Common Bulbul and frugivorous birds

Common Bulbul nesting in Arusha

Thanks to doubtful comments from a colleague, next up in the common bird series is going to be the Common Bulbul. These birds are probably the most widespread birds in Tanzania and should be a familiar sight to everyone, with their dark blackish heads, brown back and tail, and dirty white underparts with yellow under the tail. They're typical garden birds, and often ignored. However, it is often the common birds that we know most about, because they are so easy to study. And common species, simply because there are so many individuals, often have a very irritant role to play in ecosystems - the common bulbul is no exception!

How to survive the Serengeti: predation, food and body size

Serengeti Lions eat a diversity of mammal species.

Another paper describing one of my favourite talks from the TAWIRI conference back in December has just been published, this time in the Journal of Animal Ecology (available to some here). Grant Hopcraft and colleagues are interested in why different grazing animals use different parts of Serengeti in different ways when they all eat grass. Why, for example, do you usually find buffalo around riverine areas, whereas gazelles tend to be on the open plains? At some level it's obvious that where you find an animal is the place that it survives best and there are two aspects to survival that most ecologists would agree are crucially important: where will you eat? And where will you be eaten? Although both buffalo and gazelles eat grass, and both are eaten by large cats, it's quite possible that grazers of different size will be affected differently by these same two processes. And that's what Grant and colleagues set out to test.

The Serengeti Story 2: the great migration

Lion admiring the massed migration on the plains, near Naabi, Dec 2011

The second part of the Serengeti Story is the tale of the great migration, the defining heart of the Serengeti Ecosystem. At the broadest level, this is an easy enough  thing to understand - thre are two very important environmental gradients across the ecosystem and the wildebeest (and zebra and eland and gazelles, etc.) are trying to maximise their access to important resources. So, let's start with the two important gradients: rainfall and nutrients, the remaining two of the big four we didn't cover in the first part of the story.

Average Serengeti Rainfall, adapted from here

Starting with rainfall, the broad pattern is for lots of rain in the north and west, and (much) less in the south and east. Perhaps more important still is the seasonal difference in rainfall patterns - most rain falls during the wet season, of course, and the wet season rainfall shows a similar pattern to the overall pattern. But dry season rainfall is the key - the far north and the far west have an average of 400mm of rain even during the dry season, and what's more that's fairly reliable rainfall - the rest of the ecosystem is either compeltely dry, or only ocassionally it by a shower every few years. There's also only one permanent river in the ecosystem - the Mara river in the north. So dry season rainfall means there's green grass to eat, and the Mara river means there's water to drink during the dry season in the far north - an obvious reason for migrant animals to be on the Kenyan / Tanzanian border during the dry season. (In fact the animals move around quite a lot at this time, following local patterns of rainfall and often crossing and recrossing the Mara river throughout their time up there.

A small crossing of the Mara: local movements, not migration, Sept 2011

As the rains become more widespread in November the animals quickly move south, heading away from the woodlands to the short grass plains of the Serengeti NP / Ngorongoro CA border. Why? Well, this is where the other important gradient comes into play, that of nutrients. And this is best understood by looking at the geology of the Serengeti ecosystem in the figure below. Orange areas are 540 - 1500 Million years old, grey areas are recent (within 65 Million years - most only 3 Million years old), Pink areas are over 2500 Million years old and tan coloured bits are also relative recent alluvial (flood) bits, derived from earlier shorelines of Lake Victoria.

Geology of Serengeti, detail from Ordanance Survey map, Saggerson 1961

Broadly speaking there are three geological areas in Serengeti - the southern areas with very recent soils formed on top of the ash deposits from the crater highlands (which form a hard pan that plants can't get their roots through, and only having shallow soil - as illustrated in this picture below froma cutting just east of Naabi gate), the western areas and the north-eastern areas. The north eastern areas are characterised by rocks formed over 2500 Million years ago, whilst the western areas have some more recent deposits from the rivers and different shores of lake Victroia. Unsurprisingly, the nutrients from the ancient rocks in the north have long-since washed away, leaving the north in particular extremely nutrient poor, whilst the short grass plains of the south are very, very rich. Particularly in phosphorus and calcium, both particularly important nutrients for pregnant and lactating wildebeest. The recent soils of the west are rich too, but mainly in Nitrogen, important, but not especially when pregnant. So here, immediately is a massive pull for animals away from those wet, but nutrient poor northern woodlands, down to the dry but nutrient rich grasslands of the south. Obviously they can only get here when it's wet, so timing their breeding to the rainy season on teh short grass plains is a great idea. What's more, predation down here is much lower too, as the hard pan and low rainfall prevents trees and lions have a much tougher time hunting away from the rivers and woodlands, which is great for baby animals.

