[Music] Hello and welcome to the Animal Behavior Podcast. I'm Matthew Zippel. In March of this year, I visited the Kalahari Research Center in the southern portion of the Kalahari Desert in South Africa. And this research station is the home of a number of long-term projects, including the famous Kalahari Miracat project. And if you've ever seen an image or a video of a wild Miracat or read about a paper about wild Miracats, the Miracat project was almost certainly the source. And on my trip, I got to spend quite a lot of time with two incredible researchers, Tim Kluttenbrock, who founded the Miracat project, and Jenny Tong, the director of the Max Flunk Institute for Evolutionary Anthropology, who I was traveling with. And it seemed ludicrous to have these two great minds at my disposal and not record conversations with them. So we've got a two-part series for you that we're going to call "Dispatches from the Field," where I interviewed first Tim and then Jenny. Today, you'll get the first part of that series, my conversation with Tim. Tim Kluttenbrock is perhaps the foremost zoologist and memologist of his generation. He's done groundbreaking work in numerous long-term studies that he's established, most famously the Red Deer on the Isle of Rum, and the Soy Sheep on St. Kilda, as well as the Miracats of the Kalahari. And while his fieldwork has been focused on studying individuals and their lives within societies, thanks to his encyclopedic knowledge of mammals' natural history, he also has an amazing ability to make connections and find general principles across species. It was an incredible joy to spend time with him, and I hope that comes through in our conversation. Two quick notes. First, we recorded this conversation on location, walking around the Kalahari with a group of Miracats. So that means that you'll hear a speaking and slightly hushed tones, so it's not to disturb the animals. And you'll also hear some of the sounds of the Kalahari in the background. Birds, as well as Miracats, and an occasional gust of wind. As always, you can let us know what you think of this episode at
[email protected]. Here's my conversation with Tim Plutton-Brock. I think just to start, if you just maybe describe where we are, and why we're studying Miracats, how you decided that this was going to be the field type for you. I worked on Red Deer for the first half of my career, and I worked basically on under variety of religion assistance. And a lot of that work was concerned with contrast between the sexist, the operation of sexual selection, the consequences of sexual selection. I felt by the sort of 1990s, I'd been going on these for 25 years, and I felt it was time per change. I was interested in polyandry, and I thought it'd be interesting to look at polyandras birds who were row reversed. And so I investigated initially the possibility of working on buttonquail. And my thought was that since people are used to breeding quails in captivity, and the way you do that is well known, I thought it'd be quite feasible to bring buttonquail into captivity. So I came out South Africa because there were people who were breeding buttonquail there. At the same time, I thought it would be quite interesting to look at complicated breeders, because there are a lot of wonderful studies of complicated breeding in birds. And there had been early studies on mongooses, but that stage no one was working very much on mongooses. And so I thought I'd go and see mucats in the Calahari, and I got in touch with John Skinner at Pretoria, and he arranged for someone to take me up to the Calahari Krimsburg Park. And there'd been a French film team there who had a bit-to-aided group. And so we found the habituated mucat group, and it was just a big plain sand with various holes in it, and they were prospecting for insects. And you would see what each animal in the group was doing, and they were very well habituated, so you could walk in the middle of the group just like we're walking in the middle of this group now. And I thought wow, this is it. This is so good. I almost do what the boom people are doing. And we started in mucats in the Calahari Krimsburg Park in 93. They habituated quite easily because the park's very open, and when a group ran away from you, they couldn't really get out of sight, and so you could drive a vehicle after the park it a couple hundred meters from them, and then gradually press them. And so we got to be able to walk with groups quite easily. And at the same time we started a study actually here outside the park or in Ranch Land. Here you couldn't pursue the groups as you could in the park with a vehicle. We initially tried to induce them to come to us and to allow themselves to be weighed, using everything. We tried to find food that they would eat, so we tried everything. We tried peanut butter, we tried various sorts of cooked meats. We knew that they commonly had pluvas eggs in the field. And I thought that they might like hens eggs. So we tried that, we broke hens eggs that weren't interested. And it even got to the stage where sometimes we tried actually breaking a hens egg on their heads so they did round down their face, we forced them to lick it off, but they dislike that too. And