m a wildlife artist living in South Africa. I’m very well acquainted with hartebeests … we have some living in our municipal nature reserve here. They are remarkable-looking anteloped! The horns, the face, even the shape of the body, and the short, brush-like tail … even the way they run … everything is unusual and distinctive.

Some comments.

First, the specimen you photographed seems quite faded! Living animals are rather rich reddish brown, with glossy black patches on the face, the limbs, and the tail.

The horns you have further on in the article, do indeed look like Khama horns.

Over here we simply call them Red Hartebeests. Actually, the name does not come from the heart-shaped horns. It comes from the Dutch words ‘Hert’ which means ‘Deer’ (you even have the word ‘hart’ in English for deer), and ‘Beest’ which means ‘Cow’ or ‘Ox’. In other words, they called it a kind of deer-like cattle. The thick neck made them think more of cattle than antelope … just like with Wildebeests (‘Wild Cattle’ in Dutch).

If the Khama is regarded as a species, then it is divided into the two subspecies as you say … but when it is relegated to being a subspecies of the African hartebeest, what happens to its own ‘subspecies’?

I wonder if there are any museum specimens of the extinct ‘subspecies’ or version, A. caama caama (or whatever it would be named if A. caama itself is a subspecies of A. buselaphus)? Even over here in South Africa there’s very little info on it…

Another very remarkable and very ignored Alcelaphine that we have here, is the Black Wildebeest, Connochaetus gnou. Compared with the Blue Wildebeest (which many people might believe is the one and only Wildebeest) it gets about zero publicity.

Seeing as the Hartebeest, Alcelaphus buselaphus, has so many regional forms here in Africa … I wonder, if they survive and diverge even more, we would at some time have a great many separate hartebeest species. I’m wondering how long that might take … how long does a species take to fully evolve … a hundred thousand years, perhaps?

At any rate, thanks for your great articles!