Soak-away near Naabi showing the hard pan that limits tree growth, but makes grass very fertile

So, now we've got the important data we need for understanding the broad-scale movements of the migration. During the dry season, you've got to be near the Mara, in the far north. Once the rains come you want to move as fast as possible down to the nutrient rich grasslands of the south, where it's wise to give birth. But then once the rains stop, the bad news is that even though the grass stays green for a while, the standing water at Masek and Ndutu is so rich in nutrients that it's actually toxic - so even though the food is still there and still good you've got to start moving off as soon as the rain stops. But instead of heading straight back up the the north, it makes sense to move west, where there's still relatively rich grazing and water remains in the Grumeti and Mbalageti rivers. So come late May the migration moves away from the short grass of the south and heads into the Western Corridor, staying as long as the grass remains before gradually filtering north again as the good grazing is eaten in the west. (That date has got later in recent years, as there's now a lot more grass left in the Grumeti Reserves, thanks to a policy of burning only after the migration has been through - which explains why those northern camps have had some tough starts to the season in recent years!)

Movements of individual wildebeest caught near Seronera (blue circle) from here

And so you have the broad pattern - a triangular migration in a clockwise direction, covering between 500 and 1000kms, and one of the most amazing wildlife sights anywhere on earth. But, as always, the broad scale picture isn't all there is to it. Individual animals take some remarkably different routes around the ecosystem, as some data from gps collared indivudals shows - all these animals were caught near Seronera at the same time, but all have done different things - the dark blue one is particularly interesting, and none of these animals came down the eastern side of the NP at all. Why not? No-one knows - maybe simply because they were all passing Seronera instead. More recent work in the Masai Mara has made even more exciting discoveries, with animals I'd have assumed previously to be local migrants into and out of the Mara showing some extraordinary movements, even joining the main Serengeti migration in some years, but not others - look at these maps from here (they're updated very regularly, as the animals are still out there!)

The first of these spent a year in Kenya, migrating from wet season home in the west to the east and back, but then joined the main Serengeti migration this year and is somewhere in the NCAA today, whilst the other left Kenya last year and headed off to Loliondo for the wet season, before returning this year to wet season home in the north east! What made these animals change their routes from one year to the next? It will be fascinating to try and find out as more data on the movements of individual animals become available. Clearly, understanding the broad scale pattern is only a tiny fraction of the question as a whole and we've lots more to learn.

Anyway, I hope that's a pretty good introduction to some of the Serengeti Story. It's far from static, and there's still lots more to learn, so we're bound to return to the issue in subsequent posts, but I hope this is a good start at least. Meantime, Happy Christmas!

The Serengeti Story, part 1: history

So I guess this is the post I've been putting off longest. Not because it's not interesting, but because I know I'm going to forget some crucial component. But I'm just back again from a fantastic trip (thanks to all the guys at Dunia!) and decided it's definitely time to bite the bullet. However, it's going to be a long story, and I'm going to split it in two sections so I don't spend all night here (and so I stand a chance of remembering what I've forgotten before I consider the story told!). If you want more details on any of these things the essential references are the excellent series of very technical books edited by Tony Sinclair and colleagues you can get from Amazon. I've cut and pasted a few of the graphs from 'Serengeti III' into this post, hopefully 'fair use' for education...

I always start telling the Serengeti story with a bit of history, since it helps us understand how scientists have uncovered some of these things. There's no really obvious beginning to the story, but let's start with something we've already discussed on Safari Ecology - the introduction of Rinderpest to Africa in 1887. As we saw in that post, this had a massive impact on wildlife throughout Africa, the disease reaching Cape Town by 1897. The Serengeti migration was decimated, and when it was finally erradicated from the wildebeest population in 1963, there were still only around 250,000 wildebeest (see the plot below).
As you can see, once rinderpest was erradicated the wildebeest population exploded, reaching it's current total of somewhere betwen 1.2 and 1.4 million in about 1977, and this is the huge change that has let us understand so much of what happens in Serengeti.

Now, by now we should all know the 'Big 4' of savannah ecology, so it shouldn't come as a surprise that such a huge change in herbivory had a massive impact on the ecology of Serengeti, perhaps most obviously on the amount of another of the big 4 - fire. The figure below shows very clearly how the rise in numbers of wildebeest reduced the amount of fire in those northern woodland areas (essentially the woods from Seronera north).

This is clearly down to the very simple fact that wildebeest eat grass and grass is what carries fire through the savannah - more wildebeest means less grass which means less fire. And a change in the fire regime, of course, will alter the ecology too. So introduction around 1890 and then erradication of rinderpest in 1963 led to a massive change in both grazing pressure and fire frequency. It's not surprising, therefore, that massive changes occurred in Serengeti during the 1900s, most obviously the change in woodland cover. If you dig through old photos of the Serengeti / Mara area you can find some fantastic images of change. Tony Sinclair did it and came up with this beauty from 1944, that he then returned to in 1983 and took the subsequent photo (I've borrowed them from his talk available online here).