then one very hot day I tried to get the dominant female to eat hens egg by breaking a hens egg. I'm not actually breaking it on top of her head but breaking it above her head. So the yoke and the white grand-owner face. And she was disinterested, but it sort of slipped off and slipped on to the side of the burrow. And it was a very hot day and it actually cooked in the sun. So one had something looking very like a fried egg beside. And two subordinates came back to a burrow and they sniffed this and they immediately started eating with gusts there. And then the other members of the group joined them and they ate it. So we discovered that we had, we finally had the thing that was attract them, which was hot boiled egg and we discovered they loved hot boiled egg. So we moved from one group to two groups to four groups. So for the last 30 years we've normally had between six and twelve groups. The moment we've got 14. So this one that we're with at the moment, which is Nasi La, is the only one, only group that has kept going over that whole period. It's now a sizable group, 29 individuals in the group at the moment, one dominant female with one dominant male. And then a pile of non-breeding helpers, the subordinate females, sometimes try and breed commonly with wandering males from other groups because they are normally the offspring of the breeding pair and like lots of mammals, they avoid females, avoid breeding with individuals that are familiar to them. Commonly you get asked in here's what's the most important thing we've discovered about mere cats. And I always said the most important thing we've discovered about mere cats is they love hot boiled egg because that opened the way to weighing them regularly. My wife, Daftler, who's biologist, appointed at the day, okay we didn't need to stop just way them once a day. We could weigh them several times down and measure their rate of weight gain. So we developed a regime where we weigh when we're working with a group. We weigh everyone when they get up if we can and we weigh them again. We've had lunch time to get the morning weight gain and we weigh them in the evening before they get good to get the afternoon weight gain. We weigh them the following morning which gives us a weight loss overnight so we think of that as a weight sequence. We've now been doing this with mere cats for 30 years and we've got large numbers like thousands of individuals where we've weighed them at least once a week, commonly multiple times in a day over one or two days per week for large numbers of individuals from birth to death. So we've got these measures of weight gain and weight and condition for thousands of individuals throughout the whole lifespan and we've found that these differences in weight gain and weight tell us an awful lot about how they do previously about their survival, about their reproductive success. We are able to compare dominoes and subordinates and so on. So I think the one of the unusual aspects of the study is that we have this very very fine great measures of growth and weight. So cooperative breeding was part of the motivation for being here. So cooperative breeding was definitely part of the motivation for breeding here.
The cognitive breeding is in a sense a similar system to social insects, with its most similar really to some of the what we call primitively social insects, but Darwin recognized that the whole issue of the presence of helpers or workers that didn't breed and spent their lives helping other individuals breed was a real challenge to the idea of natural selection and how natural selection worked. That has led to considerable interest in corpse of breeding, particularly in birds. And so when we start off with this I was well aware of that interest I was well aware of the bird studies, but at that stage there have been few studies of mammalian corpse of breeders which were really inconsidual details. So for example, while dogs some wonderful studies the kind of data that one can collect on wild dogs is I mean they're difficult animals to work with. I realised how we worked on primates previously. I realised there's some fantastic opportunities that when you could work with animals in group you could walk around in, where you could access to the pups easily, where you could weigh animals and that way you could collect samples of skin or blood or fur or feces or urine sort of as well. In contrast to lions and hyenas who could walk among them without the wanting, without the risk of the theletio they have a comparatively short lifespan so that very few animals live more than 10 years and I think there are very few mammals which provide this whole range of opportunities in the market. So I think that we've been immensely lucky and the sense that as we've pressed them, as we've got them more and more habituated, we've been able to do more and more with them. So I want to talk kind of about some of the specific cooperative behaviours. So can you just describe the pup feeding behaviour? One of the unusual aspects of Mierkats is that they are sort of one cooperative behaviour like feeding but the next they do a pile of different things. They put up sentinels. So there's commonly one individual who goes to a raised