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Broadening the understanding of ecology in East African habitats, mainly savannahs

Colin Beale
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  • May 1, 2013
  • 04:43 PM
  • 56 views

Indian house crows and invasive aliens

by Colin Beale in Safari Ecology

Indian House Crow, not the prettiest... Thanks to Dick Daniels There are very few birds I don't like to see, but today's common bird is an exception - the Indian House Crow, Corvus splendens. Actually, that's probably slightly untrue, as I have been to India and I was perfectly happy to see the species there. In East Africa, however, this is not a species I'm ever happy to see. Not because there aren't interesting things to say about it, of course, but because it really belongs in India and seems to cause a number of problems elsewhere.First though, identification is fairly simple: the house crow is a medium-large all black and grey bird, usually found in flocks in towns all along the coast and, in some areas, invading inland too. It is very loud, with a persistent "Carr, Carr, Carr" call that is the constant sound of Dar es Salaam bird life... There are few confusion species in East Africa, the only other common species of crow being the black and white Pied Crow, which often hangs about with the house crow.So, what is interesting about house crows? Well, I guess the most basic fact is that they originally come from India and south-east Asia. They were actually first introduced to East Africa in the late 1890s, when they were apparently deliberately introduced to control rubbish on Zanzibar (note, that is certainly the oldest citation I've provided so far on this blog- hurrah for the British ornithologists' union digitising old issues of Ibis!). Which just goes to show quite how stupid we humans can be sometimes. Yes, they eat rubbish (lots of it too), but noone was thinking that they might also like to eat other things too- like all the eggs and chicks of small native birds, or even young chickens (meaning outdoor chicken farming is now impossible where they're very common), or young maize cobs, or, etc., etc. They also carry and possibly spread some rather nasty diseases around too. For a full list of impacts, I suggest going here.Thanks to J M GargNow, it turns out that house crows are an excellent example of an invasive alien species, and if you're not currently interested in alien species but are interested in conservation or even human well being you really ought to be! The EU estimates that invasive species currently cost the world economy $1.4 trillion each year' through their impacts on human health, through reducing the potential of land for agriculture (or making farming more costly), through altering the water cycle, etc., etc. and that's before we consider impacts on biodiversity, which is where most of the costs are associated with house crows.  Invasive species are a massive global concern for conservation, certainly in the top three causes of current extinction risk for threatened species globally.Given their massive impacts, you'll not be surprised to hear that there's been some research on invasive species, and it would seen that the house crow is a pretty perfect example. As we've noted already, the species was deliberately introduced to East Africa, but for a long time its population was fairly restricted, and only more recently have the numbers and distribution increased massively. This slow establishment, followed by eventual massive increase is absolutely typical of successful invaders.  Much of it can probably be explained by the fact that a population starts from just a very few individuals, and even exponential growth takes several generations before very large numbers suddenly seem to build up, but some of it might also be to do with the initial individuals not being perfectly adapted to local conditions, and evolving to local conditions takes several generations- but certainly can be rapid among some invasive populations.  Secondly, it turns out that it is extremely difficult to predict what species will become invasive- not all introduced species establish in the new area, not all established species grow much in population, and not all alien species that establish well actually have major impacts (it turns out that each stage has a roughly 10% chance). And remarkably little will tell you whether a newly arrived species will be invasive or not, the main key being whether or not it has been invase elsewhere. And again, the Indian House Crow is a particular good example: in recent years it has managed to reach nearly everywhere, and has established populations in 4 new continents!So now we know that the house crow can become invasive, it is interesting to know if we can predict the areas where it could possibly live, assuming that eventually it will find it's way there- as those involved in the current efforts to eradicate the species from Tanzania know, once established, it is extremely difficult to remove an invasive species, you're much better off acting before the population gets a chance to become established in the first place. Knowing where the species lives naturally, and the conditions it like to live in should give some useful information here, so recently a number of researchers have tried to build the same sorts of statistical models of distribution that we've discussed before here, and then plot where else in the world there are similar conditions, where the species is perhaps most likely to persist. Again, it's been done for house crows - and it's pretty impressive how far they might spead in the end - figure from the paper below!But as often with these models, I'm still not totally convinced - we know there's a population in the Netherlands, which only has a very low probability in the map above - but I would be totally astonished if it can live in the Netherlands and not through Spain and Portugal, for example. In fact, several of the studies that have compared the conditions in introduced parts of a species range with those in the native range find surprisingly little agreement, with introduced species often occupying different climatic regions than the area they evolved in, so I'm not totally convinced of the value of these exercises (probably partly because the models used to predict distributions are not always statistically or ecologically sensible).Still, all told, I hope you'll agree there is something interesting to say, even about the Indian House Crow!Main Reference:... Read more »

Duncan, R., Cassey, P., & Blackburn, T. (2009) Do climate envelope models transfer? A manipulative test using dung beetle introductions. Proceedings of the Royal Society B: Biological Sciences, 276(1661), 1449-1457. DOI: 10.1098/rspb.2008.1801  

  • April 10, 2013
  • 05:25 PM
  • 86 views

Common birds: Rattling cisticola and why birds hold territories?

by Colin Beale in Safari Ecology

Rattling Cisticola, near Arusha, March 2011. Something of a birder's bird? Returning to my recent theme of common birds, what could be more suitable as the rains begin than a look a Cisticola chiniana? Whilst small, streaked and brown might make this something of a 'birder's bird', I'm happy to think there's plenty to interest everyone in this species too.First the identification. Let's be honest, Cisticolas can be something of a challenge to identify! It doesn't help that there are seven pages of nearly identical looking small, streaky brown birds in the fieldguide! Happily, there are better ways to identify Cisticolas than their looks - the key is always to listen. Most Cisticolas, and rattling is no exception, have fairly distinctive calls and once you know it their 'tee, tee, churrurrurr' call is a constant sound in the bush (click the link to find a recording on xeno-canto), especially during the rains when they breed. In the unlucky event that none are making any noises, you can usually be fairly confident in your identification of any moderately sized, streaky cisticola present in the drier bush regions as rattling simply because they're so common! They are surprisingly variable in size (sometimes appearing really rather small) and colour (from very grey to warmer brown - but never with bright chestnut on the wing or head) though, so don't be too taken in my any one feature if they're not calling.This is a bright individual from Serengeti (thanks to tanzaniabirds.net)So, what is so interesting about this common bird? Well, the most interesting things I know about them concern their communal territories. We're so used to birds (and other animals, of course) holding and defending territories, that we often don't think to ask why they do it? Not all birds do, of course: our last common species, the red-billed quelea, is a colonial species that benefits from information sharing with others in the flock and never defends territories. Other species that we see in Tanzania (like willow warblers) show no territorial behaviour here, but are intensely territorial at other times of year. And most of the birds we think of as territorial usually defend a territory as a pair with one adult male and female (sometimes with sons and daughters from previous years sticking around to help out in cooperative breeding). Yet the rattling cisticola often holds communal territories involving two or three males and several females (and multiple nests within the same territory) where the males can be unrelated to each other. Watch carefully those males you see singing at the moment (and take the opportunity to make sure you learn the song, because it really is the easiest way to identify them!) and you'll soon spot that some are working together to defend territories against others. So why is this? Why should rattling cisticolas (and possibly other cisticolas that you see in groups) choose to not only defend territories, but defend communal territories with unrelated males?Obviously, this problem needs breaking down into two parts: why defend a territory in the first place - all that singing and fighting is pretty time and energy consuming. And if defending a territory is a good thing, why let others into yours and defend together? The former problem has been studied for a long time, and the answers are fairly obvious: territories are all about securing access to resources, be they food, females, nest sites or other scarce things. As such, there are two very important requirements that need to be met, before territoriality of one form or another can evolve: competition, and defendability. If some resource is extremely abundant, it's obvious that there's no use wasting effort fighting over it: red-billed quelea's grass seed food is so super-abundant when ripe, they have no need to fight each other for access, and as we saw before, there are real advantages to sharing information. If, however, you eat insects that are hard to find and that live in a rather harsh semi-arid environment, there's more incentive to fight over your spot - especially when you've worked hard to identify the best places to find those insects within you local patch. So competition is very important to the evolution of territoriality. Equally, there are some foods that are so spread out, that even if competition is very high, there's no feasible way for you to defend them. For example, if you're a seabird fishing over large areas of sea, you have to come all the way back to shore to nest and feed your chicks, meanwhile you obviously aren't able to defend your resources, so again there's no point in wasting energy trying to do so.Obviously, food availability varies throughout the year. Equally, a bird's requirement for food varies during the year- when feeding young, the amount of food an adult has to find is much greater than when only worrying about it's own needs. Consequently, the impact of competition is likely to change during the year, being greatest when trying to feed young. At some point, for some species, this might mean that the advantages of defending a territory may increase, and may outweigh the costs- leading to a pattern of seasonal territoriality like we see in the willow warbler. Of course, birds also have some options available to them to reduce the costs associated with territory defence - it's hardly efficient to constantly fight your neighbour over resources, so neighbours may 'agree' to leave each other alone, or simply identify accurate indications of how likely they are to win fights, without having to it all the time. Which, as I discussed on the post on why birds sing,  is one of the functions of song. So, these ideas work well to explain why birds might defend territories in the first place, but why should the Rattling Cisticola form communal territories?A much duller individual from Tarangire (Daudi Peterson)A number of ideas have been suggested to explain this behaviour. Firstly, there's the idea that you may well already be familiar with from knowing something about lions:  that without friends helping to defend a territory, there's no way you'll be able to do so on your own. Equally, there's the rather interesting idea that when competition gets particularly tough, any 'agreement' that may normally exist is torn up. For example, it's all very well agreeing not to constantly fight your neighbour, if you know that there will be a suitable territory for you somewhere else - but if all the territories are full, and you have no chance of breeding, nor much of surviving without one, you'll keep fighting even much bigger neigh... Read more »

  • April 3, 2013
  • 05:25 PM
  • 107 views

How to protect lions?

