Lecture 3 - Cradles or arks, automated transcript May 28, 2021 --- Hi. So this is the last in a series of three lectures looking at Tropical diversity. And in the first lecture, we looked at by geography and why we get different species in different places. In last lecture, we looked at why the tropics are so warm and wet the ecosystem that gives rise to particularly rainforests and why and the human threats that face those ecosystems. And why that's important in this one, we're going to think about why even though we had a look at why the tropics are so warm and wet and ever. Think about why that, why the tropics are so bio diverse because even though they have high rainfall, they're kind of warm all year round. That doesn't necessarily lead to the extraordinarily high amounts, high amounts of diversity that we see in these regions. So why? That we get and such high amounts of biodiversity in the tropics. And why do we get this thing? Which is called the latitudinal biodiversity gradient, which we see in the present day. So that here is illustrated by this work which looking at terrestrial vertebrate species. So the concentration of diversity around the world varies by color. So, very high diversity, we see in red, and then goes through green, it goes to yellow and green down to Blue, which has a areas of very low do. Varsity. As we can see the high diversity areas of the world are very highly concentrated in central and southern Africa, in Southeast Asia, and particularly in South America. And the kind of the deep Amazon of the western side of the Amazon. So why do we see that? Why do we see higher levels of diversity in the tropics? That does that decreases as you move away from the tropics towards the poles? There are three kind of large hypotheses for the latitudinal biodiversity gradient, which I'll continue to refer to as the LBG. So we have higher rates of speciation in the tropics, which is the Cradle hypothesis. Then there's less Extinction in the tropics, which is the Arcon Museum hypothesis. And then we have the last one here which is a mix of the two where you get both these things are going. So we see on these graphs here where blue lines is kind of originated. No speciation rates, some left hand side. They would be higher in the tropics and Extinction rates would be flat across all latitudes. And then we have the graphs which show this for the other two hypotheses as well, which I won't go into too much detail because I'm sure you can and working out. As well as the use those hypotheses. We have to think about kind of what's, what would what factors that cause either higher, or lower speciation or Extinction rates across that latitudinal gradient. So why would we see these higher and lower Extinction rates? The first one is a geographical, which looks at the greater area so I've misspelled area their area extent of the drop. And which allows it to create support more species compared to other regions. So if there's greater area, then maybe there's going to be lower Extinction rates because if we think about a species area relationship, perhaps if speciation events are our constant everywhere. So the second hypothesis are the our hypothesis, but less Species will go extinct in the tropics because there's a greater area that can support them. And we have a look at climatic and also historical, I'm going to go through these in turn now and think about why the evidence that each of these. So once when too long, the A on cause it's fairly self-explanatory, but as the Earth is, almost is a sphere for all intensive purposes. It slightly slightly wonky, but it is essentially a sphere. As we talked about in terms of the speed of the earth rotation is also greater area at the equator. That's so when you think about a map of the world, you tend to think of the kind of classic one, is the Mercator Projection. And what I mean by a projection is that because the Earth is a sphere to be able to display it on a flat surface. You have to slightly alter what it looks like in order to do that, in order to kind of unroll the surface and put it on a flat surface. And so the classic way of doing that is called McKay to projection. And what that does is it sort of makes the land closer to the poles and look a lot bigger, and that's why I Greenland tend to looks huge because it's quite close to North Pole. And that makes this kind of forget that that places that are close to the equator. There's lots of land area because they kind of get squashed made it a little smaller on those projections. So the as an idea about whether if there's greater area in the, in the tropics, Except marine and terrestrial Realms, then then, are we able to support more life? In this place is just do, in fact, the grades area available and there's also thinking here looking at the distribution of tropical temperate. Paulo zones is symmetrical around it around the Equator. So, polar and temperature regions are disjunct separated by intervening tropical areas. In contrast, northern and southern tropical zones are adjacent. And so, what this means here is that, so we think about the Earth from equator up to the polls, we have tropical regions in one band around the middle. So, tropical species can theoretically move kind of along North and South between the Tropic of Cancer, the Tropic of Capricorn, without having to go into a different massively different climatic environment they can stay within that warm and wet climate. However, if a temperate species in the north wants to move. Temperate species in the South, it has to cross that tropical band and it may not be able to do that because then we have to cope with the conditions in the tropics. So perhaps that has meant that extinctions. Haven't been as high in the tropics because there's been able to be dispersion around within that one band. Whereas the polar and temperate species can't migrate to the other region because of the tropical realm being in the way. So this is is a just