 It's pretty obvious that the woodlands vanished sometime between these two photos were taken and more detailed work suggested a rapid decline in woodland cover from about 1945 to 1980 - just the sort of delay you might expect from the increase in fire around the turn of the 1900th Century, given that fire doesn't kill savannah trees above 2m tall, so any established trees would gradually die of old age some time later.

Interestingly, as a direct consequenc of the decline in trees the national park authorities changed their fire management strategy in the 1970s from late burns at the end of the dry season and in anticipation of the rains, to one of early burns which tend to be cooler and rather less damaging to tree seedlings. At the same time, of course, the wildebeest population was recovering and the fire was declining in frequency as a consequence, so this change was probably less necessary than it seemed at the time (though everyone at TANAPA has since forgotten that the current fire strategy is a relatively new one, of course!). And as you might expect, more recently the trees have returned. Again, Tony Sinclair has some fantastic series of photos of these changes too, this from relatively close to Seronera:

(There's a whole lot more of these sorts of photos available on the web if you search for Tony's various talks.) And so the woodlands returned to Serengeti, as a consequence of the return of wildebeest and subsequent decline of fire. [It's interesting too, that savannahs globally are getting woodier, so there's a chance that this change is also related to global change too, not simply a local Serengeti effect - we might return to this in the future...]

But the story's not quite complete yet, as there's a neat twist at the end involving elephants. During  the 1970s and 1980s there was massive and nearly uncontrolled poaching of elephants throughout Serengeti, ending abruptly with the band on ivory trading in 1989. It's had a massive impact on elephant numbers in Serengeti:

At the same time, however, across the border in Kenya poaching remained under tight control, with no such dramatic change in elephant numbers. Such large herbivores can have a massive impact on the vegetation and the story in Serengeti is a particularly interesting one - Elephants walking across grassy plains often 'weed' out the tree seedlings instead of eathing grass. In woodlands they tend to leave the seedlings and concentrate on adult trees. So if there are lots of elephants it can be rather hard to turn grasslands into woodlands, even if the fire frequency is reduced. The difference between Kenya, where elephant numbers remained high throughout the period, and Tanzania, where they crashed at just the same time the fires declined, is stark. And elephants being rather clever animals, they knew where the border was and they were safe. So here's one last picture of Tony's from northern Serengeti / Mara, where the international border is clearly defined by woodlands.

Amazing to see the impacts of elephants so clearly, but also amazing to see how two different habitats (grassland and woodland) within the savannah biome can be stable under exactly the same environmental conditions - these days elephants are common both sides of the border and yet the woodlands remain in Tanzania, thanks to the different way elephants behave in grasslands from woodlands. So the history lesson ends with an important lesson about how important the initial conditions are to how a savannah looks - to turn a grassland to a woodland you need to reduce fire frequency (which can be done by increasing herbivory), but you also need to at least temporarily exclude elephants. All very complicated...

So, that's the history lesson and the broad overview of some population changes as a whole. The next post will continue the Serengeti Story by, I hope, explaining what we know about the migration and the regional differences across the ecosystem today. Hopefully it won't take so long to create either!

TAWIRI Conference discussions

I've spent most of this week at the Tanzania Wildlife Research Institute (TAWIRI) conference here in Arusha. This is an event that happens every two years and involves a very high proportion of researchers active across Tanzania, so it's always a good place to hear about interesting things going on in these areas. I thought I'd give a few of my highlights today. The two talks that most exicted me were from two different aspects of ecology - one by Dr. Grant Hopcraft on the Serengeti and how climate change might impact wildlife there, the other also related to Serengeti, but this time by Dr. Dennis Rentsch from Frankfurt Zoological Society on the economics of the bushmeat industry. I know both of these folk fairly well, so was able to press them for lots of extra information about both talks, and what I'm going to descibe here represents both their presentations and some of the other stuff we talked about - I hope they don't mind me putting this information out before it's all polished and published!