by Colin Beale in Safari Ecology

Lions: just big kitties really! There have been a couple of lion stories in the news in the last week or two, and enough interest in them that I felt compelled to write something. First there was a paper by Craig Packer and many coauthors about lion populations in Africa, their current declines, and the possible role of fencing in protecting them. Then, shortly after, there was a letter in the New York Times by Tanzania's own Director of Wildlife, asking the US government not to list the lion as endangered, as lion hunting is crucial to their conservation in Tanzania.Tarangire Lioness - a surprisingly resilient population.So, I thought it would be useful to bring the two issues together and look at some of the facts behind them. So, let's start by looking at Craig's paper on lion populations. As several others have already noted, this is an important paper - it has collected as many medium and long term population data as could be found, from lion populations all across Africa and received a lot of attention (click the links before and here!). in some places like Serengeti (where Craig, his coworkers and his predecessors have been busy recording lions annual for a very long time) the data are good. In other places, like Waza NP in Cameroon, the data are much patchier, but by collecting everything in one place we can see for the first time quite how much the population has been suffering in recent years. And there's a clear picture painted of large-scale, continent-wide declines. This isn't really anything new (though sad to see it so plainly), and will come as no surprise to anyone who knows anything about African conservation.Lions are still outside protected areas in much of Tanzania!Where the story gets more interesting is in the analysis of the population trends. As well as population trends, the authors have estimated how many lions there should be in a protected area, based on the number of herbivores and (where that's not available) the rainfall and nutrient availability. They can therefore compare observed lion densities with expected densities and identify populations that are below capacity, even if the trends aren't changing much. And alongside each of the population trends the authors have gathered a set of additional information for each protected area: human density in the surrounding area, an index of official corruption, operational budget, whether or not there is commercial hunting, area protected and other details of the protected area management, including whether or not the protected area is fenced. Using these pieces of information it is fairly straightforward to identify correlations between population trend and conservation actions. This analysis identified a number of correlations: both the degree to which lion densities matched potential lion density and the trend in lion population are positively correlated with whether the protected area is fenced, whilst the degree to which actual and potential lion populations matched was also correlated with the management and the trends correlated (negatively) with whether the park was run by the state (bad) or privately (good).Pale blue line is total lion density, dark blue is adult density, red is estimated maximum densityThis is potentially an important result. Regular readers will probably have noticed that I'm not generally a fan of fences, so data suggesting they're of major benefit needs some careful examination. I actually had a look at these data a while ago, as Craig was kind enough to send me an earlier draft of the paper, which enabled me to reach my own conclusions, of which I'll share a few here. Those who know me won't be surprised to know that the first thing I turned to were the data themselves. In the Supplementary Material there's a wealth of information (which is fantastic to see), including plots of the population densities in individual reserves. I've cut out the Serengeti information here so you can see (one of the best) examples. When I look at this, I can see that lion density as increased over time - some of that is due to the return of the woodlands, and the increase in herbivore populations - and that it appears to be pretty much at the theoretical maximum at the moment, but we can see fluctuations that relate to drought and disease. So it seems slightly surprising to me that the important table that summarises these results (Table S1) suggests populations are only at 85% of capacity and that the trend is a slight negative one. (The reason for this, of course, is that they only use the last 10 year of data, and the fact that the log scale is important is ignored when calculating the percentage) but what was obvious to me from this figure and, particularly the other, much more patchy datasets, is that any estimate of trend and/or population is only a rough estimate, with lots of annual variation and measurement error, but the variation in estimate is not considered in this analysis - it is (incorrectly and unnecessarily) assumed we have perfect knowledge of both population and trend, something I've often criticised. If this uncertainty is appropriately considered, I'm quite convinced there will be absolutely no significant correlation of any of the measured variables with population trend (though the correlation with population size might remain - but I'm equally sceptical here, because if there really are lion populations nearly 300% higher than the theoretical maximum density (as they appear to be from Table S1), then I'm sure the theoretical maximum density is incorrectly calculated, no matter how strong published correlations may be. This is a particular concern since fenced sites tend to have lower predicted lion densities than unfenced sites).This statistical issue aside (and the related one that assuming exponential growth when populations are manifestly not on the exponential part of a logistic growth curve is strange), there are other, more profound issues here too. For me, the crux of the matter is that these are correlations, and correlation alone can never identify causation. T... 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Packer, C., Loveridge, A., Canney, S., Caro, T., Garnett, S., Pfeifer, M., Zander, K., Swanson, A., MacNulty, D., Balme, G.... (2013) Conserving large carnivores: dollars and fence. Ecology Letters. DOI: 10.1111/ele.12091  

PACKER, C., BRINK, H., KISSUI, B., MALITI, H., KUSHNIR, H., & CARO, T. (2011) Effects of Trophy Hunting on Lion and Leopard Populations in Tanzania. Conservation Biology, 25(1), 142-153. DOI: 10.1111/j.1523-1739.2010.01576.x  

  • March 14, 2013
  • 07:39 PM
  • 179 views

Common birds: Red-billed Quelea, commonest bird in the world?

by Colin Beale in Safari Ecology

1000s of quelea at a dam on Manyara RanchI've been struggling to think of the next common bird to do something interesting with, until the obvious solution came to me, possibly the world's most abundant bird, the red-billed quelea.So, first the identification. The most obvious thing about red-billed queleas are, as the name suggests, a large red beak! Apart from that feature, females and non-breeding males are rather nondescript, small sparrow-like birds. Breeding males are rather brighter, with the red bill surrounded by a black face and variable amounts of orange on the top of the head and breast, with otherwise sparrow-like brown streaks on the back and wings. Perhaps the most useful identification feature though is the fact that you almost never see just one, but flocks of tens, hundreds or thousands of busy quelea all searching for grass seeds or drinking at waterholes.Red-billed Quelea on right in non-breeding plumage. What else do you see?!So, what are the most interesting things to say about red-billed quelea? I think the most interesting things about quelea are to do with their numbers - if you've a good enough internet connection, have a look at the clip here for a typically impressive flock. There are likely to be well over a billion pairs of these little birds: estimates vary greatly, but I tend to trust the 1.5 billion pairs quoted here as a reliable minimum estimate. (If you like back-of the envelope calculations, you'll be impressed to hear that at 20g each that's around 60,000 tons of red-billed quelea across Africa!) With nesting colonies reported in the millions, you can imagine the food that these birds need to consume: 4 million adults, feeding a further 8million young. Each bird needs around 3.3g of grain per day (a figure I'm sure is an underestimate - I also found 10g online, but doubt that too...), giving a daily demand of nearly 40 tons of grain per day. If they happen to nest near your crops, I'm afraid the impact could be disastrous. It's no surprise, therefore, that governments and farmers across Africa control these birds. There's a range of control mechanisms in use, from large-scale spraying with fenthion (which obviously kills large numbers of non-target species too, to say nothing about the risk to humans when they are inadvertently sprayed too) to the extremely spectacular (but rather less effective) fire-bombing of nesting colonies (I know there's a video of it online, but I can't find it now. Anyone help?). This massive impact alone surely makes these birds one of the most interesting things out there, but I find it interesting to think about why there are so many in the first place.Quelea always roost together at night - an chance to exchange gossip?The answers to this are quite interesting and can be tackled at a number of levels. Most simply, there's a lot of grass seed out there, and if you can eat human grass seed like sorghum and millet as well as wild grass seeds, you're certainly going to do very well. But finding grass seeds isn't always easy - rains can be patchy, meaning grass seeds ripen in patchy spots: one field might be good for a few days, then another field somewhere else. Finding this patchy and briefly-available type of food is pretty tricky. To solve this problem, red-billed quelea are a very good example of birds that appear to support the "Information Centre" theory. This theory suggests that the optimal strategy to find food under these conditions is to scatter into small groups, search about, and then come back together, share information about what you find, and then follow any other individuals that have found food. Quelea are pretty good at this - they roost together overnight in huge flocks, and also roost during the heat of the day in smaller groups, often visiting water holes. During these roosts, not only are they fairly well protected from predators (if you are one in a flock of 100s of 1000s, you're very unlikely to be the one that gets eaten this time) but they can exchange information with one another. Why should a bird that has found food share this information with other birds though, and not just sneak off to eat it's fill on it's own? Well, if the food supply once found is much more than one or a few birds can eat alone there's little cost to sharing this information. And with seeds, before long they fall off (or get harvested) and lost - so there's only a short period of time when ripe seeds are available, and when they are, there's far more than any individual bird can eat. So with little cost involved in sharing information, the benefit of sharing with your relatives (helping relatives is generally good for your genes too), or even with any other individual likely to share similar information with you in the future, is certainly going to be a good idea. It becomes much more likely to evolve when individuals are identifiable to each other (a sort of ;you scratch my back, I'll scratch yours)- and this is one of the reasons currently being investigated as to why male quelea are so obviously different in how red they are, not (as often the case) simply to attract females, but to make themselves identifiable when in a vast flock. There's lots more work to do in this area, to test the theory completely, but I think that there's some good evidence for quelea that it might really be an important reason why the species is always found in such large flocks.Another level to explain the large flocks at, is to look at their breeding biology. Quelea are really rather interesting as Africa birds, because they have relatively large clutches: four eggs are fairly common, though two and three are more usual. That's a somewhat higher than average clutch size for similarly sized African birds, where small clutches are the rule (see the earlier blog on African birds doing this slowly to understand why that is generally the case). And even more remarkable is the speed with which they breed. Once adults build up the correct protein reserves they start to nest: incubation is 9 or 10 days, fledging is at 16 days and independence a day or so later, start to finish in 5 weeks and the population has doubled! With more than one breeding event per season, possibly in distant locations, and first breeding at only 9 months old, the species has the potential to rapidly increase in numbers when conditions are good. There are lots of other things we could talk about weavers too, of course - most recent research on them has focused on ... 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Bayer R.D. (1982) how important are bird colonies as information centres?. The Auk, 99(1), 31-40. info:/

  • January 31, 2013
  • 04:19 PM
  • 141 views

Common Birds: the case of the Baglafecht Weaver and missing forests

by Colin Beale in Safari Ecology

Male Baglafecht Weaver, Mt Kilimanjaro  If you live on or near an East African mountain, you're very likely to have Baglafecht Weavers in your garden. Like most of the other true weavers, they're a basic black and yellow colour. The first thing to look at in weavers is usually the colour of the eyes and legs: in Baglafecht weavers you'll always see a yellow eye (easy to see against the surrounding black feathers) and pink legs. Males and females differ slightly: males in the population in northern Tanzania and Kenya have only a black mask on the face, with yellow on the top of the head right down to the (black) beak. Females have an all dark head. In northern Tanzania the back of both sexes is essentially black, with some yellow wing edges, in other areas of Tanzania the back is greenish/grey and not as strongly contrasting. Juveniles of all forms are rather greener and lacking in black, but still have the yellow eye. Like other weavers, they weave their nests from grasses in colonies of 5-15 pairs (not usually in very large groups) and males in the breeding season are pretty noisy with their rather scratchy and squeeky song! Baglafecht weaver nests aren't the neatest of affairs...Map thanks to the Copenhagen DatabaseOne obvious fact that I could have chosen to talk about would be the famous weaver nests themselves. However, I think I'll save that for a different species in time, since there's so much to talk about there and Baglafecht Weavers hardly have the tidiest woven nest of the family... Instead, the thing I think of whenever I see these birds is the remarkable insight they give us into how species evolve, and some African pre-history. Already above I've hinted that there are some identification challenges with this species, populations in the northern highlands being different from those in the south. And in fact the confusion is even greater than that: there are currently a total of eight subspecies described, many rather distinct. (See some of the variety of three Tanzanian subspecies in pictures on the TanzaniaBirds.net site here) In fact, Baglafecht weavers occur from Nigeria in the west, to Tanzania in the east, north to Ethiopia and south to Malawi. And yet, although it has a very wide distribution, within this area it is far from everywhere, restricted to the higher hills and mountains, some separated from one another by 1000s of kilometres (see map). Now, each of the hills and mountains contains is home to only one of the various races: Northern Tanzania has the race often called Richenow's Weaver, whilst southern Tanzania has Stuhlman's,and each other set of hills has it's own. And it's not just Baglafecht weavers that show this pattern - plenty of other birds other animals and plants also show similar patterns, present on many African mountains, but not in between with slightly different populations in each area. Yet it's pretty puzzling when you think about it: ultimately, all the individuals of one species share a common ancestor so at some point in time all these now isolated populations must have been continuous. But how can that be so, given that they're restricted to mountain forests?Female Baglfecht Weaver, ArushaThe answer, of course, takes us into the past, quite far into the past, in fact. Just 12,000 years ago northern parts of the world were emerging from an ice-age that had lasted around 10,000 years. Most places currently with temperate climates where then very much colder than today. In tropical Africa, however, the impact was slightly different: it may have been a bit cooler, but the same conditions that brought ice to the north, resulted in drying in Africa. Using pollen collected from deep cores sampled from the bottom of lakes, scientists can identify the plants that lived around those lakes at different times in the past: the deeper you are in the core, the older the sample. And although there are still lots of gaps where there aren't sufficient lakes or people simply haven't had time and money to look, we've now got a pretty good idea of what happened to Africa's biomes during that last glacial period, about 18,000 years ago. Crucially, we can see that the plants currently associated with Afromontane forests occurred at significantly lower altitudes than they currently do, suggesting any of the species around at that time would have had to move. And, of course, there has been not just one ice age, but several over the last million or so year, meaning animals and plants will have been repeatedly moving up and down mountains - a process that has been thought to explain the interesting patterns of distributions among the montane bird communities, at least since Moreau studied birds in Amani.However, recently advances in DNA sequencing means that we can analyse exactly how different each population is from another one, and there's a handy process called the 'molecular clock' that lets us actually determine to within a few 1000 years how long it has been since two populations were last in contact. (This 'clock' works because there are some parts of DNA that are not really subject to strong selection and accumulate mutations at a regular, and known, rate: roughly speaking a 1% difference in DNA corresponds to a 1 million year split.) Now, I don't think anyone has done this yet for Baglafecht weavers, but they have for other birds like starred forest robins and olive sunbirds. In these (and other) cases, it turns out that most of the populations diverged rather longer ago than we thought: one set of splits occurred around 1.7 million years ago, another around 1.3 million years, and a last just under a million years ago. These times correspond very nicely with periods when we think Africa was undergoing even more extreme drying that during the recent ice ages, suggesting that it's these extreme drying events that allowed the montane forest much lower than today, spreading in an arc that connected all the current fragments.So by looking at the current pattern of distribution in Baglafecht weavers, we can deduce that Africa once (not even that long ago in geological time!) looked very different than it does today, with low altitude montane forest across much of the central region. And we can realise that our current impacts on global climate are very likely to result in some equally dramatic changes in biomes across the world - who knows where the Baglafecht weaver may have to live in 50 years time!Main references:... Read more »

