Tuesday, November 14, 2006

Giant killers: macropredation in lions

By now you have probably heard that episode 2 (‘Great Plains’) of the BBC’s series Planet Earth (currently in its second series) included amazing footage of the elephant-killing lions of Savuti in Chobe National Park, northern Botswana. While most people ‘know’ that elephants are immune to predation thanks to their size, nobody has told this to the Savuti lions. Hunting at night, when the elephant’s poor night vision puts them at a major disadvantage, the lions co-operate as a pride of about 30 individuals to bring down and dispatch elephant prey. It is amazing. But, as usual, the media is leading us all horribly astray.

We should make clear to begin with that these are not just any old lions, behaving spontaneously or opportunistically: they are a specialised, highly experienced population that have, uniquely, become elephant killers. While there are some major questions as to how the Savuti lions learnt to do this, Planet Earth didn’t, unfortunately, touch on how old this culture is, or how it originated. It is thought that the Savuti lions have learnt over time to kill bigger and bigger prey, each time winning success by the virtue of their large pride size. Lions elsewhere can – opportunistically – kill Cape buffalo Syncerus caffer* (weighing c. 1000 kg) and sometimes hippo Hippopotamus amphibious (c. 1500-3500 kg), and it has been speculated that, after learning to successful tackle and kill hippo, the lions became bold enough to begin regularly taking juvenile elephants, eventually moving up to adults. And if you’re wondering: YES, the Savuti lions have been recorded attacking and killing adult elephants.

* Though note that some lion populations are specialist buffalo-killers. In Tanzania’s Lake Manyara National Park, George Schaller (1972) reported that an amazing 62% of all lion prey was made up of Cape buffalo, with 81% of this 62% being adult male buffalo. Buffalo-killing is also important to the lions of Kruger National Park, and studies here have shown, significantly, that male lions are not just frequent and successful hunters: they are also the lions that are best at killing buffalo (Funston et al. 1998).

A few opportunistically recorded events may have encouraged the lions to view elephants as potential prey. In their National Geographic film Ultimate Enemies, wildlife film-makers Dereck and Beverly Joubert recorded a case where, after a fight with another bull, a defeated elephant lay, wounded, on the ground. Understandably, the elephant’s misfortune became the hungry lion pride’s gain. Wildlife photographer and travel writer Leigh Kemp recorded a case where an old, weakened bull that collapsed and became unable to stand was discovered and eaten (while still alive) by opportunistic lions. It is tempting to suggest that these events and others like them might have been catalysts in encouraging the development of elephant-killing in the Savuti lions. Numerous other instances of elephant-killing have been filmed and documented by the Jouberts, and in 1997 they published a book covering this behaviour in depth (The Lions of Savuti: Hunting with the Moon, published by National Geographic).

The Lions of Savuti: Hunting with the Moon records something like 15 years of observations, and even in 1990 the Jouberts were estimating that about 20% of the Savuti lion’s diet was made up of elephant. I would love to know if the behaviour goes back further than this, as personally I find it highly unlikely that this behaviour really is something that the lions have ‘just learnt’. Historically, Africa was filled with a lot more lions than it is now, not to mention elephants, and given the extraordinary behavioural flexibility of lions* I suspect that elephant-killing is something that lions have practiced many many times in the near and distant past.

* If you’re read Bruce D. Patterson’s outstanding The Lions of Tsavo (Patterson 2004) you’ll know that studies of the Serengeti-type lion that we’re all so familiar with (e.g. Schaller 1972) have ‘created an orthodoxy around lion biology that applies poorly to the species elsewhere’ (p. 138).

The fact that the Jouberts were photographing and filming this behaviour negates one of the claims that have appeared as a result of Planet Earth’s coverage: this being that the BBC were the first to film this behaviour (not that anyone working for the BBC has said this, so far as I can tell). In fact Ultimate Enemies, showing scenes of night-time elephant predation by Savuti lions, was broadcast in North America in 2004. This is not to downplay the BBC’s commendable efforts, however, and it is clear that obtaining the sort of footage they did is tremendously time-consuming, dangerous, and requires a monumental amount of effort. With panicked elephants lumbering around in the dark, and surrounded by hungry and aggressive lions that routinely kill animals weighing many hundreds of kilos, the camera teams were in the middle of the bush, in the middle of the night, in small jeeps with open sides and windows.