position, stops feeding and watches for predation, giving a variety of alarm calls when they see predators. They collaboratively dig out bolt holes across their range within a group has probably around 1000 bolt holes in their range and we believe that they actually have a good map, a memory map of where they all are. They don't forage cooperatively so everyone forages independently so there's no cooperative foraging there but they defend each other and when danger comes they'll attack. And then there are a slew of cooperative activities associated with breeding, when pup's are born at a burrow, several females will lactate for them, not just the dominant female, but the dominant female does most of the work. Helpers of both sexes babysit and babysit, the each group has a range of the region of 3 to 5 square kilometres and they commonly travel several kilometres away from the breeding burrow during the course of the day. So the breeding burrow, it's highly risky to leave it unattended and one individual or two individuals stays at the burrow and protects it against any intruders, sometimes fighting for death because neighboring groups will attack breeding areas and try and kill the pup, neighboring Mekak groups will try and do that. And then they feed pups for the first three to four months of life. So individuals, they don't feed them at the burrow when they're around three weeks old, pups start to forage with a group when they're really quite small and they're fed by their parents and by the helpers giving away somewhere between 30 and 40% of all the food they find. The helper would go of sniffing for food in different places, digging for food, finding, finding grubs or small scorpions or, or in some cases, small vertebrates. And they'll come and then disable these and then bring them to a pup. You can hear them, or you hear begging all the time, "Wa-wawa-wawa, I'm begging for food." The begging stimulates the helpers to feed them. And as pups get older, they start to be able to feed themselves and they stop begging and the adult starts gradually sees feeding them. But all the individual feed, females feed a bit more than males do and the females adjust what they do rather more sensitively to the aged stages of pups. And despite those high level of cooperation, at some point that turns between the dominant female and others to complete dissolution of that cooperation. It's not exact at some point. It's always continuation. So the dominant female doesn't tolerate other females breeding. And if other females become pregnant, she communicates with the most of the group, flat out attacking them. I mean, if they say they do eventually leave, but she commonly choose them at the base of their tail initially and then finally loses her temper in the sense that it ends up with an all-out attack. We've never seen females leave voluntarily. Females are always evicted. And if you look at the group, there are no females older than about three, three and a half years. So all females get evicted eventually by the dominant female unless the dominant female dies. In evolutionary terms, what you've got is a dominant female who meets with a dominant male, usually with the same male, because he's almost always an immigrant. In some cases, when the dominant male dies, you get another male assuming dominance, you get a natal male assuming dominance. But the dominant female doesn't breed with him on the rest of the other group members don't breed with neither. They share a very strong avoidance of breeding with familiar individuals. And males leave voluntarily, normally, between the ages of about two and four. And they initially start visiting other groups trying to mate with subordinate females who commonly show quite a lot of interest in them, but the group chases them away and kills them sometimes if it catches them. So that roving males have to mate surreptitiously with willing subordinate females behind bushes. But if the subordinate females get pregnant, start to breed, they're quite likely to get evicted by the dominant female. They then always, and subordinate females do sometimes produce letters, but they share much lower reproductions of success. The reasons why dominant females kick out pregnant daughters is the pregnant females killed pups. So if you're a dominant female and you've got pregnant daughters in the group, those pregnant daughters will kill any pups you produce. And vice versa, if pregnant daughters produce pups before the dominant female, the dominant female will kill their pups. And then you'll have to interest the dominant female to remove pregnant daughters from the group. So I think I'm probably biased. When I think about social groups, I think of multiple kin groups, you know, living together. Can you walk us through kind of the evolutionary logic of how strategies are going to differ in groups where you have several kin groups living together versus everyone being more closely related? Yeah, well, you come from a booboon back then. So you're thinking in terms of multiple females breeding in a group and multiple males breeding with them. When the outcomes of a situation like that, we've got multiple breeding, male, multiple breeding females. The individuals in the group are not very closely related to each other. So the average coefficient of relatedness in the sort of animal that you work with booboons is commonly as not much higher than 0.1. Whereas here, many of these animals are full ships. The male and the female can last and breed together for several years, including the immigrant male who is unrelated to everyone apart from his actually own children. The average degree of relatedness in the male care groups is about 0.4. So that's, I mean, if you think of it as an average, that's something like four times the average degree of relatedness in the booboon. So,