  • January 23, 2013
  • 04:13 PM
  • 181 views

Common birds: Ring-necked Dove

by Colin Beale in Safari Ecology

Ring-necked Dove (Cape Turtle Dove) in Tarangire, photo from here Two more challenges have been set since the last one, and I'm hoping to rise to each! The first was for the African Collared Dove, however that species (Streptopelia roseogrisea) from the drier north of Africa in the Sahel and in the Middle East is not found in Eastern or Southern Africa and I suspect request was for a Ring-necked Dove Streptopelia capicola which is indeed one of the commonest birds to be seen in the bush across much of Africa.The sound of the Ring-necked Dove  is one of the constant backgrounds to a safari in the bush (if you don't know it, the "work harder, drink lager" refrain is available here) and it's actually this distinctive song that is the easiest way to identify the species from among a number of confusingly similar species. The ring-necked dove is a medium sized, grey dove. It has a black collar around the back of its neck and is a paler grey white below, with pale edges to its tail. Unfortunately, that description is would cover just about any of the close relatives of this species, and (as well as listening to the calls) you need to look rather closer to identify the species correctly. Firstly, look at the eye: if it is dark and not obviously surrounded by bare skin, you're probably looking at a Ring-necked Dove. White (not grey) edges to the tail and a generally pale grey would confirm the identity in eastern and southern Africa. If the eye is pale yellowish, with a red ring around it and there's a warmer brownish wash to the back and neck that contrasts with a grey head, you're probably looking at an African Mourning Dove (call) and if its got a dark eye in a bare purple/red patch of skin, and is overall darker looking, with grey tail edges, you're looking at a Red-eyed Dove (call: "I am a Red-eyed Dove").Another borrowed picture, fora  grey bird, they are rather attractive!So, once the identification is over, what is there interesting to say about this species. Well, I think there are two things to point out generally about doves, one of which I've mentioned before on the blog, the fact they're one of the few bird groups that can drink without lifting their heads (see the post here for the full story!). The other interesting thing is their ability to feed their young chicks what is called 'pigeon milk' for the first few days after hatching. Essentially, adult doves of both sex produce a substance in their crop that is fed to young chicks in place of normal food. Although this substance has evolved completely independently from mammalian milk, it has a number of remarkable similarities which has been the subject of a couple of interesting recent papers, starting from the extraordinary fact that the process is all regulated by exactly the same hormone (prolactin). Whilst mammalian milk is a liquid secretion that is probably a modified form of sweat (nice, huh?), crop milk is actually made of specially grown cells that are shed from the lining of the crop (for a particularly gruesome picture of dissected 'lactating crops, have a look here...). That difference aside, the content is pretty similar: a very high component of protein (around 60%) and the rest of the solid matter being mostly fat. Amazingly, the similarities don't end there - an important function of mammalian milk (and probably the main reason why formula isn't as good for babies as breast milk) is to pass on antibodies to the baby, which help protect the baby as it's immune system develops. And it turns out that exactly the same things happen in crop milk - chickens fed on pigeon milk developed a stronger and better functioning immune system than those fed on a similarly nutritious diet, but without the antibodies. Moreover, mammalian milk also includes bacteria, which effectively 'seed' the guts with useful bacteria that help digestion - the same is true of pigeon milk. It seems that 'milk' has evolved multiple times with parallel results -  it isn't only pigeons that produce cop milk, flamingos and (male) penguins do too, but it is rather remarkable!These general facts about pigeons are certainly interesting, but not specific to the ring-necked dove. The most interesting thing I know about ring-necked doves in particular, is that in some areas of Uganda they are perhaps merging into a new, hybrid species. Often, two species that are closely related and usually live in separate regions illustrate an interesting process known as "Reproductive character displacement" if and when they later come back into contact with one another. This is the process by which two, usually closely related, species appear more different in the zone of overlap, than in areas where only one of the two species lives. Often closely related species can hybridise (male and female of two different species will mate and produce young, the hybrids), but these hybrids are usually not as fertile as 'pure' individuals, and may have other problems too. So it's in the interests of the two species to be able to accurately identify mates of their own species. Where all the grey doves in an area are of one species, you don't need to discriminate much to find a mate of the right species, but where you overlap with a closely related species, you need to be more careful about who you mate with, and evolution might tend to exaggerate whatever slight differences exist. If, for example, one species has a slightly higher pitched call than the other, where the two overlap this difference may be  much greater than elsewhere. If, however, there's little cost to hybridisation (which sometimes occurs if the hybrids themselves are able to mate with other hybrids), the opposite can occur and a new species with features of both can occur - it seems a relatively rare way of forming new species (except perhaps in plants), but might just be happening with these doves at the moment. All very interesting...So, hope that's something interesting about ring-necked doves. Baglafecht weavers are next... Main References:Gillespie, M., Stanley, D., Chen, H., Donald, J., Nicholas, K., Moore, R., & Crowley, T. (2012). Functional Similarities between Pigeon ‘Milk’ and Mammalian Milk: Induction of Immune Gene Expression and Modification of the Microbiota PLoS ONE, 7 (10) DOI: 10.1371/journal.pone.0048363... Read more »

  • January 6, 2013
  • 04:39 PM
  • 174 views

Common Bulbul and frugivorous birds

by Colin Beale in Safari Ecology

Common Bulbul nesting in ArushaThanks 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!Bulbuls are fairly generalist- they eat invertebrates very happily (often feasting early morning on moths around security lights) but probably their biggest impact is through their fruit eating. Now as is often the case I could find little specific to Tanzanian bulbuls, but the species is very widespread and is found in the Middle East as well as across Africa. There its role in seed dispersal has been studied, and it turns out to be rather important. Obviously, plants grow fruits to encourage birds and other animals to eat the fruit, swallow the seeds and then drop them elsewhere with a nice helping of dung to start them off. I'd always assumed that with the exception of some special fruits eaten by large animals like elephants, the bird dispersal was a nice optional extra for plants - without birds I assumed the seeds would germinate perfectly well, just rather closer to tteamsother tree than if a bird had carried the seeds away. Well, it turns out I was wrong, and there are two reasons why this is so. Firstly, it seems like the process of passing through a bird's digestive system is extremely important to many seeds, without which they barely germinate. In the Israeli study, for example, one of the trees studied has a 2% germination rate of fruit that hasn't been through a bird (i.e. 2 of 100 seeds germinated), but 50% of those that had passed through a bulbul germinated, a much more acceptable rate. The same was true for several other tree and bush species studied. Bulbul at Olduvai GorgeSecondly, it seems that many Tanzanian riverine forests need birds to eat their fruits to protect them from beetles. This work, carried out by Tony Sinclair's group in the Serengeti, studied what happened to seeds from riverine forest trees if they were eaten by birds, and if they weren't.The story is simple - if the seeds fell with the fruit still intact virtually all were predated by beetles. If, however, birds had eathen the fruit and the seeds fell uncoated, the beetles ignored the seeds and the trees germinated. So without fruit-eating birds in riverine forests there'd be no riverine forest. And one of the problems with a lot of riverine forest at the moment is that too frequent fire has removed much of the understory, leaving grasses that the baby trees strugly to compete with and removing the undergrowth that many fruit-eating birds nest in,a nd once the birds are gone, the forest follows. So frugivores like the Common Bulbul are an absolutely essential part of the riverine forest habitat too.Now, of course, I have to confess that for both of these important roles, it doesn't seem to matter too much what species of bird does the eating, so this story could have been told for many other fruit eating birds so you might suggest that I'm cheating by using it as an illustration of the Common Bulbul. But as I mentioned to start with, common species, by being abundant, have a very important role to play in many of these processes. Moreover, another study from Israel showed that Bulbuls are much more likely cross open gaps between trees, meaning they're most important at spreading fruit seeds to new locations, so I'd suggest it is a really good species to choose to illustrate this important process...Main reference:Sharam, G., Sinclair, A., & Turkington, R. (2009). Serengeti Birds Maintain Forests by Inhibiting Seed Predators Science, 325 (5936), 51-51 DOI: 10.1126/science.1173805... Read more »

  • September 2, 2012
  • 03:09 PM
  • 255 views

Cycads and more botanical revolutions I've missed...

by Colin Beale in Safari Ecology

Lake Natron Cycad, near Loliondo. Cycads look ratherpalm-like, but are not true flowing plants at all.Back in April I headed to Loliondo for a few days with a bunch of guides from Thompson Safaris. Along the long and bumpy route I was really pleased to spot some Cycads and jumped out to take a few photos. Spotted in action, I was forced to explain why I was taking pictures of some random tree. My answer at the time was based mainly on the evolutionary history of plants that I'd been taught at school and then probably on into university: Cycads form a remarkably early split from the branching evolutionary tree of seed-bearing plants, their ancestors somehow linking ferns to the much more modern flowering plants.  I was also keen to see this particular species (according to the IUCN commonly known as the Lake Natron Cycad) because it's one more of those remarkably restricted range species that fascinate me.Now I have a confession to make - once again I was completely wrong! It seems that there's been another revolution in the world of botany that I'd completely missed and the latest evidence suggests that the group of plants we call gymnosperms (which includes the Cycads, as well as other more familiar groups like the conifers) are not really steps on the way from the 'lower' plants like ferns to the flowering plants, but rather a whole side-branch in evolutionary history. I've spent a while trying to put the full story together, and my conclusion from all the various papers I've been reading, is that it's not yet clear exactly how the new tree of plant life fits together, but it is pretty certain that most of the gymnosperms form a separate group to the flowering plants, and that as a group they were the dominant vegetation at around the time land vertebrates first evolved and right up to form the forests roamed by dinosaurs. They then went through some major extinction events apparently caused by climate change, so that today this once dominant group of plants is far rarer across most of the world than the much newer flowering plants.So, whilst cycads aren't any longer considered some sort of botanical 'missing link', they are still a very interesting relic of a past landscape. And my reading discovered a few more fascinating things too. A few of the interesting things to remember - all Cycads come as either male or female plants and, obviously, only the females have the big cones that hold the seeds in this group. Those seeds, attractive as they may look, aren't all that good for you - in fact they are really rather toxic even after extensive preparation, and are blamed for a range of brain diseases in human cultures that eat them, such as in Guam. The leads are also rather toxic (full of cyanide in fact), but there's something odd going on with one of the other Tanzanian species that is common on the coast, where the Zanzibar Red Colobus seem to be very keen on eating the leaves apparently without ill effects. And the other thing that interested me is that most (possibly all?) Cycads have an association with nitrogen-fixing bacteria, just like the legumes. That's of interest to me because the habitat the Tanzanian species are found in are usually very low nutrient systems and I wonder if there's possibilities Cycads have some similar roles in their habitats to that of Acacia in the savannas.All in all there's plenty of interesting things to say about Cycads, and I'm pleased to have found this rather rare species. I was also rather concerned to hear from several people that there's been a massive recent export of these plants to gardens around Tanzania, by people who headed to Loliondo to visit Babu. There certainly didn't seem to be many plants around in the few suitable spots I looked in, and as it's already considered Near Threatened, my guess is that this is, sadly, yet one more species on it's way up the endangered status.Main reference:M.J.M. Christenhusz, J.L. Reveal, A. Farjon, M.F. Gardner, R.R. Mill & M.W. Chase (2011). A new classification and linear sequence of extant gymnosperms Phylotaxa, 19, 55-70... Read more »