What did we actually get to see? The answer to this is both positive and negative. To begin with, it seems that the lions used psychological warfare to intimidate and confuse the elephants: loud roaring in the dark. This behaviour has been recorded in other lion populations and also in leopards, and it seems that the idea is to scare prey into making an ill-thought dash for ‘safety’. Paying particular attention to juvenile and adolescent elephants, especially those that were separated from the rest of the herd, the lions were then shown attacking the hind legs and haunches of fleeing elephants, biting and clawing and hanging on to the pursued animal. And that… is about it. Here’s where we come to the negative, particularly problematic, part of this whole story.

We empathise with elephants. And, somehow, seeing them being killed and eaten by big cats is, for many people, just wrong. That may or may not be a justifiable point of view, but what is undeniable is that elephant-killing is protracted, unpleasant, and gory. Consequently almost none of the actual killing was shown. By clawing and biting at the elephant’s legs, the lions hamstring a chosen elephant, and also use the combined weight of multiple individuals to bring it down. This apparently happens surprisingly quickly. From spotting an elephant, to pursuing it, to getting it to collapse: all can take as little as 30 seconds. Once an elephant is down, some of the lions work on clamping its trunk shut, and I presume that they might also attack the throat and mouth. Like it or not, we can assume that lions at the other end of the animal will now begin feeding. The elephant might take about 30 minutes to die. It does not sound nice, or look nice.

I empathise with elephants, and do not enjoy the thought of them being killed. But the fascination that I have for animals makes me want to know more about what actually happens. This is a natural act of predation: sure, it’s not pleasant, or pretty, but I want to know what happens. For me, the footage was ultimately disappointing, then, in showing bugger all (worth noting here is that views on the screening of acts of predation are starting to change. See Finally: big cat kills uncensored and uncut).

What makes this all the more frustrating is the implication from some that the lions are downright nasty, committing an evil, murderous act that is heinous and unjust. An article – titled ‘The killing fields’ – that appeared in Times2 (a supplement to the British newspaper The Times) described the footage as ‘possibly the most shocking natural history footage you will have seen’. It went on to state that ‘If you have any sentimental feelings about lions, prepare to lose them’. I’m sorry, but that’s crap. The appreciation I have of lions and their amazing behavioural flexibility and unique social system is increased by the knowledge that they have learnt to kill elephants. Yes it’s gory, and – no doubt about it from our point of view – upsetting and even horrific, but it is an amazing thing that we should wonder at.

Coming eventually: agamas, tupuxuarids, fake Chinese turtles, temnospondyls for beginners, kinglets and the passerine supertree, more on sea snakes, anguids, giant eagles and plethodontids, those lost tree frogs, storks and (one day) rhinogradentians. For many of these posts you'll have out check Tetrapod Zoology at its new location here.

Refs - -

Funston, P. J., Mills, M. G. L., Biggs, H. C. & Richardson, P. R. K. 1998. Hunting by male lions: ecological influences and socioecological implications. Animal Behaviour 56, 1333-1345.

Patterson, B. D. 2004. The Lions of Tsavo: Exploring the Legacy of Africa’s Notorious Man-Eaters. McGraw-Hill, New York.

Schaller, G. 1972. The Serengeti Lion. University of Chicago Press, Chicago.

22 Comments:

Anonymous Roger B. said...

Another amazing sequence on 'Planet Earth'. The kill itself wasn't shown - it was presumably protracted and very bloody.

2:22 PM  
Blogger Hai~Ren said...

I always thought that lions killing elephants had been known for quite some time; it was featured in National Geographic a decade ago, no?

Still, for a felid that lacks sabre teeth, it's amazing (and gruesome) to see such a bloody spectacle. One imagines the sabretooth cats were more efficient at quickly dispatching similarly thick-skinned prey. (That is, if they really did hunt proboscideans)

10:24 PM  
Anonymous Ville Sinkkonen said...

yeah...I'm pretty sure I have seen some national geographic documentary that showed large pack of lions taking down adult elephant.

11:33 PM  
Anonymous karen said...

Living here near Chicago...we can visit the Lions of Tsavo (they are here at the Field Museum)...