a theory. I don't want to too much here, but there is a paper that's in the recommended reading at the end. Looks at some of the evidence. This so climactic again. So we could just to remind ourselves of what these three hypotheses are. And so here, we're going to look at why speciation might be higher in the tropics due to climatic factors. and also, actually this kind of thinks that little bit about why Extinction, No. So I will, yeah, we just think about why speciation that behind in the tropics due to climatic factors. So, hypothesis a here. So one of the reasons might be that there is low seasonality. So as we talked about in the previous lecture when we're thinking about the earth, going around the Sun honest light with its access on a slight tilt in the northern and southern kind of as you go towards the poles and northern and southern regions of the world. They switch from Summer to Winter as they are facing the Sun and then facing away from the Sun drawing us, right? Orbit, that's the way I look for around the Sun. However, in the tropics, they tend to be almost always facing the sun really. So there is low much lower seasonality because there isn't that Kilt to and from towards and away from the Sun. So this potentially result in species with highly restricted environment appliances and limited dispersal ability, across environmental barriers leading to population, fragmentation and speciation. So this is saying is that if there is low seasonality then species don't need to deal with changes in conditions there. For they don't need to, don't need to have wide environments tolerances so they just they can adapt to a very narrow range of conditions and still survive because we don't have that seasonality. The challenges them every year, so if they don't have large environment stances, they maybe they can't cross barriers. So they can't cross mountain ranges, they can't cross other things like that. So that means that you kind of get smaller ranges of species that live. And especially if it's kind of small ranges and narrow environmental Sciences, which means that you can get speciation, because they inhabit smaller areas. So, more species can fill the area. Basically, so that's one. The other one is to think about insulation. So, again, tropical either greater conservation of solar energy as we talked about in the previous lecture, which promotes increased productivity, this might lead to larger Bible population sizes of Specialists and higher plant diversity, and this is kind of similar again, but It's thinking about how with greater amounts of solar energy available. Plants can be more productive and in that case, they can support larger population sizes and and may be able to diversify more quickly. So, so it kind of links to previous one a little bit. So if plants are able to exist in more dense, the population size is a larger over time. There's a larger chance Of speciation occurring, just due to the fact that there's, there's more individuals, there's more genetic diversity rising, and they may be more speciation. And then, if there's And these two don't necessarily occur independently. So, if there's large population, size of plants that can have marrow environmental tolerances, and they can evolve to be Specialists to certain to certain environmental conditions and we can get this kind of diversification and speciation on a relatively small, spatial scale and it says higher plant, diversity. That in turn could support more Specialists herbivores are which and then specialist carnivores. So if we Much more diversity at the bottom of the ladder with plants then that can lead to higher diversity at higher trophic levels, which are dependent on those Specialists. So we kind of get very high specialization so animals and plants that have narrow environmental sciences and also special quite specialist diets and things like that. So this leads to quite a few hypotheses that we could test. I won't go into lots and lots of them here because it's quite a lot to cover in this lecture. So if we think that this last one here or actually both of these, You'd Stay A that potentially, we could predict that tropical species would on average have smaller ranges than similar temperate and polar species. All that they would be more tropical species. We have parallel environmental Collins's that temperate and polar species, or that there would be more specialist rather than General species in the tropics. So, this is my parties that would arise from this. And that's just worth highlighting that with a lot of these hypotheses for why we predict Go for with a lot of hypotheses for why that for the processor that given rise to particular ecological patterns. And so this is kind of theoretical ecology. Why we see the patterns we do you want to be able to outline hypotheses that would allow you to test whether or not those processes actually gave rise to the patterns? We see. So it's always worth thinking, oh, well, this person has suggested that this process has given rise to this pattern. Okay. Well, what particular aspects of that process would would we be able to test? That would allow us to say that that's true. And a different process has not given rise to that pattern. So does your hypothesis give it does your proposed method process, give rise to testable hypotheses and this case, it does. And we're not gonna look too much in detail at that. Now, we're going to look instead about whether or not there's evidence from different tax that are whether or not species Jason 10 seems to be higher in the tropics. So what we're looking at here is fossil data from three different groups. So we have formant, foraminifera, Nano planets, and radiolaria, and these are very small organisms that live in there. See So fos on the y-axis, here is first occurrences. So this is the first time that a particular species has occurred in the fossil record. And on the x-axis, we have latitude. So where was that particular