M.J.M. Christenhusz, J.L. Reveal, A. Farjon, M.F. Gardner, R.R. Mill . (2011) A new classification and linear sequence of extant gymnosperms. Phylotaxa, 55-70. info:other/

  • August 7, 2012
  • 06:04 AM
  • 290 views

Do fires stop the Serengeti migration?

by Colin Beale in Safari Ecology

Dr Kate Parr lighting a controlled fire in the Serengeti Ecosystem There's been a bit in the East African press recently claiming that Tanzania has been deliberately setting fires in the Serengeti NP to block the migration. The Tanzanian National Park Authority (TANAPA) have, of course, denied this. Reading the articles and press releases there's obviously both some serious ignorance and some seriously bad journalism going on here, and I thought it might be useful to share a few of my observations.Each fire burns differently, so to study their impacts you must observe closelyFirstly, let's look at the fires themselves. It is actually easy to see where fires are burning and how large they are - almost real time data are plotted on the NASA website and with a little fiddling I managed to set the parameters for the week in question for everyone to see here. Each cell is 500m square, and a fire may have occurred anywhere within that box, but has to be of a certain size before it is detected. Where there are lots of overlapping cells you can be pretty sure there's a big fire happening across a large area. From this map you'll see (a) that it's hard to see any fires in northern Serengeti during the week in question that could come even slightly close to being big enough to block the migration and (b) that there has been some really big fires elsewhere in Serengeti during the period, particularly around Seronera. So the TANAPA statement that the fires in question were under half a km were not exactly accurate, but nor is there anything big enough to ever create an obstacle for animals even if fires could do that.Has this tagged tree seedling been killed by the fire, or will it resprout in the rains?What about that shoddy journalism then? Well, you only have to compare the press release from TANAPA with the headlines like "Tanzania refutes allegation of fire deliberately set in Serengeti" to see there's no doubt who set these fires at all, nor that they were deliberate. And,in my experience, if you talk to rangers in northern Serengeti about why they set fires you might well find (sad to say) that they state that they set fires to keep the animals in Tanzania. It's not what's in the fire management plan, but I've failed to find many rangers on the ground who can give any of the real reasons why and when they burn. Why do they think it will keep animals in Serengeti? Because fires should promote nutritious grass growth. Does it work? Certainly, for a while. Immediately after a fire, whilst it's still smoking, animals come on and lick the ash. A week or so later, there's a flush of new growth that keeps animals very happy for as long as it lasts - which depends on how many animals there are and how much moisture there is in the soil. Now, the question is, given that the wildebeest population in Serengeti is food-limited, is the reduction in poor quality grazing (by burning it off) compensated by the smaller volume of higher quality forage? And the truth is that we simply don't know because researchers have mostly been asking the wrong questions. Instead of studying a range of alternative management options, they have simply compared early burns with no burns. Does that mean early burns are better than late burns? Does that mean burns every year are better than burns every two years? Sadly, we simply don't know. But I do know that change in the fire regime within Grumeti reserves is responsible for the relatively later arrival of wildebeest in northern Serengeti for the past few years - Grumeti tries to prevent fires there until the migration has passed through, and the increased food availability has kept the animals in the protected area much longer than before.Highly technical beer cans can help tell you how hot the fire was![BTW Where the other random statements about the reasons for the migration that the TANAPA spokesperson has come up with - e.g. avoiding inbreeding - I really have no idea! Any suggestions, do let me know - what scientists tend to think about the drivers of the migration at the moment is summarised in this blog post... I do agree with them that few animals in Kenya by late July is hardly unusual - presumably wishful thinking on the behalf of tourism folk in the Maasai Mara!]Overall, this just makes me more confident than ever that what we really need is a good, controlled experiment on the impacts of different fire regimes across the Serengeti ecosystem, and I'm really glad we've got it started (that's one of the things I've been busy doing whilst I've no been active here for a few weeks! Sorry...)Main Reference:Shombe N. Hassan, Graciela M. Rusch, Håkan Hytteborn, Christina Skarpe, & Idris Kikula (2008). Effects of fire on sward structure and grazing in western Serengeti, Tanzania African Journal of Ecology, 46 (2), 174-185 DOI: 10.1111/j.1365-2028.2007.00831.x... Read more »

Shombe N. Hassan, Graciela M. Rusch, Håkan Hytteborn, Christina Skarpe, & Idris Kikula. (2008) Effects of fire on sward structure and grazing in western Serengeti, Tanzania. African Journal of Ecology, 46(2), 174-185. DOI: 10.1111/j.1365-2028.2007.00831.x  

  • June 14, 2012
  • 09:51 AM
  • 460 views

East African Butterfly families and corrupt, singing caterpillars

by Colin Beale in Safari Ecology

Citrus Swallowtail, Papilio demodocus, is very common in TanzaniaWe're rarely short of butterflies in Tanzania, but they're a sadly overlooked group. Except, perhaps, when they're swarming by the million as earlier this year most people will, at best, only notice a few in passing. For a hugely diverse group (there are over 18,000 described species), they fall into a relatively small number of readily recognisable families. Unfortunately, all the nice identification books are out of print (and wickedly expensive to buy on ebay!) for East Africa, but there are some resources out there that will help once you've figured out the families. The relationships between the families have recently been the subject of some serious work. It turns out that the family relationship were rather difficult to pin down because they all evolved relatively quickly in the Cretaceous (yes, dinosaur time, 100 - 75 Million Year Ago). But our best guess at the moment sorts them into 4 main groups split into a total of 26ish main family groups, only a few of which are at all diverse. So it's not too hard to get to grips with the main families, and the main change to the traditional taxonomy, if you've been into that, is that the big group Papilionoidea is actually two, rather distantly related groups. I'm going to describe some of the common families here (together with some of my favourite stories about them - yes, including corrupt, singing catterpillars) and hopefully will be able to show how the various families fit together at the end. So, here goes...Graphium antheus, the Larger Striped Swordtail. Note 6 legsPapilio orphidicephalus, the Emperor SwallowtailStarting at the bottom of the new taxonomy (the first major split from the butterfly clade) we've got the Papilionidae, the swallowtail family. We've got lots of flavours of these here, from the standard yellowish ones like the Citrus Swallowtail which is virtually everywhere, to the nice Graphium genus of greenish blue ones. They're typically big spcies, often with long tails, but it's the caterpillars that are most distinctive - often brightly cloured and striped, they have a special forked organ behind the head that distinguished the group from all other butterfly groups. Most interesting thing about this group? Check the mocker swallowtail story in the caption...Papilio dardanus, the Mocker Swallowtail. This is probably a male, but the females come in at least 14 different forms, as mimicsof toxic species, but also as mimics of the male. Mimics of toxic species escape predation, mimics of males escape sexual harrassment! They're fascinating things...Coeliades anchises, the One-pip Policeman (check the single dot on the white triangle!) is not a typical skipper, but still shows curved antennaePardaleodes incerta shows the strange resting wing position of skippersNext off is the Hesperiidae, or skipper family. If the swallowtails are the birds of paradise of the butterfly world, this family are the little brown jobs... They are a really big group and readily identified: all have curved antenae, instead of straight-ended clubs in all other families. They're usually small winged and big bodied, looking rather more like moths than other butterflies, and they're pretty unusual too in the way their wings are held, often folded in strange ways when resting. A hugely diverse family, many of these skippers are highly host-plant specific and often show strong evidence for co-evolution with their hosts.The Flat, Celaenorrhinus galenus often sits underneath leaves perches on the undersides of leaves and shows curved antenae tipsColotis evagore antigone, the Tiny Orange-tipRaffray's White, Belenois raffrayi is a typical whiteThen we're into the whites, or family Pieridae. As the common name suggests, this is the grou... Read more »

Heikkila, M., Kaila, L., Mutanen, M., Pena, C., & Wahlberg, N. (2011) Cretaceous origin and repeated tertiary diversification of the redefined butterflies. Proceedings of the Royal Society B: Biological Sciences, 279(1731), 1093-1099. DOI: 10.1098/rspb.2011.1430  

  • June 4, 2012
  • 02:07 PM
  • 335 views

More on management of protected areas: the human dimensions.

by Colin Beale in Safari Ecology

Public relations are a huge part of conservation workIn the previous post I described two of the ten lessons that we, a bunch of conservation managers and researchers from eastern and southern Africa identified at a workshop in Serengeti. I started with the big lessons on making sure you start with boundaries that make ecological sense - and what can happen particularly to migrations if that's not done. There's more to learn on that score too, but I'll skip to one of the most important lessons we identified, that will come as no surprise to anyone working in the field: don't neglect public relations!There are two groups of people that we identified as really important here - the general public, and the local population around protected areas. In many areas, poor relations with neighbours can be traced right back to the exclusion of people from their traditional grazing areas during protected area formation. This comes back to a point we've made before on the blog - that people were once part of these landscapes and the lack of current habitation is an artificial imposition of what is mainly a colonial-era romantic ideal of empty Africa. No wonder local people often feel hostile to the park, hostility which in turn leads to more illegal activity and a more negative perception of human-wildlife conflict. But equally, poor relations with the wider public mean that many Africans view their National Parks as rich white people's playgrounds, not for them at all. Such perceptions can lead to a widespread lack of support for conservation and, ultimately, a lack of political will to continue protection. In South Africa there's a lot going on to combat these problems - school visits are encouraged, entrance fees are very reasonable and accommodation is available for a variety of income groups. As a consequence, even in some of the areas where local hostility was historically high, tribal groups such as the Makuleke who have won back land they were disposessed of when Kruger was formed decided not to resettle the area, but to contract the park authority to continue to manage for ecotourism on their behalf. Other areas are allowing people to access the park to harvest thatching grass and medicinal plants, etc. And in Botswana's Okavango Delta, widespread antipathyto wildlife and tourism among neighbouring communities in the 1990s was reversed following implementation of a community-based natural resource management programme with revenues from tourism reaching community members (often through employment opportunities): poaching decreased dramatically and several mammal populations stabilised or increased by 2004.The only Tanzanians I usually see in parks like Manyara are guides like these!We've still got lots to learn in east Africa! Here, park fees are very cheap for nationals, but that ignores the fact you need a 4x4 to get around in the parks - which is well beyond almost everyone's means. Just as importantly, although the parks here raise money for the central government (50% of park fees go straight to the treasury - an idea that was completely unbelievable to our southern African visitors!), only tiny amounts make it back to the local communities - less, even, than the small amounts officially allocated: 1.8% of income was spent locally by TANAPA, when I last looked. We've got to do better than that here, and I'd love to see more Tanzanian's enjoying the parks for themselves too - can we persuade any of the safari companies to make their cars available at cost price to locals during the low season? Can we raise money to fund that somehow?Subsistence poaching is still a major problem for many Tanzanianprotected areas - note the snare on this giraffe.Related to this issue, but rather different too, was another mistake identified in southern Africa - insufficient law enforcement. Now, I have to confess that I was under the impression that southern African parks pretty much epitomise "fortress conservation", with big fences, highly trained and well equiped anti-poaching units and all the rest. But such efforts have only really managed to contain subsistance hunting for bushmeat - it's done little to halt the poaching of high value, commercial good like rhino horn and elephant tusks. And sadly, once these animals decline there are real impacts on ecosystems - it's only recently that white rhino numbers have got high enough for us to realise that they are important at forming species grazing lawns that are then used by a whole host of other animals. So what to do? Well, it certainly isn't wise to stop the law enforcement operations that are ongoing, but it clearly isn't enough on their own and more sucessful models have been shown - both the Botswana example I've already given, but also the story in Namibia, where where poaching of elephants and rhinos outside protected areas was almost eliminated initially by the employment of community game guards, and ultimately through the establishment of conservancies where control over the use of wildlife resources was delegated to representative local councils. There's lots to learn here in east Africa - but it's not going to happen without some major changes in the political landscape too, with corruption endemic in many places.Pastoralism is probably the wisest land use in many buffer areasThe human dimension we're bringing out in these two issues is also very much there in another issue: that of buffer zones. In southern Africa there's been little regard for land-use in the areas around protected areas and, given that most parks are fenced, that might not seem like a problem. But our southern partners really saw this as a major issue for a whole nuber of reasons. Primary among this was the fact that now conservation is higher up the agenda, and a really money-earner too, it's not possible in most cases to enlarge the parks or protected areas, except in a few areas where land remained relatively unconverted thanks to happy accidents, like some parts of 'greater' Kruger and Addo Elephant Park. But it also means that in some areas of the parks certain management tools aren't possible - you can't have fires where residential housing is riche on the edge of the park. And we also now think that although many of the areas not in protected areas weren't core areas for wildlife, they filled an important ecological role as population sinks - as the population in the core areas increased following a few good years, animals would move out and into those less good areas, stabilising the population in the core areas, so when bad ti... Read more »