Wildebeest and zebra migrating through Grumeti Reserves, Feb 2010

Grant knows rather a lot about Serengeti and, in particular, the herbivores of the system. His work has focussed on how nutrition impacts herbivores and his talk fitted well into the overall theme of the conference on climate change, by asking how climate change will affect the nutrient content of the grasses and how this might impact the animals that feed on them. You might think it's crazy to suggest that climate change impacts grass quality (i.e. nutrient content), but actually it can have some pretty profound impacts indeed. Grass growing in high rainfall areas gets very tall very quickly, but also tends to be poor in nutrients - it might be that the grass can only collect the same amount of nutrient from it's roots, but in wet years it grows faster, so there's less nutrient per leaf than in dry years when the plants can't grow as much and pack all the nurients into a smaller volume. So more rain means lower quality grass, but more of it, less rain would mean less, but higher quality grass. In fact, lots of people showed plots of rainfall in Serengeti and demonstrated that the area is getting wetter (though I also suspect there might be shifts in the dry season length which could be even more significant, but no-one really talked about that), so we should be seeing more, lower quality grass. What is the consequence of this? Well, according to Grant, perhaps it means different things for different species, since all the herbivores prefer slightly different combinations of nutrient quality and grass quantity. In particular, hind-gut fermenters like zebra are happy with lots of relatively low quality food, whilst wildebeest are typically selective ruminants and need higher quality grass. Now, Wildebeest in Serengeti are food limited, not predation limited or anything else, so a decline in food quality might be bad for them - but they are, of course, interested in quantity too, particularly during the dry season when any rain is going to provide grazing which is clearly better than no rain at all. So a wetter Serengeti, if it impacts the dry season too, is probably going to mean more food at this crucial dry-season food shortage period, and we can expect that even in a wetter dry season the rain will still be scarce, so the grass will be relatively nutritious. So on the one hand poorer-quality forage during the wet season might be bad news, but more grass in the dry season is certainly going to be good news - which effect wins out isn't yet clear. My money will be on the dry season effects, but we'll wait to see! On the other hand, it seems pretty unambiguously clear that a wetter Serengeti will be good news for zebra, provided again that the dry season remains at least a bit wet too. So more zebra will always be good - though how that will affect everything else is also tricky to forsee. Does more zebra mean better facilitation for the wildebeest? Or might there be more competition? Who knows, as usual, more research needed (and if you want to fund Grant on his next project, do let him know - he's searching for money right now!).

The migration reaches Seronera, Nov 2010. Don't get eaten!

Spot the snare: many animals are poached in Serengeti. Moru Jan 2011

Meanwhile Dennis has been working on bushmeat trade on the western side of Serengeti for many years now. His approach to studying what is, after all, an illegal activity has been to deal not with the hard end in the park of finding and apprehending poachers and trying to get them to tell him how many animals they hunt (they're very unlikely to give an honest answer in such circumstances!). Instead he's focussed mainly on trying to work out how much bushmeat is being consumed in the villages around the Serengeti by asking them about the various protein sources they eat during the week. Although there might still be some resistance to tell the absolute truth in this context, it's likely his numbers are underestimates of the full impact of the harvest (especially as it doesn't include any of the meat that gets exported from the region commercially). Underestimates they might be, but the numbers are still staggering. In the villages surveyed, the average number of meals of wildebeest eaten per family per week was 2.4. Obviously that goes up during the period when the wildebeest are migrating through the particular village, and down when they're far away, but 2.4 meals per week is the average for the villages immediately around Serengeti NP. And knowing the number of households in each village, plus the number of villages Dennis estimates that somewhere between 90,000 and 100,000 wildebeest are harvested (illegally) from Serengeti each year. To put that into context, it's equivelant to a harvest greater than the entire wildebeest population of South Africa each year!

At between 500 and 1500TSh / kg (depending on seasonal availablity), and assuming a conservative 100kg of meat per animal that gives a a total market value of $2.5 - $8.5 Million per year. Compare that to TANAPA income from Serengeti gate fees 10 years ago (the latest I can find online) at about $5.23 Million, and we're talking the same size economy. (Bear in mind that these TANAPA fees are used throughout the national park system to subsidise less well visited parks, so Serengeti NP actually has an opperating budget of only around $2Million per year.) That's a pretty remarkable figure on it's own, but Dennis went on to talk about how consumption is related to price of other meat in the area - if the price of beef goes up, more wildebeest is eaten. Which suggests that it might be possible to reduce the amount of wildebeest eaten, if you bring the price of beef down. Now unfortunately I wasn't quick enough to get all the figures off Dennis's slide to do the calculation here, but I think I'm right in saying that if you want, say to halve the wildebeest harvest, his figures suggest you need to bring the price of beef down by about 3 times as much - so 50% of 50% of 50%, which is an 87.5% reduction in price. That's probably going to be tricky to achieve, unless you fill Serengeti with cattle, which is hardly going to help! So you're rather stuck there. Instead, the only effective solution is to make the wildebeest more expensive - and Dennis suggested you can do that either by giving poachers alternative employment and dry up the supply of meat, or by even more strictly enforcing the regulations within the park. But bear in mind that this is a sustainable harvest - there's no impact of this level of poaching on the wildebeest population overall. The problem is the bycatch - people want to trap common wildebeest, but instead their snares catch resident game sometimes and have had a missive impact. So instead of strictly enforcing current regulations, perhaps TANAPA should be looking at ways to encourage sustainable use and minimse the negative off-take. Perhaps making a few million $$ in the process. What do you think? Should we go this way? Or how should we feed these people?