Rock in that period of time? What last year was at? So we don't have to just look at what I asked you to stop now. Because if that rock is from a hundred million years ago, then we need to be able to tell where it was at the time that it was formed, but we can do that with geological methods and we can look at how many first occurrences in the fossil record. There are Different latitudes. So here we can see that for form and infra it seems like speciation may be higher in the tropics because we get more much more first occurrences at low, latitudes at these tropical latitudes, however, for the other two groups, it's not clear. So if Nana Plankton, it doesn't seem that there are, there are many more verse occurrences. And for radiolaria, there's many more first occurrences at the Barre. Hi, Southern latitude. So in this kind of Southern polar regions, so it seems to vary by taxa but it's just evidence that that maybe this is important for some tax or not others. So again as we always talk about so they like when we talk about the effects of climate change and things like that. So it's not only that certain species may and there's this taxonomic variation in the importance of different things. So for this, there may be tax on it variation in the importance of different processes, giving rise to the The biodiversity gradients, so it may not be the same explanation for all taxa. Finally, we're going to get onto historical here. So again, this is a reminder of the three hypotheses. So, are the historical factors that have either led to higher speciation rates and Tropics or lower Extinction rate or both? so a way of doing this again, we can look at the fossil record I'm to provide evidence whether or not these historical other historical factors at play with that affect the speciation and the extinction rates. So, the first thing to look at is whether or not this latitudinal, Brett biodiversity gradient that we see today, so higher the rest in the tropics lower in the, polar regions has been present all through the history of the world and we can look at that. Using the fossil record, we can estimate it. Using fossil record at least so this shows in the late Cretaceous and this is the kind of layout of the continents of the world and it contagious, there doesn't seem to be any relationship between diversity and latitude gradient. So here, looking at this diagram, we have Take into account, the land area that was present to each laughs dude, as well to see whether or not there. That's why that's why we see the because if we have more land than is likely more species due to our species every relationships, but we can see here that the red line, which indicates relative diversity. So, towards the right is high diversity for the left is lower. There isn't a big peak at the tropics, and a big decline in the polar regions and instead, The amount of diversity, we see seems to be kind of largely controlled by the land area that there is available. So that's why it seems to be a Peak at around 50 or 60 degrees north. So if there hasn't been Allatoona gradient through this, the same last June of biodiversity gradient Through Time. What period of time has it occurred in? Because it is definitely present now. So has it been a has occurred in the past? So again, we can plot not just looking at one period in time where we looked at the late, Cretaceous there instead, which is roughly 72, 66 million years ago. Instead, we can look at the diversity. Gradient through a longer period of time, so Here we have the red line is temperature. So lower is low temperature. Higher is obviously higher the blue shading show. I have ice house world so times in the Earth's history where Isis has been present. So it in a nice Health world. Now, technically because we have ice sheets of the poles and then we can compare this with the times where we had either, pull with declines in species, diversity, which is what we see now. So there's lower species, diversity in the polls or whether or not they were high diversity in pain. In temperate regions. So here the Open Circles show times of high diversity in paleo temperate, regions and solid circles your pole with declining species diversity. Which again, is what we see today. As we can see, we tend to get this poll of declining species diversity in Icehouse World period. So times in the Earth's history, where that's been Ice sheet present at the polls. So maybe, then the reason we see this last unit, biodiversity gradient, has something to do with temperature, and/or, ice, sheets, and when we think about this. So, that's so we're kind of, we now have a potential mechanism, but how does that translate into either? Different speciation rates or different Extinction rates? So again, we will come back to this idea of Niche conservatism. So again, this is Chandni, introductory lectures. This means that basically it's quite difficult for organisms to evolve to fill a different Niche. It can take a long time so that means that the ancestral niche of a clade. So a group of species that contains all the descendants of a particular species. Can the term, the regions and habitats to, which the creek can spread and those, which will persist in the face of environmental change. So we looked at Tropical frogs in one of the previous examples. So, if tropical frogs arise in the It may be difficult for them to evolve to inhabit the temperate regions and the polar regions. So, they're kind of concern may be constrained by that evolutionary history to live in tropical regions. So if this is true, then disturbance is a non tropical regions could lead to extinctions while speciation continued in the tropics and one of these disturbances. Maybe I sheets tropical species come up move to inhabit non-tropical reasons easily. And then again, this gives rise to some hypotheses. But basically, what we think about here is so there's now they're going to the interplay between this biological or ecological process of