  • June 2, 2012
  • 03:18 PM
  • 317 views

On managing protected areas...

by Colin Beale in Safari Ecology

Spot the scientists! Prizes for anyone who can name at least 4... In  a very rare burst of finishing things of, I've managed to submit two papers this week (wow!). One is on climate impacts and I'll blog about it in time, the other is something I've been working on for a some time that reports the deliberations from a workshop that I was invited to 18 months ago now, at Sasakwa Lodge in the Grumeti Game Reserve. This was a fascinating experience, and not only because it's the only way the likes of me will ever get to stay in Paul Tudor Jones' house and be looked after like a real guest! We brought together several senior researchers and conservation practitioners from Tanzania and Southern Africa, to see what would happen. And what did happen (as well as the lodge running out of whisky), was an attempt to identify the ten most important lessons for conservation that could be learnt from the mistakes of southern Africa. As they say, it's a wise man who learns from his mistakes, but it's an even wiser one who learns from the mistakes of others! So, given that the population pressures in east Africa are now similar to those experienced in southern Africa when lots of conservation interventions started to happen down there, we thought it would be a good time to see what we could learn.Workshops like this are the only chance for mere mortals like me to see inside this sort of place! Though it took days before I could walk on the zebra skins...It was a fascinating experience - and a lot of fun too! In honour of having finally submitted the paper (with, of course, no guarantees that it will be accepted, but at least it's off my desk for a bit), I thought I'd explore some of the 10 lessons here. Firstly, it's important to realise that conservation approaches in eastern and southern Africa are rather famously different - traditionally, Southern African protected areas are perceived as highly managed systems often with e.g. fences, artificial water holes, strict fire regimes and culling programmes, whereas East African (here meaning Kenya, Tanzania, Uganda, Rwanda and Burundi) protected areas are mostly unenclosed and have traditionally had a 'hands-off' management policy. Southern Africa was, of course, rather similar until the middle of the 20th century, when pressures from population and cattle ranchers (to prevent disease spread) led to most of the protected areas being fenced, which had a whole series of knock-on effects.Kruger, whilst rightly famous, has fences that cut once large mammal migrationsThe problem really started, not necessarily because a fence was put up, but because the fences were put up in the wrong places - and lesson one is that boundaries for protected areas need to be correct: As with protected areas in many parts of the word, the boundaries of southern African reserves were designated on the basis of pragmatism under increasing pressure for land: political boundaries, existing land use, tsetse fly presence, disease and opportunities presented by lack of settlement were more important considerations than ecological integrity. Indeed, at the time of gazetting, information about ecological boundaries of the protected systems was often lacking. Even South Africa’s largest two parks do not represent self-contained ecosystems. Before the formation of the current Kruger Park, Sabi Game Reserve was in the southern area and  provided the best habitats for rarer antelope, and a high-rainfall dry-season range for other mammals. But these wetter areas were excluded from the park when Sabi and Shingwedzi GR were merged to form Kruger NP. This cut the seasonal routes of migrant mammals, leading to smaller resident populations and vegetation change generated by year-round grazing pressure and disruption of fire patterns. Infrequent, but vital, movements in response to extreme conditions (e.g. to rarely used drought refugia) became impossible, leading to increased variation in annual survival.If we want to keep sights like this in East Africa, we have to work outside PAs tooHere in East Africa, many protected areas were designated based upon the experience of hunters, who were mainly aware of dry (hunting) season aggregations of animals. Even Serengeti NP was gazetted without knowledge of the migratory routes. Thus many protected areas in East Africa protect only dry season refugia - even the Serengeti NP, Mara Reserve and associated game reserves don't completely contain the wildebeest migration - each year sometime in May / June, they leave the Grumeti area of western Serengeti, and cross village land to reach Ikorongo GR and back into the National Park.If this ecosystem is to be adequately protected, these village lands should be protected too or we risk disrupting the entire ecosystem.Once borders are fixed, you've either succeeded or failed in protecting wildlife, and our group identified, in particular, the almost total failure in southern Africa to protect migratory mammal populations, and maintain adequate corridors between protected areas and another major mistake. You might not know that Southern Africa formerly had mass migrations of mammals, most spectacularly of huge numbers of springbok moving between the north-western Karoo and Namaqualand. Black wildebeest and blesbok probably migrated seasonally between the western Free State plains and the Maluti mountains. In Botswana's central Kalahari, a wildebeest population of c.200,000 animals collapsed when their dry season movement was blocked by a fence. Today, only in northern Botswana, where no fences exist, do substantial herbivore movements persist. In addition to being totally amazing, mass migrations have important influences on vegetation dynamics that cannot be maintained by resident populations, with year-round grazing commonly leading to woody plant invasion.  The minimally protected Simanjiro is birthing grounds to many Tarangire wildebeest. To maintain the Tarangire ecosystem, we need to work in this area.In East Africa, migratory mo... Read more »

  • May 31, 2012
  • 03:33 PM
  • 469 views

Why do scorpions fluoresce and other such trivia...

by Colin Beale in Safari Ecology

Scorpions - everyone's favourite! I have to admit that I find scorpions a bit creepy. Not only do they have too many legs to begin with, but some of them seem to accelerate from stationary to far too fast in no time at all. And, of course, some of them (a tiny minority, it's true) can be really rather nasty when they're pushed to it. However, despite the slight wariness they inspire in me, I do find them absolutely fascinating creatures. One of the big things that puzzled me about them was their bizarre fluorescence under ultra-violet light. If you've never been it, it's well well finding someone who's got a fluorescent light and taking them to the bush at night. You'll be amazed not only at how the beasties glow, but by quite how many of them there are! In many savanna habitats you'll see them every 3 or 4 metres. Hopefully enough to convince you never to walk in the dark without shoes! Why they should do this has been a mystery to me, but some new research published this year by Gaffin et al (find it here, but you have to pay...) has, perhaps, started to unravel the mystery.In normal light...And fluorescing under ultra-violetThe first thing to know is that scorpions don't like light. They're mostly nocturnal, and hide under rocks or in burrows during the day. Despite their defences, rather a lot of things like to eat scorpions and they're safer hiding away during the day. Shine a torch on them at night and they'll often head for the nearest hole too (which is why you don't realise quite how many there are around until you look with a fluorescent lamp!). Now, scorpions have eyes that can obviously sense light. In fact, most (all?) have eight eyes - a big pair on the top of the head and three pairs around the side. These eyes in the centre are, in fact, one of the most sensitive light gathering tools in nature and they can see perfectly well for hunting using only starlight. In fact, none of the eyes give clear vision, but they certainly do allow them to sense light and dark. However, the new research has shown that even if you block all these eyes, scorpions can still respond to light by running away. It seems that some species have light receptors in their tails, but by shining lights of different colours Gaffin et al have discovered that some eyes are particularly sensitive to the blue-green colours given off by a fluorescing scorpion. And as shining even faint ultra-violet light makes the scorpions themselves fluoresce blue-green, they are very sensitive to UV (ultra-violet). Fluorescence is a pretty good way of measuring light: as you'll know if you've ever tried making a room really dark when there's strong sun outside, you have to shut out even the tiniest bits before you make a very noticeable impact. Fluorescence, on the the hand, because it starts at much lower intensities is much more graduated. So what the scorpions may have evolved is  a 'whole-body' light measuring device which is exactly what you want if you want to scuttle backwards into a crack or hole whilst keeping your eyes (and pincers and sting) all facing the way of a predator. As the sun is the only significant natural source of UV, scorpions have a very sensitive way to detect when they start to find a suitable hole without needing to look.I'm not sure all the details are quite worked out yet, but I can see that there's some sense to this, and it's the best answer I've seen yet!Eight legs in a circle are perfect for vibration detectionAs well as fluorescing, of course, scorpions have a number of other handy tricks that I think are rather neat. Firstly, they have structures called pectens on the underside of their body. (Strangely, I don't have any close-ups of the underside that show them, but they're small to medium-sized hairbrush-like structures under the body, near the legs.) These they use to detect chemical signals - they're bigger in males than females and males certainly seem to use them to follow pheromone signals from females - though they also probably help when hunting prey. Pretty neat.And the other useful trick they have for helping hunt is a phenomenal ability to sense and locate vibrations. As you have no doubt noticed, it's possible for us to hear something and look in the right direction to see what caused it - we do that by measuring the difference in time it takes for the sound waves to reach our left and right ears, but we're not very good at it really. Owls have perfected the ability and can use these aural queues alone to hunt in total darkness. But for scorpions it's the legs that do the job - and they've got eight to use to give extremely precise distance and direction sensitivity. It's estimated that using very sensitive hairs on the legs they can pick up a vibration of less than one millionth of a millimetre and use the difference in timing it takes for a vibration to travel from one leg to another to pinpoint the source so they can turn and run exactly the right distance and direction to catch their prey, despite very poor vision. Now that is truly awesome!Main reference:Gaffin, D., Bumm, L., Taylor, M., Popokina, N., & Mann, S. (2012). Scorpion fluorescence and reaction to light Animal Behaviour, 83 (2), 429-436 DOI: 10.1016/j.anbehav.2011.11.014... Read more »

Gaffin, D., Bumm, L., Taylor, M., Popokina, N., & Mann, S. (2012) Scorpion fluorescence and reaction to light. Animal Behaviour, 83(2), 429-436. DOI: 10.1016/j.anbehav.2011.11.014  