The landscape of fear

Serengeti Landscape of Fear - where would you feed? On the green by the river where predators hide? In the bare bits on the plain with no grass left but a good view? Or risk the woodlands somewhere in between?

Lions are often in thickets (N. Serengeti)

But sometimes on kopjes... S. Serengeti

where you might also find a cheetah! N. Serengeti

It might sound like the sort of novel you's find abandoned at a camp by passing visitors, but understanding what ecologists mean by the 'landscape of fear' - how predators have impacts on the ecology of a savannah that go well beyond their direct predation events - is such an important concept that I'm going to break my usual rule of not talking about the big five! Actually, understanding the concept is simple - it's about looking at a landscape and working out where you'd be (most) scared to be walking. Long grass? Yup, scary. Thick riverine vegetation? Not for me! Nich bushy kopjie? I'll give it a miss, thanks. You get the idea - any place you might think about looking for predators whilst on a game drive, is going to be a scary place for herbivores too. And it's not simply a function of the numbers of predators that are present, but how efficiently they might be able to hunt within that habitat - I'd be much happier walking a short grass plain with a high density of lions than I would walking through some tangled thickets with a much lower density of lions. (Obviously I'm also a bit warier of buffalo and elephant than most herbivores have to be, but if you've done a few walking safaris you'll have the idea anyway.)

So what? These patterns are so obvious, we don't really think about them, or think they have an important part to play in very much - but we'd be wrong. In places where top predators have been removed, we rapidly see changes in the behaviour of herbivores and, soon after, we'll see changes in vegetation. Perhaps nowhere more famously than in Yellowstone National Park in the US (described here) - when wolves were eliminated elk and bison were released from their major predator and the populations changed - they didn't change in numbers very much, becausee like Serengeti's wildebeest and zebras (and, of coruse, elephants and the rest of the mega-herbivore group) they're limited by bottom-up processes of food availability, not top-down processes like predation. But they changed in behaviour, spending much less time looking around for predators and not moving around very far from their favoured willow patches. Which mean that after 50 years of no wolf predation, those patches of willows were in a bad way - it looked possible this form of riverine vegetation would vanish forever. Until 1994, when wolves were reintroduced. Within a matter of months the female elk and bison were spending significanty more time looking around, avoided open areas and only stayed in one place for a little period before moving on. And, in time, the riverine areas started to regenerate. We'd witnessed a 'trophic cascade' - removal of a top predator had had a massive impact on vegetation and landscape, the impacts 'cascading' down from top predator through the herbivore to the basal layer.

Leopards like riverine too, C. Serengeti

And lions often hunt by rivers and small ridges, Tarangire

These Fringe-eared Oryx have spotted something from their vantage in the Tarangie plains

The same processes are in operation on our east african savannahs all the time. Lions are far, far more efficient predators in areas where they can conceal themselves (in bushes, around kopjies, along even small river lines and shelves) than in the open - and as we know they plan their hunts accordingly. Similarly, leopards are best looked for around kopjies and riverine forests, where they're both able to avoid lions and can hunt sucessfully. Even cheetahs are often around kopjies to get a good view. So these are scary places for animals and, as we've seen so often before, if you start altering grazing pressure (one of the big 4 processes in the savannah), you'll see a change in the vegetation - riverine forests get a headstart if there's a big predator population living in them, scaring all the wildlife out! There's even the possibility that this processes becomes a positive feedback - as bush increases, so does predation success, making bushes even scarier, leaving fewer herbivores to keep the bushes back, meaning the bushes spread further, etc. And on the other hand, grazers like to graze areas where they have a good view - short grass - but if there are enough of them, their very grazing ensures the grass stays short, reinforcing the benefits.

Of course, things are complex for a herbivore - you can't simply decide never to forage in a wooded area because you might get eaten, because maybe half-way through the dry season you'll have eaten all the grass on the plains, and all that's left is in those scary woods. So you can either starve to a certain death in the plains, or head into the woods and risk predation, but at least stand a chance of avoiding starvation. Animals must constantly be assessing and weighing up the costs and benefits of foraging in high reward (grass under legumes like Vachellia is often of higher nutrient content than elsewhere) but risky areas, versus the safer but less beneficial areas on the plains. Not only will seasons make these decisions change, but so too will the details vary during the day - it's far more important to be in the plains at night than it is during the day, resulting in a evening movement of animals out of woods and onto plains - woodland edges are a great place to be at sunset!