Niche conservatism and the the formation and spread of ice sheets. So if There's no ice around the world then speciation can occur. Similarly, everywhere speciation Extinction, occurs, summary everywhere. So we don't get the latitude and biodiversity gradient as we see here. How's a nice house? World's eyes will always kind of be more present in the polls because of that seasonal variation, whereas, because they Tropics receives more insulation, more solar radiation, there's less chance of being nice to the polls. So that means that as you go through these Ice House, world's speciation and kind of can carry on in the tropics, but lots of species will go extinct and speciation and normal ecological processes and evolution process. This may be disturbed, weather is where the land is covered in ice because it isn't that normal threshold buyer and available for the species to live in. So, in the tropics in his hypothetical situation, we have normal speciation, and normal things happening, whereas, in their areas are covered by Ice, everything's been Disturbed. And then even in times when ice retreats to the polar regions, it may be that the tropical species that evolved in the tropics. They can't move to inhabit on tropical regions or it takes a really long period of time. So, say in today's world, we have ice sheets, but they are restricted. Mainly to the polar regions. So why then are the temperate regions? Not More species, rich or species rich in the tropics. Why do we see lower diversity in the temperate regions? Because that the ones that aren't covered in ice or times like the polar regions can be well, maybe it's because the tropical species haven't had a kind of stuck there and they haven't had the evolutionary time to be able to diversify and spread into those non tropical regions. So, therefore, the interplay of ice sheets and Niche. Conservatism could have given rise to this latitudinal biodiversity gradient. And so if we think about the last unit bird biodiversity gradient, we see in the modern day, we want to have a look at these different time periods. So the pliocene between 5.4 and 2.4 million years ago, was a time of global cooling following the much warmer miocene, which came before it. And during this period we saw spread of grasslands and savannas. So the kind of buy-in that we see in the bottom here. And so that led to potentially if they were tropical forests around much of the world. Before that, in the warm I seen as these kind of grasslands potentially spread down from higher latitudes, a lot of those tropical forest species may be the gone extinct orbit, move towards the poles and we get this Guild of long-legged grazes that inhabit. These grasslands but potentially of more significant is the spread of ice sheets from the pole downwards which would have Extinction of lots of the more warm adapted species, that would have been present in those latitudes beforehand. But as towards the end of the pliocene, as these ice sheets started to form a started to move down towards the lower latitudes. That's where we start to get internal these Icehouse worlds. And then we move from the placing of the pleistocene. So 1.8 million to ten thousand years ago where we have periods of glaciation but on the glacial cycles of the next slide. So during this time as there was much more ice around down too much. Lower latitudes. Than we see today, many Northern species occurred far to the south of the present distribution during glacial period as the I spread towards the equator from the poles. And many species would have had to have migrated down towards the lower, latitudes towards the poles to escape that ice. And one size retreated, they move north as against, we kind of get this latitudinal toing and froing of lots of these species and this is looking at how this been glacial Cycles during the pleistocene. There were glacial Cycles during the pleistocene. So we have periods of very low CO2 levels when they would have been high ice levels and then periods of higher CO2 levels when we get called interglacials. So in these interglacials it doesn't mean that the ice definitely necessarily disappeared. All together especially with look today and so we're currently in this integration, but we do have ice present, we may have massive humans, may have massive disturbance. So if we see right at the right hand side of this graph, we see a sharp spike in CO2 levels, right? At the right hand side, as you move into present day. So that's caused by anthropogenic, CO2, emissions and pollution, and greenhouse gas emissions, which is leading to climate change. But this just shows how we've had these kind of glacial Cycles during the pleistocene, which would have had, I spreading to the polls in a way, which would in constantly disturbing the species are present in the polar and temperate regions as they were came into contact with these ice sheets. So when this is paired with a niche, conservatism it, maybe this is why we see the latitudinal, biodiversity gradient that we see today because the tropics never been covered by ice sheets in this time period. So kind of speciation and normal ecological, Evolution Dynamics can continue in the tropics while in temperate and polar regions. Those areas of been continually disturbed by these glacial Cycles, so many species name on it stinks. It's and also speciation may have been Disturbed. So that might be, we see the last funeral, biodiversity gradient. We see today due to this, the kind of interplay between this climatic history and also Niche conservatism, which has meant that the tropical species can't quickly. Spread out the tropics to inhabit those different climates and environments. You get in the temperate and polar regions. It's just looking at how I started climate change. Has also affected other things, so it's not just kind of gently as not, just temperature and rainfall. And so we look at southeast Asia. During the pliocene pleistocene, the landmaster climate