And yes I have Patterson's book about his adventures in capturing them. Quite the tale of the Ghost and the Darkness indeed.

:-)

11:50 PM  
Blogger Hai~Ren said...

In what ways are the Serengeti lions different from those living in other parts of Africa? Is it to do with their social lives?

9:40 AM  
Blogger The Ridger, FCD said...

Well, it's time to break out all the footage of adorable baby lions, I guess. Lions gotta eat, too, you know?

I remember a large brouhaha over showing them killing zebras on US TV (decades ago now, granted) so I can understand the relunctance to show the endgame of this elephant kill. But I also understand that lots of people have a professional interest in seeing how it works. (I have to admit I'm stunned by the notion of 30 lions working together to do *anything* - I suppose I'm one of those laymen whose idea of lions comes strictly from the Serengeti.) Maybe the dvd can have it as an extra? (half joking)

11:05 AM  
Anonymous Anonymous said...

Hi Darren,

Watched the sequence with interest. It harked back to my own 'theories' on how big-brained, large-eyed Homotheres hunted mammoths in Ice Age N. America.

With regard to the Tasavo man-eaters, I'd also recommend a book I recently read called Man-Eaters of Eden, which actually considers that the lions of Kruger National Park are by far and away the most successful historical man-eaters. Check it out.

11:12 AM  
Blogger Darren Naish said...

Hai-ren asked...

In what ways are the Serengeti lions different from those living in other parts of Africa? Is it to do with their social lives?

It would be easy to write a whole blog post on the variability within lions (maybe, one day, I will). Briefly, lions across Africa and Asia differ tremendously in body size, in skull shape, in size, extent, colour and presence of the mane (not all lions have manes), and in social system. The Serengeti-type lions that we are most familiar with live in prides of about 6 females defended by coalitions of 2-4 males, and these males have thick, tawny-coloured manes. This isn't typical for all populations. In the Tsavo lions, for example, single maneless males defend prides of more than 7 females, and there are various other deviations among the species.

1:18 PM  
Anonymous Anonymous said...

I agree with your comments about not taking too sentimental a view about natural hunting. But conversely I am annoyed by the way that some wild life presneters identify too strongly with the predators. The series 'Big Cat Diary' provided plenty of examples of commentators 'justifying' big cat hunting behaviour with the 'its only nature' response, while taking a very different line when the herbivores strike back. I recall one scene where an elephant seemed to get fed up with a pride and charged them, trampling many of the cubs to death. That is just as 'natural' - and the commentator's remarks betrayed a onesided viewpoint.

1:24 PM  
Anonymous Dan Koehl said...

Interesting remarks, I linked to your blog from www.elephant-news.com.

It leads to further questions, like wether this occaur in Asia with tigers, and wether the Amur Tiger was present in russia 5 000 years ago, and if they and the Cave lion were predating on mammoths...
predating on mammoths, since the

7:52 PM  
Anonymous Carl Buell (OGeorge) said...

Great post Darren. As a member of a species that still tortures and kills other members of the same species over minor religious differences, I find my own squeamishness interesting. I've seen animals do things that are absolutely horrid to watch, but very interesting as behavior. I have my own special place in my head and heart for elephants, but I know for a fact that I would have hunted them mercilessly had I been born in a cave 10,000 years ago. I'm rather glad I live in a time and place when I can afford to be a little squeamish.

10:38 PM  
Anonymous Anonymous said...

Has enough work been done on lion DNA to clearly distinguish the extinct cave lion from the African lion? The maneless Tsavo lions could be relict -- relict is the wrong word -- cave lions, which were depicted as maneless; it could be they're all lions and the species is wildly variable, too, and I'm curious if anyone knows.

-- Graydon

11:50 PM  
Anonymous Filipe said...

Fantastic, is there any other difference besides pride numbers?
What about the teeth?

I wish they would make a site with all the details of the kill. A bit like National Geographic did with leopard seals (caution, it's gory).

8:16 AM  
Blogger Darren Naish said...