There's territorial system, lots of groups occupying different ranges throughout the habitat, so the whole habitat is saturated. So dispersing from group anyway is always extremely risky, lots of individuals die on a dispersal age. So females' best chance of breeding is to inherit from our mother. So in a sense what you've got here is a lot of subordinate females, but subordinate females when a sense if you put it bluntly they're all sitting here hoping something will come and eat their mother. And that's their best chance of breeding. If you can get to the dominant position in a meerkat group and hold on to it you've immediately hit the reproductive jackpot. So our most successful females have produced somewhere around 80 or 90 babies. They've reared 80 or 90 babies during a lifespan which compared to the top level of a baboon or almost any primate that you can think of is way, way higher, an order of magnitude. So one of the results is that the females compete like fury for the dominant position. Females are more aggressive than males. In many of the animals with breeding systems this kind females have unusually high testosterone levels. Females are fierce animals holding on to the boners, the reproductive boners that the dominance gives them. And the dominant female, she's dominant to all other members of the group, normally including males. But you don't see the same sort of level of dominance here. You don't have a rigid and very obvious hierarchy like you're doing baboons, really in either sex. Outside of the dominant, everyone else is pretty much the same. Everyone else is much the same. And even among those you don't really get a big difference. It's this high variance in relatedness and these big differences between king groups that give you in high-innocent baboons the sort of coalition-ary support that you see. And it generates a situation where an awful lot depends on support from your kin. In mere cats an awful lot depends on the size of the group. And a single female can seldom be real perhaps successfully on my row. And for perhaps to grow and have a good chance of survival, we want to re-simply high ratio of help as to perhaps. So it's similar principles but at a different organizational level. With you you've got in a sense a super group, with lots of subgroup, competing subgroups in it. Here we have separate groups each in a separate territory with group size maturing a lot. At the surface level, cooperative breeding, socially insects feel very similar. Yes. How are they not similar? Well, individuals source burn for life, there's obligates to reality. I think one of the important things is just the size of the colonies so that if you have a colony of those tens or hundreds of thousands, your individual chances of breeding are miniscule. Here everyone has a chance of breeding. When you move to socially insects, the chance for an individual being able to get into a breeding niche unless they're costeted and set up for that and the competition is reduced by the workers. And so unless they're sort of prepared for the role, then the chances of getting there are vanishingly small. So for those animals, there's no, even if you're a worker, which is not total sterile, your individual chances are very, very low. So there should be more conflict in mammals? There should be as much more conflict in mammals. They do share a number of very important characteristics. But I think the comparison I draw is if you said, "Okay, just think of insects as out is generally." I'm just too ignorant. I don't know enough about the whole breadth of social insects or the whole breadth of insects really to know what the right comparison to make is with mammals. I know enough about birds and mammals to have an idea of what I'd like to compare with birds between birds and mammals. But as one moves to all invertebrates, what would one say about sociality among invertebrates? Which range from all organisms? It starts to become very vague and it's difficult to identify the questions that one really wants to answer and so make the comparisons that you want to make without ignoring major differences in constraint. So I want to turn back a bit to the site here and the approach to doing science. So can you describe the volunteer structure and the kind of large scale data collection approach you've taken? Because each group has a range of about three to, in some cases, 10 square kilometers, it's difficult to move from group to group. I've always wanted to have a statistically adequate sample group. I've wanted to have at least six groups, ideally around a dozen groups because one of the restrictions of many