of Southeast Asia was dramatically influenced by changing sea levels. So as you get greater ice coverage, we get more and more water. It's locked up in those ice sheets, so we get drops of your Falls and sea levels also as the Earth. You also get water becomes more dense. So that the also factors into Hawaii, sea levels, drop when the earth becomes cooler. As we can see here, we're kind of moving down into the this one yet. So we go back to the start here since the current Coastline. And this shows the depth of the 30-meter Contour around land masses. So if sea level drop by 30 meters, then this is how we would see where we would see the coast of the drop by 50 meters. This way, we see it. So it dropped by 75 the drop by 120. So in these Ice House periods, In glacial periods are even those more ice around sea level dropped. It allowed more migration of species around southeast Asia because there was land bridges between a lot of the islands here so that just may also influence why we see this species we do today so the biogeography of species in Southeast Asia. And this environment, major environmental change is expected to have major effects on the spatial distribution and genetic diversity in many taxa. So during interglacial periods here we have all these conditions and then in glacial periods we have low sea levels, reduce levels of precipitation. So that is why there may have been as so this this kind of history may have had a major impact on these biogeography of species that we see in Southeast Asia today and also the genetic diversity So, it's also worth thinking, a little bit about how this interplays, with the the, by jog up here Wallace, which we talked about. So, The Wallace Line goes, it's characterized by a very, very deep bit of ocean. So, even during the kind of periods when sea level would have a very low there, may not have been a land bridge in many places or any at all across that Wallace Line. So that may be why there is such a stark difference between the Flora and fauna on one side of The Wallace Line, and the Flora and Fauna on the other. Because even during these glacial periods, when sea level was much lower, it may have been that trash or species, still couldn't cross that very deep bit of water because there wasn't a language available, so that might be one reasons why? So again, that's thinking about the interplay between evolutionary history between climactic history and how that's giving rise to the five geography of species that we see today. I'm just it's just looking at the the major biomes are present here in the last glacial maximum so around 22,000 years ago, I think so this is what kind of the biomes that would have been present in that time look like and so we just do deal, we do still so you have rainforests in some areas but it's a bit more concentrated. So this is just kind of you an idea of how biomes would have been changing throughout this period through these glacial and interglacial periods. So that's the kind of evidence for why? How there could have been. How historical factors could have given rise to the the last unit biodiversity gradient and how the history of glacial interglacial periods over the recent relatively recent history of the world may have given rise to greater greater diversity in the tropics. However we have to think about About kind of the potential objections to this as well. So this is one of them so that have their kind of really been climactically stable regions in the tropics over this time period. So yes, they may not be affected by ice but what other of climatic shifts meant that they wouldn't have been stable enough for that to make sense for the kind of speciation and normal evolutionary relationships to continue as they would normally in non glacial periods. This work which again sized in the notes for this. I'd so the locations of climactic we stable regions are likely to vary considerably across and within Millennia. So have they even been long-term climate refugia so as we can see here, there's been the haven't really been many places that have been spent a lot of time in extremely stable conditions, This date, this diagram here just shows the data from from that paper, which looks at how stable certain readers have been over over long periods of the Earth's history. Well, the kind of more recent periods of the Earth's history So the different areas so that's been split into coming up with a stability score. If you want the kind of detailed how this is done, please read the paper and they split regions into eight kind of stability classes where the blue regions of the a diagram here are the most stable. And then red are the least stable. So we do get kind of a gradient where polar regions have been, sorry. Tropical regions have been more stable than polar regions. However, if we look at D here, which shows the amount of time that the two most stable classes, the areas that have been put in these two most stable classes, the amount of time that they've spent in these stable conditions or very stable conditions. So, Sorry. So the p25 class areas the kind of second most stable areas and we're looking here, how long the p25 regions class have been how much time they spent in in Dairy in stable condition. Sorry with green for land and blue phocion. So we can see that actually even for this kind of stable region, not quite a loss of that that Massive land hasn't actually spent that much time in stable, unstable conditions. So there's some areas that are spent, you know, over 90%, but quite a lot of that density curve. That is is kind of below 50%. And then, if we think about the very stable conditions, actually very small amounts of land. And the ocean have had spent a lot of time in very stable conditions. So, so again, I recommend the pay-per-view a bit more interested. Now, this has been calculating how this works, but basically also saying that they're actually even though the polar are the proper region. Sorry, are