Tarangire Wildebeest treck from the woodlands to the plains every evening

Spotting predators on Serengeti's short-grass plains is easy - no fear here!

Anyway, once you've got the idea of trophic cascades and the landscape of fear, you'll start seeing how it works all the time, and I hope I've given you enough here to start thinking about at least. (You'll get a much more in-depth and very readable discussion of tropic cascades and the landscape of fear in this nice article here.)

Rift valley geology and soils

Many visitors to East Africa are looking forward to seeing the rift valley, but often aren't quite sure what they're seeing when they get there, especially here in Tanzania, where it's a remarkably complicated feature and doesn't show the typical east and west escarpments of a rift valley - only some obvious western edges. This means, of course, that here in Tanzania it's impossible to point out exactly when you enter the edge of the rift valley, which is a bit confusing and disappointing to some travelling from Arusha to, say, Manyara for the first time - you're definitely in the rift valley at Manyara, and the western escarpment is obvious - but when did you actually arrive there?! Now, I'm not a geologist and am not going to go into huge detail about the rift's formation here, just the general idea should do. But I am an ecologist, and the presence of the rift valley has huge consequences for the ecology of East Africa too, so I might go into more detail about that!

Rift valley scarpment above Lake Manyara

Firstly, what is the rift valley? Well, some very readable details are available here, if you want the full thing. In summary, it's a great series of cracks in the earth's crust that can be traced right from eastern Turkey, through the Middle East and down trhough Ethiopia, Kenya, Tanzania, Uganda, Rwanda, Burundi, etc., as far south as Mozambique. Here in East Africa there are two parts to it - the western, or Albertine Rift, than runs through Uganda, Rwanda and Burundi back to Tanzania, and the eastern rift, running through western Kenya and the middle of Tanzania. These cracks are around 20-30 million years old (oldest in Ethiopia) and are believed to form because there's a large plume or two of magma (molten rock) beneath the earth's crust that pushes up on the crust, creating large bulges (the Ethipian Highlands, and the Kenyan/Tanzanian higlands are both pushed up from below) and, in places on top of the bulge, cracking the crust and leaving a rift valley. Some times, of course, the magma has burst through as volcanos, with erruptions still fairly regular in various locations. The older volcanos associated with this (such as Monduli, and the Crater Highlands) date from 20-30 million years ago, though the major faults (big escarpments) are only about 2 million years old. It's a divergant fault, splitting the African contient in two and gradully moving even now - eventually it seems likely that this fault will split Africa in two - but I don't think we'll be seeing that for the next few years at least...

Oldonyo Lengai is spectacular from the air!

So, that's (very briefly) what it is. The important things from an ecological persepective are it's incredibly recent geological age - compared to most of Africa, these mountains and plains are new - even the oldest are only 20-30 million years old, and volcanic activity has probably been pretty much constant since then. Now, this age is important, because (as a first approximation), material recently thrown out of the earth is full of unusual chemicals that, over several million years, will be washed away. Consequently, soils derived from new rocks are usually nutrient rich, whilst older soils derived from older rocks have been washed clean (leached) and are generally rather nutrient poor. And as we know, nutrient availability is one of the big four processes driving savannah ecology. I couldn't find any very fine-scale pictures of this, but I've found a global map of nutrients available to plants in soils here that I've included below.

The important thing to note is that in general, soils in Africa are incredibly nutrient poor (yellow and orange on the map), but that there's a clear green bit associated with the rift valley. That's the consequence of volcanic activity in this part of the world. This large scale doesn't show too much, but focussing in on the underling rocks will give us some idea about nutrients too - so here's a map I've edited from here that shows the geology of East Africa.

In this map you can see the fault lines creating the escarpments nicely, but you can also see how the only areas with relatively recent rocks are those associated with the rift, from northern Tanzania up through Kenya, except for the Tana river area, where the recent rocks have other origins. Note especially that Serengeti/Mara only has recent bedrocks in the southern, short grass plains, and Tarangire only just gets into that new complex right in the northern tip. In both places these nutrient rich soils explain in part the migrations we see, with calving always happening on nutrient rich grasslands. It's also obvious why wildlife densities in the rift valley are so much higher than elsewhere in Tanzania - and indeed Africa. The scarcity of nutrient rich soils, and hence food availability, probably limits animal populations in many of these other areas, right down to South Africa. No doubt we'll come to this in more detail in the future, but for now, what's probably enough - as well as being a spectacular geological feature, the nutrient rich grasslands associated with the volcanic activity help explain both the number of animals up here, and their seasonal migrations. Geology matters!