  • April 3, 2012
  • 03:34 PM
  • 488 views

African Vulture Declines

by Colin Beale in Safari Ecology

I saw this hooded vulture in Tarangire this weekend, so they are still around!I've spent a bit of time over the last few days analysing some of the data from the Tanzania Bird Atlas project on vulture declines in advance of a workshop happening soon in the Maasai Mara. The Asian vulture decline is quite possibly the fastest decline in any bird species ever recorded, with more than 95% of the Indian population of Oriental White-backed Vultures dying between 1988 and 1999, from one of the commonest large raptors in the world to one of the rarest. It's now well know that the cause of that decline with the veterinary use of a drug called Diclofenac which, happily, isn't in quite the same usage here in Africa - sick o dying cows tend to be eaten here, not treated with drugs and then left for the vultures. But although the declines haven't been as steep and there are still plenty of vultures in places here in East Africa, there's still a problem.Changes in Egyptian Vulture distribution (after accounting for observer effort) from 1980/1990s to 2000s.Red are where the change has been observed, black is where the models fill in observer gaps.Juvenile Egyptian Vulture (note tail shape) on the Mara River,only recent evidence of breeding in TZ Sep 2011In the Maasai Mara, studies are suggesting around 60% decline in vulture numbers, whilst in South Africa and west Africa the problem is even greater, with near total loss of vultures in many areas. Monitoring in Tanzania isn't our strongest point, but the data that the Tanzanian bird atlas have produced show some obvious patterns here too. Look at these maps above I generated today of Egyptian Vulture, for example, showing the changes between the earlier records from 1970 and 1980s to post 2000 records. It's clear that until 1980 you had a chance of seeing this species across much of northern Tanzania. Since 2000 you've only really stood any chance at all in the Serengeti Ecosystem, and to be honest more recently it's extremely rare, with just a handful of records each year from this area. Whilst this is far and away the most extreme decline, my analysis today suggests there's also something happening with Ruppell's and White-backed in Tanzania, so we shouldn't be complacent.Ruppell's Vulture, Serengeti, Sep 2011So, what's going on? Well, most of the declines are probably due to poisoning. There's no shortage of meat in the parks of East Africa, so we can rule food out (although this isn't necessarily the case with declines in West Africa, where large mammal populations are virtually zero on many areas too). Poisoning occurs for all sorts of reasons, deliberate or accidental: people leaving poison out to (illegally) kill lions are likely to kill vultures too. In the past poachers are said to have diliberately have targetted vultures, since their movements may give anti-poaching teams an idea of where recently killed carcasses lie. And more recently, there's evidence that killed animals are being used to supply traditional medicine demand from elsewhere in Africa: if it weren't so damaging, I'd find it fascinating to learn that because vultures have such good eyesight, practitioners of juju in South Africa have attributed them with foresight and use their body parts to divine football results! One question we haven't really managed to address yet, though, is why the smaller species (Egyptian Vulture and Hooded Vulture) seem to have been declining first and fastest? Are they simply more exposed to poison, being rather more closely associated with humans? Or what? Any ideas gratefully received!Lappet-faced Vulture, Serengeti, Sep 2011Some, of course, might think it doesn't matter and we shouldn't care. But, as I've already said on this blog when talking about how to identify the various east Africa vulture species, vultures are really important ecologically as keystone scavengers. Take them away and other animals (and potentially people) suffer: populations of other scavengers such as dogs can increase, leading to increases in rabies (and the potential for infected animals to bite people, or spread this and other canine diseases to wild dogs, etc.). Human anthrax case increases are also attributed to vulture declines in some places. They also show other scavengers the way to dead animals, enabling a lot of opportunistic scavengers to benefit too. So I suggest we should be concerned. And if you're in Tanzania, do let us know where you see some of the rarer species - and in September I'm expecting there'll be a vulture counting day organised, so keep looking here and see how to get involved!Main reference:... Read more »

  • March 31, 2012
  • 09:54 AM
  • 548 views

Revising climate impacts on African vertebrates

by Colin Beale in Safari Ecology

A few weeks ago I wrote a piece on climate change and African vertebrates. As I usually do, and especially in this case as Raquel had pointed the paper out to me, I let her know that I'd written something and asked her opinion. After quite a few emails back and forth we confirmed that I'd misunderstood a figure in the paper that I'd thought was the crux of the matter, but it turns out to have been not as useful at all.In light of these discussions, Raquel and colleagues have now produced an addendum to their paper that contains the figure I thought I was looking at and, although I still have some issues with the work, it makes much more sense to me now! In the interests of getting all this information out there, as well as my pointing out the mistakes in the original post and Raquel posting a comment there, I thought her ideas were valuable enough to reprint in full from the comment as a new post, with some more discussion here. So, here's what she has to say:I’m glad that our study has prompted Colin to start blogging about climate change – or, more precisely, about bioclimatic envelope models and their uncertainties under climate change. There has been much discussion around these models and the conceptual frameworks that underpin them.  But we all (seem to) agree that these models have various uncertainties, and that their results and the use of these results are contingent on a series of assumptions.  The focus of the study is on uncertainties First of all, the focus of the study was to explore model uncertainties. We examine three sources of uncertainty in projections of potential climatic suitability for African vertebrates (more on the projections themselves below).  Just as Colin and many other researchers argue, it is important to try to identify and quantify the uncertainty surrounding the results from models.  Doing that helps us to know where we should put our efforts to reduce the uncertainty, and can guide the interpretation and use of results.Our study explores the variability in projections from the choice of modelling technique, climate model and emissions scenario.  It quantifies the relative importance of each, and maps them spatially.  Of these three sources of uncertainty, the modelling technique was the major one, but the importance of climate projections increases with longer time horizons.  The study further cautions against projections onto novel climates, where these models are less reliable.  For thevariables used in our study, climates beyond the current climatic range were pervasive in northern sub-Saharan Africa by the end of the century. If we cannot identify the most appropriate model or climate scenario, the safest is to use a range of alternatives.  How to then interpret the ensuing range of projections has been a topic of debate. We too caution that the simple averages depicted in Colin’s cartoon mayimply loss of information.  But one of the consensus techniques we apply overcomes this problem, to some extent, by first grouping similar projections, and then averaging each group.  Our study compares these and other techniques to combine projections.  The aim was tocontribute to the debate around building consensus as one possible way to explore the uncertainty that surrounds ensembles of projections – another way being a probabilistic approach. What are we modelling? Now to the projections we used to explore uncertainties.  If we ignore model uncertainties or assumptions, it is easy (I should say inevitable) to interpret model outputs as saying something that they can’t say, or to use them for something that they cannot be used for.  If we closed our eyes to the assumptions behind our study, we would say that we are forecasting the fate of African vertebrates by the end of the century, and showing on the map where species will remain or disappear.  Reading Colin’s blog, it may seem this is what we were saying.  But it was not.  The assumptions are clearly stated, reflected in the interpretation of the results, and revisited in the discussion.  Our models are not built to provide forecasts of actual distributions but rather potential areas of climatic suitability for species.And these two parameters, just like Colin argues, can be very different indeed. At the large scale of our study, climate is one determinant of species distributions. But climatically suitable areas and actual distributions are different because of a range of factors operating at different scales.  We mention some factors highlighted by Colin, such as dispersal, fire, and biotic interactions.  These factors were not accounted for in our projections of climatic suitability, yet they would surely need to be considered in projections of species distributions.  When assessing shifts in climatic suitability, there is uncertainty also in the species data used.  Past history and biogeography can limit the extent to which the species data available reflect the full climatic tolerance levels of the species.  We also mention the contingency of the results on the climatic variables selected.  Given the large number of species, we have compiled a set of bioclimatic variables (including important seasonal variables, such as precipitation seasonality or the precipitation of the driest quarter), and selected uncorrelated variables that explain a large part of the variation over the study area.Colin bases most of his discussion about the plausibility of the projections on the maps of climatic suitability retention shown in the blog (Figure 7 in the paper).  Colin assumed that these maps depict local persistence as a proportion of local species richness.  But in fact what they show is local persistence in relation to total richness (as the caption in the paper explains). Persistence calculated in this manner is thus much lower than otherwise, and emphasises areas of high current richness. Maps with the more commonly used metric of in situ persistence, as we present in an addendum available on the list of publications of our group (http://www.ibiochange..../Garcia-et-al-2012-Addendum.pdf), show a very different picture for African vertebrates. The blog also reproduces the maps of turnover that we used to show projected changes over time for all species.  Our percentage of species turnover, for which a reference is provided in the text, was calculated as the ratio of the sum of local gains and losses (of climatic suitability) to the sum of local baseline richness and gains (of climatic suitability). Evaluating the projections is difficult, among other reasons, because the models might rightly predict that a site is suitable when the data indicate absence.  What our study does is simply to select one measure of accuracy and compare the relative accuracy of the different projections.  The focus is on relative, not absolute terms. One thing to remember also, when interpreting projections like these, is that the patterns shown are mostly for wide-ranging species (the ones that could be statistically modelled).  Our projections refer to about two thirds of the species and do not necessarily reflect the patterns for small-ranging species.Fit for the purpose?Only highly complex models could begin to tell us where African vertebrate species might die off or survive under changing climates.  Until we have the knowledge and data to build these super-models for thousands of species, we are left with imperfect models explaining part of the reality.  If we are clear about what part of the reality the models are explaining (assumptions) and where the imperfections are (uncertainties), they will serve a purpose. And the purpose in our paper was to explore the challenges posed by the variety of climatic projections and modelling techniques available when examining broad patterns of climatic suitability for these species. Percentage of species predicted to retain climatic suitability under each emissions scenario. The proportion of the local numbers of species of amphibians, snakes, mammals and birds that are projected to retain climaticsuitability in each location are shown for the median ensemble of allbioclimatic envelope models and for the ‘maximum consensus’ general... Read more »