relatively more stable? It may be that they haven't actually been that stable over this period of time. So can climate refugia. Does that make sense as an explanation for why we see that higher diversity in the tropics? Because even the most stable region of the world aren't really that stable over this period of time. And it may be that there still would have been disturbances to the to the life exists in those regions. Finally, we're going to look at the the last hypothesis where there's going to be speciation where speciation rates are higher in the tropics but also Extinction rates are lower there as well. And so, this kind of is a hypothesis that both of these processes have been driving this pattern. And so again, we're going to have a think about what hypotheses that that theory would give rise to. So can we think about the hypotheses? We could test that, that would give evidence for that theory at the expense of others. So, so are there, are there certain things that the out of the tropics Theory would give rise to that. The other two wouldn't on a certain patterns. We could observe in modern diversity. That could only be Explained if the out of the tropics theory was the real one. So here we thinking about phylogenies and so one of the hypotheses is that the shape of phylogeny use would would look a particular way if there was the this out of the tropics model. So if we were thinking about just topics' cradle or Tropics as Museum the following News. If we think about these conservatism and these are the kind of the two diagrams at the top of this, that these three are the other phylogenies that we'd expect. So, wouldn't see much Crossing of of tropical species out into the extra Tropics out into the temperate and polar regions. We would do follow Jesus species mainly maintained within their own zones. However, if we think about the out of the Tropic model, The case will be slightly different. Look at the top one here. Tropics are is cradle. There's going to be greater speciation in the tropics, and that's what we see here. So, there's more species being formed in the tropics and there are in the other to think about the middle. One speciation rate to the same in all three places. So we're getting lots of branches. However, lots more, the species go extinct in the extra tropical regions. In our out of the tropics model. We get greater, speciation the tropics, but we also get less Extinction. And what this means is that if there is less Extinction, yes, we can still have knees conservatism and Take longer for tropical species through out of the tropics because they are ancestors and adapted to those temperate or polar conditions, but because there is less Extinction over a long period of time. Certain branches of that follow, G will move at the tropics into the extra tropical regions. So this is one of these hypotheses that we can test now and we can think about if this is true, then it provides evidence for this out of the tropics model. So this paper again, site in the notes for the slide and also want to recommend reading Owing to this lecture. So this theory is supported by an analysis of Bibles. These are Marine Shellfish, a certain class of marine, shellfish over the past 11 million years As we can see for this for this taxer, there's evidence that there's been both higher speciation and lower Extinction in the tropics. So the top two graphs here, look at species. That had their first currents in the pleistocene. Middle is first occurrence in the pliocene and the bottom is the first occurrence in the late miocene and so a c and e. So, the grass on the left hand side, show the number of First occurrences in tropical and extra-tropical regions. So, in all three time periods, has been more first occurrences of taxpayer in the tropical regions and the extra tropical regions. So, this would suggest that more speciation is occurring in the tropics than in the extra pickle regions. So, This is gives evidence for the speciation part of the out of the tropics hypothesis, but as well as that. Look at that other graphs on the right hand side here. So this looks absolutely modern Pollard limits of taxa, with tropical Origins. So, this again, looks at taxes that formed in their place in the top players in the middle and my team the bottom. And, as we can see lots of these species, that occur that originated in the tropics now. Have Quite High latitude modern limits. So lots of it seems like lots of tax that have moved been able to move out of the tropics into the extra Tropics. So this provides evidence for the second bit of our out of the tropics theory that Extinction rates may have been low in the tropics, so the use tax have been able to get over there. Then use conservatism Challenge. And I've been able to move into the extra tropics So this is support this analysis, supports the out of the, tropics hypothesis, at least for bivalves. So, again, as I said, quite early on the lecture, it isn't necessarily the case that there's going to be one explanation for this pattern seeing across all taxa. It may be that different tags that have been affected by different processes. And there's different explanations or different explanations are more or less important for different. Taxa Okay, so just in conclusion, tropical rainforest, some of the oldest ecosystems. They have persisted in the equatorial regions, even when there have been periods of cooling, that would have reduced or fragment of these horizontal extreme. Last Jude's this long stem term stability, has likely played an important role in the high diversity of rainforest and contributed to the LBJ LBG sorry and high speciation rates are also likely to be important. So just a conclusion that it seems like there's a combination factors at Play. They are, these are the three routes papers that I recommend this reading. This is the end of this tropical diversity, lecture course, for me. So thanks so much for listening and I hope it's been useful.