Why Are There So Many Wildebeest Compared to Other Animals in The Serengeti?

Herds crossing into Kenya.

Having been on safari for the last couple months, I’m unworthy of being called a co-author of this blog considering the wonderful posts that Colin has been writing. In my travels I have been to the Serengeti ecosystem four times in the last few months, three times in Serengeti and once in Maasai Mara and of course we have followed the spectacular herds of wildebeest.

When you’re driving through hundreds of thousands of wildebeest, or watching tens of thousands plunge into the Mara river because the grass is greener on the other side, its hard to wonder why there are so many of them. Why not zebra, topi, kongoni, impala, dikdik or one of the other antelopes?

So, I thought I would explore this topic and discovered this wonderful paper online, which you can download if you want to read a more scientific explanation. (Click here )

Part of Colin’s themes has been that there are things that shape or influence the environment, and that the environment then shapes the species in it. It’s a two-way interaction that steers what happens. E.g. When there is predation on plants they evolve defense mechanisms like thorns or chemicals.

So, what is it about the Serengeti that promotes these massive herds of wildebeest?

The simple answer:

Climate and soils.

The Serengeti ecosystem extends between two geologically significant features:

In the east, are the rift valley volcanoes that blew volcanic ash over the eastern part of the Serengeti, starting millions of years ago. These became the extremely fertile short grass plains between Maswa and Piyaya.

The short grass plains of Piyaya- the volcanoes in the distance.

In the west, Lake Victoria gives the north-western Serengeti a much higher rainfall (1200mm) than south-eastern Serengeti (500mm), especially when everywhere else is dry.  

Put these two factors together and you have high quality grazing every month of the year. In the wet months of the year (Feb, March, April), the soils in the short grass plains make the grass particularly excellent grazing with extra dose of calcium and phosphorous - perfect if you are a wildebeest trying to make milk for your calf. In the dry season- well, you migrate to where its raining and you find green grass which is much more nutritious than dry grass. (Wildebeest need 30% more energy, 5 times as much calcium, 3 times more phosphorous and 2 times as much sodium when they are lactating than pregnant and the short grass plains are perfect.)

A newborn wildebeest in Piyaya. It stands within 20 minutes
 to suckle. The milk is a high-cost to the mother but she
survives because of the minerals in the grass.

So, now we understand that the whole 25,000km2 Serengeti ecosystem always has nutritious grass (and drinking water) somewhere at all times of the year. The next question we have to investigate is- why wildebeest? Why not zebra, topi, kongoni, eland etc. etc?

The simple answer:

                   Wildebeest are special.

As you might know, wildebeest belong to a tribe of antelopes called the Alcelaphines. This means they are fairly closely related and if you want to know how close, well, they are about 4 million–year-old cousins. All of them are ruminants, which means they have a four-chambered stomach that they use to digest cellulose. Rumination is a very efficient way of extracting nutrients from plants but each species will have it’s own efficiency and Coke’s hartebeest are actually the most efficient of the three species. So why isn’t it Coke’s hartebeest?

Topi in the long less nutritious grass on the Lamai wedge

We can start by looking at the mouth structure of these animals and realizing that wildebeest actually have a mouth that is perfect for eating grass that is 3cm high, which is when the grass has the highest levels of protein.

The next thing they do is chose the parts of the grass that are also more nutritious- the leaves and fresh shoots. Coke’s hartebeest and topi eat more stems and leaf sheaths than wildebeest, zebra survive on almost only stems. But there’s a lot more grass stems than grass leaves so you would rather expect zebra populations to be in the millions but they aren’t- what is actually happening, is that zebras suffer very high losses of young, so predators keep zebra numbers down.

Now, you might ask, why aren’t wildebeest populations kept low by predators?

Answer: Synchronized reproduction and rumination.

80% of wildebeest calves are born in 3 weeks in February= 250,000 wildebeest calves= 500 per hour. It is an amazing sight. In scientific terms: extreme synchronous breeding outstrips predator’s ability to limit wildebeest recruitment.

Calves are most vulnerable when they are very young but they reach a certain age when they become equally vulnerable as the other wildebeest. There is a limit to how many calves predators can take per day, so by all having their babies at the same time, more calves have the chance to live past the age where they are vulnerable. Topi and hartebeest do not have as synchronized breeding as wildebeest.

Zebra on the extra nutritious short grass plains.

As we mentioned before, wildebeest are ruminants. They spend about 8hrs a day grazing so they have 16hrs a day to look for predators. Zebra on the other hand, spend 15hrs a day grazing so they only have 9hrs to look for predators. This is because they are hind-gut fermentators. This is obviously simplified.