studies is that they're done on a single group or one or two or three groups. And one of the consequences of that is we wanted to have reasonable number of people because it's difficult for one person to work with several groups on the same day because they widely dispersed. So we opened up and we started taking on volunteers who would come for a year, usually we selling take on people. On the Miacat study, that's not prepared to come for a year. So we moved to a bigger group of people working on them, collecting routine data on life history as foraging and on different aspects. But we certainly try and cover the growth and the weight characteristics and life histories of all the individuals in all the group to work with. And that led us to a situation where we commonly now have 10 or even 12 volunteers coming for a year at a time. They get a fantastic grounding. They obviously know how to do field work back to front, but they also commonly get involved in data analysis. They're very used to putting data into our enormous database and they become adept at putting it out too and quite a number of them have gone on to do PhDs elsewhere and so on because that the background they get here fits them and gives them exactly the sort of background that anyone doing another sort of long-term study would like. So the deal is that we have put up enough accommodations that everyone has a separate room. We feed community in the evening. We basically cover the costs of all the volunteers while they were here working with us and we make a contribution to their airfare to come out if they're coming from another country. But what it means is that people can come and work as a volunteer here without saving up a lot of money to do it. And I think that's you're going to ask people who are going to come to work for you for a year like this. I think you need to give them reasonable accommodation and I think you need to cover most their costs. And in total how many volunteers have you hosted? Somewhere around 300, 400. A surprising number of the most still in academics, quite a number of gone on into sort of conservation related activities of some kind. Has that felt like an important legacy to you? No, I don't really think of legacy. Well, maybe I see an important thing to have done. Everyone tends to think that you use the term legacy. I don't really think about legacies in that way. But I think it's a very useful thing to have done. I think we train lots of volunteers. I played a role in setting up the tropical biology association which also trains people. And I've had, I don't know, somewhere between 50 and 60 graduate students. Not all of me are cats, I should say. What do you see as the kind of the most important benefits of long term studies? If you're going to boil it down, why do long term studies instead of cross-sectional? Then you can do more species or in more places, more population. Well, I think if you just do cross-sectional from scratch, you don't know where you're at. I mean, here, or ain't it with any of these systems? You need to know how old individuals are. You obviously need to know what sex they are. You need to know whether they're natal or whether they're immigrants. So in terms of social organization, you absolutely need that sort of information. And you really need more than that. You need to know the background of individuals. You need to know if they're breeding how long they've been breeding for. As that develops, I think they're huge.
benefit in knowing the famine history of individuals, you realize that events at one of the stages of a lifespan actually have cascading effects that operate throughout the lifespan. Even an adequate description of social organization requires a pile of longitudinal information. And once you have that kind of longitudinal information, you can ask a huge range of important biological questions concerning breeding success and survival, as well as individual social relationships that are dependent on knowing the history of individuals. Actually, a very small number of detailed long-term studies in birds, in the blue tits, great tits, song sparrows, and then more recently a slew of cooperative breeders have had a huge impact, to set the basis for the field. And I think the same is true on mammals, that actually the important results and the important concepts that have developed over the last 50 years, a very high proportion of those have come from a handful of very long-term studies of chimpanzees and baboons and elephants and spotted hyenas and wolves. One of the problems of long-term studies, that you obviously rely on whoever owns the land to let you go on and do it. And people were surprised with long-term studies, you know, they're happy to have you around for five years, but they may not want you around for 50 years. They don't anticipate that the longer you stay and work on on a population, the more valuable, the more interesting it becomes, the more specific questions you can ask. 2001, for a variety of reasons, down the South African round, went down to low levels. And I realised that, in a sense, a ludicrously amount, a small amount of money, we could buy the ranch we're working on, which is about 3,200 hectares. So it's an area of about