  • March 28, 2012
  • 01:17 PM
  • 446 views

On cattle in African protected areas

by Colin Beale in Safari Ecology

Typical pastoralist scene near Lake Eyasi Talking about blog topics the other day, a friend asked me about the impact of goats and cattle on wildlife. And then over here someone else started a similar discussion on cattle, which collected a wealth of different ideas, so I thought it would be a good idea to collate all this information for a different audience over here. Increasingly, discussions about cattle come up when people are visiting areas that aren't National Parks - here in Tanzania many people are surprised to see cattle (and their Maasai herders) right in the Ngorongoro crater, as well as around the rest of the NCA. And increasingly (particularly in Kenya where land laws make it much easier, but also here in places like Manyara Ranch) conservancies are being set up where communities set aside land for both wildlife and pastoralist activities. The fact that organisations like the Northern Rangelands Trust are making a real success of this, combined with ongoing concerns about displaced people and human rights issues, has encouraged people to think seriously again about whether the strict 'no people' policy of many national parks in Africa might be relaxed, and recognising this a few years ago the International Conservation Union (IUCN) relaxed their national park category definition to allow management "To take into account the needs of indigenous people and local communities, including subsistence resource use, in so far as these will not adversely affect the primary management objective". So, what are the issues here, and what are the ecological arguments? In this post I'm going to deal with cattle, and leave the goats and sheep for a future occasion.Cattle numbers in south and east Africa (data from FAO database).Do we always need to move cattle away from wildlife areas? To start, I think we need to be absolutely clear that the idea of an 'empty Africa' is a romantic, colonial-era myth: bear in mind that for at least 2 million year 'human' influences have been important in savannas (starting with fire and hunting), so the savanna plants and animals have evolved in a human-influenced landscape (those first hunters weren't hunting the same species we see today!). Now genetic, archaeological and anthropological evidence suggesting there have been cattle in sub-Saharan Africa for 9-10,000 years, including within all the major protected areas of Tanzania. As protected areas become increasingly isolated and the populations of animals are heavily influenced despite our best intentions, human management of the savanna is going to become ever more important, if we want to maintain healthy savanna ecosystems. Despite the history of cattle for 1000's of years, the numbers of cattle before the 1900s are not precisely know - evidence suggests around a 90% decline in numbers around 1900 associated with the great rinderpest epidemic and a gradual restocking since then, with a recent fairly rapid (last 10 years) increase in some countries/regions. It is probable the numbers are now higher than than pre-1900s. Despite this, increases are patchy and areas (such as NCAA) exist among the fast-growing areas where the population is actually declining. As well as an increase in numbers, movements by nomadic pastoralists have been restricted - both from fences on commercial farms, by preventing them from using traditional grazing areas within protected areas and through policies designed and/or incidentally decreasing the nomadic lifestyle (compulsory schooling, forced settlement, provision of water points, etc.). In many pastoralist communities number of cattle is still the primary indicator of wealth, not quality of cattle. Even in communities where this perception is changing, there is a 'tragedy of the commons' that prevents effective herd reductions: if I'm a herder sharing grazing lands with others and I decide to reduce my herd size to get fatter animals, that has no impact if all the other herders around increase their cattle herds to eat the food my cows are no longer eating... All these things together mean that there are places within Africa where overgrazing  (by both cattle and goats) is a serious concern, cause permanent damage to the ecology of a system, creating open soils that are easily eroded by water and wind and are then surprisingly hard to revegetate. Nothing (cattle or wildlife) can survive in these areas once the annual grasses are grazed off each year. Pastoralists seeing uneaten grass in their traditional grazing lands within protected areas are either thoroughly fed up with conservation, or go ahead and graze anyway (or both), potentially resulting in unmanaged grazing within protected areas with cattle unprotected from predators and real risks of lion killings inside protected areas.Meanwhile, wildlife numbers in the wider countryside have fallen dramatically, with noted declines around 1940s (leading to formation of the National Parks and Game Reserves by the colonial authorities) and in the last 30 years. Reasons for these declines in the literature focus on unsustainable hunting/poaching and (for the recent decline) an increase in agriculture in rangelands (with new cultural groups moving in who have a taste for bushmeat), with increased competition for grass from increasing cattle populations a relatively minor issue , despite the numbers of opinion pieces that discuss its possibility (eg here and here). The best ... 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  • March 26, 2012
  • 06:33 AM
  • 681 views

Why do savanna trees have flat tops?

by Colin Beale in Safari Ecology

Umbrella Thorn, Serengeti: An icon of the savanna? From sunsets behind a silhouetted acacia (properly Vachellia), to photos of rolling grasslands studded with isolated trees, a savanna landscape is immediately identifiable thanks to the flat-topped tree. But why is this? Why do so many Vachellia and other savanna trees have such a distinctive structure that they have become a virtual icon of the African savanna?It's an interesting question that was given some answers in a nice paper by Sally Archibald and William Bond who studied one species called the Sweet Thorn (Vachellia karroo) that, rather like some of our Vachellia species in East Africa exhibits a range of different growth forms in different habitats. In the semi-desert of the Karroo, it grows as a medium-sized ball of thorns, whereas in the savanna it has a fairly typical medium-tall  flat-topped acacia look to it and in a forest it's a tall, thin tree. These differences are meditated mainly by genetic differences within the species, but equally could be caused in other species by a variable response to the environment - it's not really important to this discussion and, in fact, much of our discussion could focus on different species if we wanted. As always when we're thinking about what makes the savanna species, we'd be well advised to start with the savanna big four: nutrients, water availability, fire and herbivory.Now, the first two processes have impacts in all biomes, whereas it's the second two that are most distinctive about savanna and where we'll start our discussion.It's not only Vachellia species that are flat-topped. Here are a range of species including Balanites aegyptiaca in Serengeti.Any tree that wants to get established in a fire-dominated biome has to have some special adaptations to cope. Thick bark is an obviously useful trait and has evolved repeatedly by trees that invade savannas globally. But another sensible trait is to grow rapidly tall: trees that are above the 'fire trap' (2-3m tall) aren't burnt back to the roots by a fire, but can resprout from the top. If you've got limited resources to use and live in a place where fires are frequent, it makes sense to put those resources into a single (or at most a few) stems and grow straight up, not branch out sideways. Once you've grown tall enough to escape the impact of fire, you're free to branch sideways. So there's one good reason why in a savanna you might find lots of tall trees with few branches low down then lots more above.Senegalia mellifera is often in drier areas and is typically a 'ball of thorns'.Next, consider herbivory. Growing tall isn't easy if you're a young tree and there are lots of browsers around. In such circumstances you'd be well-advised to put a lot of effort into protecting yourself with loads of thorns and grow in a thicket type growth form. By growing wide enough, you might eventually manage to protect some internal branches enough to completely free them from browsing pressure and they could then grow tall if that's useful. So in a browser dominated habitat, branching low down and frequently might be a much better strategy. As we know, in the savanna fire is carried by grass, so in areas where rainfall is low and grass growth small, we'll find a browse dominated system - and it's those areas where William and Sally found ball-shaped thorn trees. (Of course, in wetter areas where there's both fire and herbivory, it's a really struggle for any tree! In such circumstances I guess the certainty that all seedlings will be killed by fire if they're not tall enough, versus the possibility - however small - that a seedling my not be found by a browser would favour a fire-adapted response. Ideally backed up with some serious thorns and chemical defence of saplings!)Forest on Kilimanjaro - trees here are tall and have branches to catch the light passing through the top layerNow, in contrast to a fire and herbivory dominated savanna, in a forest trees are cramped together and their main struggle is to get sufficient light to grow. Consequently, different tree species have evolved two main strategies to survive: they either tolerate low light and grow very, very slowly but live a long time and eventually make it to the top of the canopy and can shade out other less-tolerate species. Or they grow fast and die young, rapidly filling any holes that form in the canopy when a mature tree dies, but eventually being out competed by the slower-growing species. They also need to absolutely maximise any light they can get - even if that light has already passed through the top leaves, they want to catch remaining light lower down too, making the forest floor a rather gloomy place to be. The consequences of this intense light competition are therefore that (a) mature trees tend to be very tall, always struggling to get above their neighbours and (b) they have lots of leaves at all sorts of heights in the canopy, trying to catch every stray bit of light.I love Vachellia tortilis! Another in Serengeti.So, in a typical fire-dominated savanna, once you've escaped the fire trap you're free to branch. But it's so rare that a seedling breaks out of both fire and herbivory, that mature trees tend to be at low density in savannas, with canopy cover in woodlands around 20-30%. And under such circumstances there's no going to be any significant competition for light - so there's no need for savanna trees to grow tall, and horizontal growth is enough to gather sufficient light. What's more, even moderately tall trees don't always escape giraffe browsing, but a horizontal growth-form still protects the central branches - in fact, it's common to see larg... Read more »

  • March 22, 2012
  • 01:50 AM
  • 604 views

Why is snake venom so toxic?

by Colin Beale in Safari Ecology

Puff-adders probably cause more human snake-bites than any otherAfrican snake,but are rarely fatal. This is a juvenile, but don't think it's harmless. After discovering all the amazing things about pedarin and the 'Nairobi Eye' last week, it set me thinking again about why so much wildlife is so incredibly toxic. Think about it - a little beetle small enough to crawl over you without you noticing at all, is more than toxic enough to kill a grown man - indeed, several. A snake like a black mamba can give a bite that's sufficient in toxicity and volume to kill an adult elephant. Many natural venoms aren't simply one chemical, but a mixture of nasty toxins with a whole range of activities - why go to the trouble of evolving a whole suite of nasty chemicals, when one is usually enough to kill most things? Why should it be so toxic? What's the purpose? This is a topic that's puzzled lots of people for quite a long time, and a number of different answers have been proposed with no one clear winner. The two main threads of answers have been proposed: firstly, that there's no effective natural selection on snake venom, once evolved, venom toxicity just drifts along. Others suggest exactly the opposite: that it must be a trait that is subject to extremely high selection pressures to have evolved as it has. Now, the argument for a neutral drift stems mainly from the observation that non-venomous snakes exist and, indeed thrive: if one species manages without venom, but another has it and both do just fine, it would seem clear that evolution has nothing to do with it. On the surface it's a pretty convincing idea, but I wonder too if there's some slightly dubious thinking going on here. Remember the thorn story: many plants here have thorns, but the animals eat them anyway, so why should they have evolved? I think the same sort of idea is happening here: we're looking at a situation that may seem daft today, but wasn't before.Female Boomslang - you'll probably die painfully in 2-5 days if not treated. Happily, they're rather shy.So let's look at the evidence suggesting an evolutionary benefit to the extreme toxicity of venom. Let's start by thinking why snakes have venom in the first place. They use it for two things (1) to kill their prey and (2) as a defence against threats. Certainly, for a defence mechanism alone, you don't need to kill - in fact, you might even be better off not doing so and just giving a warning that can be passed on to other potential predators. So let's concentrate on what seems the most likely initial reason to evolve venom in the first place: to kill and subdue prey. Clearly if you evolve venom for one reason (attack), it's perfectly reasonable to later use it for self defence, but it's also possible the attack reason is still driving the whole process. Now, a venomous snake finding something tasty to eat strikes quickly and usually lets go immediately, waiting for the prey to die at a safe distance. Not a bad idea if you want to take down something larger or stronger than yourself. So it's important to that snake that whatever it's bitten dies relatively quickly: too slow and the animal might first have had the chance to run away and the poor snake goes hungry. So although there might be enough venom in a mamba bite to kill an elephant, it takes hours and hours - not fast enough if the snake wanted to actually eat the elephant. So if it wanted to hunt a mouse it might only need a drop of around 0.185 mg (average of various estimates) to have a 50% chance of killing that mouse, it would probably have no chance whatsoever of actually eating the think unless a bite killed within seconds, which requires a much larger dose.Boomslang showing rear fangs typical of colubrid snakes. Don't try this at home folks...Moreover, we know there's a large cost involved in producing venom - snakes that have  recently bitten and need to re-fill their venom glands have a metabolic rate 11% higher than snakes with no such need. This also explains why some snakes when acting in self-defence don't always inject venom - the bite itself is warning enough. And we also know that some snakes do even adjust the amount of venom they use according to the mass of the animal they're biting. So there's obviously a cost of generating venom that means it would be strongly selected against, if there was really no need for it, yet there seems remarkably little evidence that snakes have lost the ability to produce venom, whilst the ability appears to have evolved in at least three different groups separately. Now, although there's certainly a cost involved in producing venom, it's not clear to me that the costs involved in producing a rather less toxic venom would be significantly lower than those of producing a much more toxic venom - once you've paid the initial cost, producing different variations on a theme might not be additionally costly. So if it's relatively easy to vary the basic molecules involved in toxicity (and it seems it is), it would make perfect sense to evolve a whole suite of chemicals, each with slightly different action making the overall combination toxic in the extreme. What's more, this also fits with what we know about resistance.Just as bacteria can evolve resistance to our antibiotics, so too can resistance evolve in prey to snake venom. Now, resistance is most likely to evolve in bacteria when the doses aren't kept high enough for long enough to kill everything - that's why it's crucial to complete a course of antibiotics (if you don't, there might be some bacteria that were nearly resistant still alive when you stop, and you've just selected very strongly for those strains that are close to evolving proper resistance). It's surely the same with snake venom: it's much harder to evolve resistance to a massive dose of a toxic nasty, than it is to a smaller dose that some individuals might just survive. We also use multiple antibiotics when we're trying to protect against bacteria developing resistance (it's much harder to simultaneously evolve resistance to two antibiotics than just one), which seems like a plausible reason for why venom is full of a whole range of toxins too. Despite this, resistance can still evolve in prey animals, so the doses need to keep getting larger and larger - just like thorns do - with the result that when used in self defence against an animals that has no evolved immunity because it's so rare to get bitten, the dose is incredibly toxic. All good reasons for why evolution might well lead to what looks like 'overkill' in snake venom. ... Read more »