Quick response to Graydon's comment: it has indeed been suggested that the maneless Tsavo lions are a distinct, or even relict, type of lion, informally dubbed the 'Buffalo lion'. Tom Gnoske and Julian Kerbis Peterhans have proposed that Buffalo lions are a basal lion lineage, directly descended from the maneless lions of the Pleistocene, and in many respects distinct from Panthera leo proper (Gnoske & Peterhans 2000, von Buol 2000). It is this contention that led Bruce D. Patterson and colleagues to collect DNA from Tsavo lions. This data shows, surprisingly perhaps, that the maneless Tsavo lions do not differ from 'normal' eastern or southern African lions, and that the Tsavo lions do not warrant special taxonomic status (Dubach et al. 2005). Their manelessness must be secondary. The unusual physical and social attributes of the Tsavo lions seem to illustrate how ecomorphologically flexible lions are - a statement that is backed up by lots of additional variation seen within Panthera leo.

Ref - -

Dubach, J., Patterson, B. D., Briggs, M. B., Venzke, K., Flamand, J., Stander, P., Scheepers, L. & Kays, R. W. 2005. Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo. Conservation Genetics 6, 15-24.

Gnoske, T. & Kerbis Peterhans, J. 2000. Cave lions: the truth behind biblical myths. In the Field 71, 2-6.

von Buol, P. 2000. 'Buffalo' lions. A feline missing link? Swara: the Magazine of the East African Wildlife Society 23 (2), 20-25.

10:29 AM  
Anonymous Anonymous said...

Darren --

Thank you!

Nice to get a definitive answer, and I suppose that the ecomorphological flexibility is every bit as cool as relict cave lions would have been.

-- Graydon

11:41 PM  
Blogger The Ridger, FCD said...

It went on to state that ‘If you have any sentimental feelings about lions, prepare to lose them’.

I think they're right. "Sentimental feeelings" are not at all what you have; they're that sappy, animals are just like people, Disneyesque feeling that people have for "Our Furry Friends". It's the polar opposite of the "appreciation" you have.

Recently the Guardian had an interview with David Attenborough, which contained this:

------
"But I'm not an animal lover," he says emphatically. His face scrunches up in disdain. "Animal lover means sentiment; a cloying, anthropomorphising sentiment. I don't love earth worms or spiders. They're rivetingly interesting and they give me huge intellectual pleasure. And aesthetic pleasure, I suppose. But that's a different thing altogether."

That's not the same as loving them? "No. It's the word 'love' I don't like. The phrase 'animal lover' - well, it just grates on me! That's why I'm irritated when people use it on me. They say, Oh, you're an animal lover!" He glowers. "I say, I am NOT an animal lover."
------

People who prettify lions in their mind - people who are Lion King about them, for instance - would indeed be horrified at footage of them killing an elephant.

12:28 AM  
Blogger Darren Naish said...

Well said; I agree wholeheartedly.

10:13 AM  
Anonymous KeithB said...

In the National Geographic series "Be the Creature" with Chris and Martin Kratt, they do an episode on "Being a Lion."

They show a Pride of lions - sorry, I do not remember where in Africa - taking down a baby elephant. They pretty much show the whole thing, and this is a show aimed at *kids*.

6:50 PM  
Anonymous Raj said...

Pretty shocking documentary. To see such a large, "vegetarian", social being cornered by a group of smaller animals. Quite unfair.

But then, man is a small, cunning animal too. Sad or ashamed to be a human? Well...

Should also appreciate the courage, ruthlessness and creativity of the pride.

Guess it would've been okay if the elephants had retaliated in some way. The sadness would've mitigated.

5:58 AM  
Anonymous Anonymous said...

I guess I might be lacking in the sentimentality department; my reaction on seeing this was pretty much "cool!". Feeling sorry for the elephants didn't occur to me.

3:11 PM  
Anonymous Anonymous said...

i love animals and i hate that people have slaughterd them we need to stop this as soon as possible

9:04 AM  
Blogger Sunil said...

Lions are doing what they are supposed to be do. However, what humans are doing to all the animal species is appalling and pathetic. China/Japan/Korea/Turkey/Spain you name the country and they are murdering animals in thousands. According to some shark experts, most of the shark species population is down by 90%. Has any one thought about consequences of Chinese and Japanese brutal eating habits? Eating monkeys, dogs, and lion & Tiger steaks. One day we humans are on the menu across the Chinese food malls and that day is not very far. Chinese are gobbling up all the animal species at very rapid pace.

5:35 AM  

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