Now, we’ve established the benefit of synchronized breeding but there are other advantages to being a wildebeest. Serengeti’s short grass plains are the best place for the females to get the nutrients they need to lactate, but they are also a great place to spot predators, which also helps to reduce the number of calves killed before they are out of the vulnerable stage.

Finally, calves are born precocial with a very strong imprinting instinct. The mother and calf learn to recognize each other immediately by smell and the calf stands as soon as it can and then stays as close to its mother as possible. The calf then also tends to run on the hidden side of the female so that predators have a harder time seeing them. The effect= reducing predation.

Wildebeest calve's coats change color to look like their
mothers at 2 months. Predation drops drastically.

There are other minor influences and for more details download the paper, but to try to sum it up in a sentence: The Serengeti’s unique climate and soils provide the perfect conditions to allow wildebeest to live in such large migratory herds because of wildebeest’s unique biology.

Rinderpest erradication

I'm often suprised when talking to guides about how few know of the great African rinderpest epidemic of the late 19th Century, as it has had a huge impact on the wildlife we see around us today. I'm prompted to write of it now for two reasons - the first being news I missed whilst busy training in Tarangire at the end of May and have only now noticed: on 25th May this year, the Food and Agriculture Organisation (FAO) announced the global erradication of rinderpest. This is surely something to be celebrated, as the first global erradication of any animal disease. And as we'll see in a moment, what a good disease to erradicate! The second reason is somewhat less pleasant, as I hear there's a measles outbreak here in Arusha at the moment. Why should this prompt me to write abour rinderpest? Well, evidence suggests that measles is actually a mutated form of the rinderpest virus that affects people. In fact, it seems most likely it evolved during the 11th or 12th century during one of the periodic rinderpest epidemics in Europe at that time, showing once again how nasty diseases have a habit of jumping across species boundaries when they get very common.

If you're wondering why I'm interested in cattle diseases at all, you've not heard about the African rinderpest epidemic, and of the impact of the disease and its control on wildlife. So let's start back in 1887, when the Italians were busy trying to colonise Ethiopia (afterall, the rest of Europe had colonies, why shouldn't they?). It seems as though some time early in the year they imported cattle from Asia that had the disease - whether as a deliberate act of biological warfare or accidently within their own supplies we're not certain (there's apparently no evidence to support the often heard claim of biological warfare), but whatever the reason it didn't take long for the disease to spread. Within they year up to 90% of Ethiopian cattle died, and at least 30% of the human population too (some estimates put it up to 60%). By 1897, after a short pause at the Zambesi and a couple of southern cattle fences, the disease had reached Cape Town, destroying 60-90% of Africa's cattle along the way. (You can read more about the human cost of this here if you want.)

Cattle aren't the only animals to suffer from rinderpest though - most ruminants were susceptible, some even more than cattle with up to 90% mortality of wildebeest and buffalo. Descriptions from travellers in Serengeti during 1898 suggest there were huge mortalities among the wildebeest, with the plains covered with carcasses. And for the next 60 years the numbers of animals in the Serengeti - Mara ecosystem were vastly reduced, held constantly in check by disease: in 1963 there were an estimated 250,000 wildebeest in the ecosystem (compared with 1.4million today) and immediately post disease outbreak possibly as few as a few tens of thousands of  individuals. The key change allowing the populations to recover was the erradication of rinderpest in the cattle around the park, following a large-scale vaccination programme. In fact, across most of British colonised East Africa at the time a plan was formed to erradicate wildlife from the area as a way of controlling rinderpest, but the colonial administration decided that whilst they could manage to shoot everything in most places Tsavo (in Kenya) and Serengeti/Mara were just too big, so around these areas they'd institute a 'cordon sanitaire' where cattle would forever have to be immunised, isolating the disease within the parks. This cordon was completed in the 1950s and by 1963 rinderpest died out of the wildlife population - the disease was so effective at killing wildlife that it couldn't actually be sustained within the population and had only be maintained by continual re-infection from cattle in the surrounding areas.

There are a lot of Wildebeest in Serengeti today - the view from Naabi hill, Jan 2011

Which led to a six-fold increase in wildebeest populations from 1963 to 1977 (and other increases in buffalo, etc), with a huge increase in the amount of grass being consumed by animals. (Read more about it here.) Of course, eating all the grass had a huge impact on how much fuel there was to carry fire in the ecosystem, and the frequency of fire started to decline in the 1970s. And as we know, fire has massive impacts in the savannah and it's decline is, perhaps, responsible for the return of acacia woodlands across much of Serengeti from the 1980s. (And so many visitors think of Serengeti as an ecosystem in the same state as it was thousands of years ago! Unchanging Africa is a myth...) But that's also a complicated story we'll deal with another time - for now, let's be glad there's no more rinderpest anywhere, let alone in Serengeti.