  • March 19, 2012
  • 02:04 PM
  • 508 views

Distribution of Ethiopian Bush-crow and the nature of explanations

by Colin Beale in Safari Ecology

Yesterday I was sent a link to a press release from the excellent BirdLife International (read it here). It's talking about some research by an international team to try and explain the remarkably restricted range of the Ethiopian Bush-crow (cute picture here, since I've never actually been there to take my own), and in it, Paul Donald the lead author makes some interesting comments:“The mystery surrounding this bird and its odd behaviour has stumped scientists for decades – many have looked and failed to find an answer.  But the reason they failed, we now believe, is that they were looking for a barrier invisible to the human eye, like a glass wall. Inside the ‘climate bubble’, where the average temperature is less than 20°C, the bush-crow is almost everywhere.  Outside, where the average temperature hits 20°C or more, there are no bush-crows at all.  A cool bird, that appears to like staying that way.” The reason this species is so completely trapped inside its little bubble is as yet unknown, but it seems likely that it is physically limited by temperature – either the adults, or more likely its chicks, simply cannot survive outside the bubble, even though there are thousands of square miles of identical habitat all around.BirdLife International’s Dr Nigel Collar is co-author of the study. He added “Whatever the reason this bird is confined to a bubble, alarm bells are now ringing loudly.  The storm of climate change threatens to swamp the bush-crow’s little climatic lifeboat – and once it’s gone, it’s gone for good.”Two things jump out from this to me (1) the nature and validity of the explanation itself: we're told the authors think there's probably a direct physical limitation, not an indirect effect on, say, a essential food source. (2) The concern about climate change removing the suitable bubble of cool in the highlands. (I should point out that the paper is available here and I'm going to base the rest of the discussion on the actual paper, not the press release - both these points are made in the discussion of the paper, however.) The first thing that interests me is the statement about other scientists failing to find an explanation for this distribution, and I wonder whether actually this point comes down to a change in what is considered a sufficient explanation for ecologists. There's also been another recent paper on the way biologists have changed how they think about explanations over the last few decades. This paper, by two philosophers of science, charts the first discussions of different levels of explanation by Ernst Mayr back in 1961 who defined proximate explanations (e.g. it got too hot and the bird's died of sheatstroke) and ultimate causes (people burnt fossil fuels, releasing carbon dioxide into the atmosphere leading to global climate change, and that was the end of the species). They're both valid descriptions, but they answer slightly different questions. Not long after then Niko Tinbergen (who my little brother did his PhD about, by the way - rather strange having a brother who studied biologists..) also refined these further saying that before something could really be explained you needed to answer four types of question what's the physiological mechanism? What's the development pathway that gets you there? What's the ecological function? And what's the evolutionary history? Now, the latest paper suggests the whole cause and effect idea might be more complex than we originally thought, with evolution acting in feedback sometimes - but that's a whole different set of ideas. Still, ecologists looking for an explanation of distribution over the last few decades may have been looking to explain distribution in these ways. Recently, I'm wondering if we settle too fast on a rather superficial explanation that doesn't really meet these criteria.Certainly, no-one could call the work on the Bush-crow an explanation for the distribution according to any of the four criteria Tinbergen was looking for. And of Mayr's original two it might, perhaps, be an ultimate explanation. But let's look at that - in this study, as many others of the type, the authors started by looking at five different climatic variables (we don't know why they chose these from the 19 they had available): mean annual temperature, mean annual precipitation, temperature seasonality, mean diurnal temperature range and annual temperature range. Now, let's ignore the results for a moment and ask what sort of explanation we could possibly derive from these chosen variables. Could any come close to identifying a physiological limit? As they're all means (IE averages) I really don't think so: by definition the actual temperatue on any one day is pretty much certain to be above or below the mean. Similarly, it seems likely that a physiological limit to mean diurnal termperature variation is unlikely - though if the variation is too high on any one day I could see a real limit. Unfortunately, there's no way to tell from an average alone whether the sort of extremes that could potentially cause direct physiological problems are present as well. So instead of a direct (proximate) limit, using means is more likely to identify an indirect effect: one mediated and perhaps integrated over time and space by something else, or maybe the mean is simple pointing to some associated measure that is correlated with it. That correlated variable could be the extremes that might ahve direct effects, or it could be the demographic of some slow-growing plant that's essential for nesting, for example. In fact, the author's rule out any indirect effect by noting that the species is a generalist feeder that likes degraded habitats - unlikely to suffer any shortage of food or nesting spaces in any habitat. So we're left with the explanation that mean temperature (which the authors show has the highest association with the observed range) might be acting as a proxy for some as yet unknown temperature-related variable that could have a direct effect.Now, even without bringing up my general concerns with this sort of analysis (e.g. that the statistics used are inappropriate for spatial data and will 'explain' any distribution you care to throw at them), I have to wonder if this 'explanation' for the bird's distribution has never before been suggested for the simple reason that it's never before been considered adequate? Indeed, is it, as a biological explanation, any more satisfactory that simply saying these birds live on hills in southern Ethiopia? Looking at one of the main figures (right) from the paper I can see two things that would seems surprising to me: firstly both preferred temperature and preferred rainfall at incredibly narrow. If this is true then, as the authors rightly point out, it would be of extreme concern given the likelihood of future change in these areas. But it seems implausible to me: annual rainfall in the 600mm range of eastern Africa is so variable year on year that the average itself is essentially meaningless to anything except a long-lived plant like a tree or slow-growing bush. I can't imagine a plausible ecological mechanism that would allow such a restricted range on a map of mean rainfall, when the annual rainfall is such that it rarely falls within this optimal range anyway! Secondly, I see in the seasonality plot two peaks - the birds apparently like moderate seasonality and quite high seasonality, but don't 'moderate-to-high' so much. Such a dislike of intermediate seasonality immediately rings alarm-bells for me: it's physiologically not plausible and, together with the implausibly narrow limits identified for rainfall suggests the model is 'over-fitted' - in other words, it's not going to be reliable for prediction of distribution under changed climate conditions.Unfortunately, therefore, I'm not convinced we've really learnt much here. Certainly, we have no explanation of distribution that could meet any of the older types of biological explanation (nor the newly suggested one), but we do have a fairly typical example of what seems to be a lowering of the barrier in ecological explanations: to me, a suspect correlation with little critical ecological discussion cannot be considered an explanation. Come on ecologists, we're better than this!Main Reference:... Read more »

  • March 12, 2012
  • 08:20 AM
  • 642 views

Why do birds sing in the morning?

by Colin Beale in Safari Ecology

Ruppell's Robin-chat: an impressive mimic. Lake Duluti I enjoyed a walk around Lake Duluti yesterday morning and came across a couple of wonderfully singing Ruppell's Robin-chats. These are great birds, with an amazingly varied song hat's gull of mimicry (of you want to hear one, listen here!). For me, one of the best things about camping in the bush is being able to lie in bed and listen to the birds waking up while it's still too dim to see them properly. The dawn chorus is a worldwide phenomenon and I'm often asked about bird song, so I thought it would be worth exploring some of the theories behind bird song, and - particularly - why birds sing in the morning. It's something that's interested me since I was introduced to the question by a friend of mine who did a PhD on the subject some years ago, and I know he reads the blog so I'm hoping he'll make sure I get the answers right!There are two parts to the question, of course: why do birds sing? And why do they sing in the morning more than at other times of day? The first I think we've had a pretty good idea about for a long time, and there are two main reasons: to attract mates, and to claim their territory. In general the two actions aren't mutually exclusive - as a male bird you can sing to both let the other males around know that you're still in your territory, and you can at the same time let females know that you're around and looking for a mate. For some species by listening to the song you can tell what the bird is actually trying to say  - Nightingales are well known in Europe for their beautiful song, but it's not uncommon to hear them singing down here during the non-breeding season too (they seem to be more territorial than many migant species). The difference is that here, I've almost never hear the typical long whistle notes that are so frequent in the breeding areas during the early breeding season (listen to them here). They also use even more of these whistles during the middle of the night before they find a mate, and it seems that these whistles are particularly for attracting females, which obviously isn't relevant when not breeding. As females might be flying over at night, singing in the midle of the night and whilsting a lot, might be a very good way to attract a female to your territory.Yellow-vented Bulbul - nesting all year round in my garden (March 2012), both sexes sing far too early in the morning for comfort...For some species song is probably more important for attracting females than in other species - particularly, I would suggest, for species where mimicry is very important. Like my Ruppell's Robin-chat, many birds are accomplished mimics, and observations suggest that the more varied an individual male's repertiore, the more attractive the male. The most impressive example of this that I'm aware of is the Marsh Warbler, a species that we mainly see on passage here in Tanzania from time to time - though it is another than sometimes sings during the non-breeding season. In what's become one of the classic studies of song mimicry, 30 individual males were recorded during the breeding season and the songs then analysed and mimicry identified. The total diversity of song mimmiced by these 30 males was 212 species, 113 of which were African species! On average, each male knew the songs of 76 different species, mostly African - one male included identifiable fragments of 84 species - an astonishing variety of songs and calls! Why do they do this? Well, it seems most likely that females prefer the males that have the most varied songs - males that can learn lots of different songs have clearly got some good mental abilities, whilst - the hypothesis goes - must be tied to their ability to find food, avoid predators or otherwise help around the nest. So there's not only a chance to let a female know that you're around like the nightingales do, but there's a real competion going on to be the best singer of all.As I said before, there's also plenty of evidence that it's not only about attracting a female: even after findinga  female and her sitting on the nest many species keep singing for a while - this is clearly about territory defence. And the first example of how effective this is comes from a nice experiment where birds were removed from their territories and the time it took other birds to come and occupy the patches was recorded: it's pretty quick, but if you play a tape recording of the song of the resident male even though he's been removed, it takes a lot longer. Clearly birds looking for a territory recognise individual songs and as long as the territory holder is still around and still singing they won't bother trying to take over - an efficient system of defence if ever there was one.Usambiro Barbet, a typical African species with duets. Serengeti, Dec 2011.Now, that's the general answer to why birds sing. But here in Africa there's an additional complication we should consider: here, many female birds sing too. This isn't unknown in the north, but it's fairly unusual. And we've already seen that a whole load of African birds engage in complex duets too, with both male and female taking part. Why is it that many more females take part in song here in Africa and the rest of the tropics than they do elsewhere? There's no good answer to this one yet, but a couple of interesting theories are being developed that are worth mentioning. Firstly, as we've already talked about, birds in the tropics tend to be rather long lived - they do things slowly, and they have a rather tougher time raising young in many ways than in more temperate regions. So here they're often also with their partner longer, and often holding territories year-round - lots more reason for females to be involved in territory defence as well as males, so that's one possible reason. The other theory is that here in Africa the seasons might be harder to define (at least in the forests), with breeding possible much of the year (certainly the bulbuls in my garden think it's fine to nest all year around). But birds don't tend to keep their ovaries or other bits of breeding equipment large and ready to breed year round - it would be a waste energy to carry around fully developed genetalia if you're not going to use them, and flight is tricky. So in a less seasonal environment, how do both male and female make sure they're in breeding condition at the same time? Well, perhaps, by singing to each other - there's certainly some evidence that song can help bring on breeding condition in some birds. Interesting thoughts though, and (yet) another area of tropical ornithology where we're well behind on theory and experiment.So. Having tried to answer why birds sing, let's tackle why we think they sing most in the morning. There are a number of (not mutually exlusive) theories here too. They all start with the assumption that singing is good (for the reasons we've covered above) - but it's costly in terms of time and energy (another reason why females might like males that sing more: they're showing honestly h... Read more »

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