five kilometres by five kilometres, about 25 square kilometres. A large area said we bought it. And that has been absolutely wonderful, because it's meant that we can manipulate the grazing pressure, we took off the cattle and sheep and replaced them with antelope. It's meant we could build accommodation for students and people it's meant that we've been able to build lab spaces and so on. And it obviously too, it's given us great security. So we're not reliant on anyone else to tell us what we cannot be caught in. And we've been able to exploit that so we're very independent. And I set up a not-for-profit trust in South Africa to own it. There's a group of trustees that some with South Africa and some and people from Europe. And the trust has now actually passed to Zurich and is run by Martin Mansor at the University of Zurich, run by me. And that happened for three or four years ago. And I think that's actually quite important. I think long-term studies is extremely important for whoever's running them to think about the future and to make arrangements for what's going to happen to long-term studies off their retires. So I think that that sort of transfer, all these transfer arrangements, it's a very important aspect to long-term studies. These are the sounds of destruction caused from Hurricane Hugo as it wreaked havoc throughout the Caribbean in 1989. Since then, there's been over 20 major hurricanes to hit the Caribbean and United States. Following many natural disasters, we see an increase in people experiencing cardiovascular diseases. But how does an external event like a natural disaster get under the skin, so to speak, to affect health? As a native to the Gulf Coast of Texas, I've experienced my fair share of hurricanes and have seen the toll they take on families and communities. My name is Marina Wadowich, I'm a postdoctoral fellow at Vanderbilt University. I study how environmental stressors are translated into molecular changes that impact long-term health. Ultimately, I hope to use my science to decrease disease prevalence in humans. And I do this by studying animals that are similar to humans, specifically a population of monkeys called Rhesus Maccax that live on a small island off the coast of Puerto Rico called Chiosantiago. In 2017, Hurricane Maria tore through the Caribbean, devastating Puerto Rico and Chiosantiago. Our team had been studying these monkeys before Hurricane Maria struck, so one silver lining to come from the hurricane was that it created a natural experiment where we could study the monkeys before and after the hurricane to understand how the storm may have affected their health and immune systems. I found evidence that after the hurricane, monkeys had higher levels of inflammatory gene expression, indicating that they may be experiencing long-term inflammation following the hurricane. Scientists hypothesized that experiencing severe stress might cause us to age faster, and we wanted to know if the immune systems of monkeys that experienced the hurricane showed signs of aging faster than monkeys before the storm. Using what's known as a molecular clock, we could predict animals' ages, and we found that monkeys after the hurricane had an average age acceleration of two years. Scaled up to humans, that's an age acceleration of about seven years. Now, I'm not saying that people who experience natural disasters are necessarily aging faster. We only tested this in Rhesus Maccax, but it gives us an indication of how the immune system may be affected by external stressors. Also, these results aren't all doom and gloom. Animals varied widely in their response to the hurricane. In fact, some showed slower aging afterwards, and we're working to understand the factors that influence monkeys' susceptibility or resilience to the hurricane and its aftermath. For more science like this, you can find me on Twitter at Marina Wadwich. Thanks for listening. So you mentioned retirement a couple times? I retired from my university chair five years ago, and became a director of research. So I was moved back to being a super postdoc. A postdoc who runs pretty ambitious projects. Well, a postdoc who runs pretty ambitious projects. I think it's really useful for people not to remain working on the same system throughout their lives. Every time I've moved systems, I've learned a lot, and I've worked on insects and birds and mammals. They're mostly our mammals. So I think moving is seriously important. I think one of the drawbacks for a lot of primatologies that people tend to stay with particular species, and if not, they spend all their lives with working on particular order. I'm very much asking my advice. I'd say don't stick with the same species. You know, work with multiple different sorts of animals and don't stick with the same species throughout your life if you're going to avoid it. Is it intimidating those starting over? Yes, we're scary. You don't know whether a project shouldn't work when you start it. But I think I suppose after you've done it once, it gives you confidence that you can do it. And I think that it's actually, it's quite a valuable thing for people. People in my generation commonly start their own projects. So I mean, I started to work on Red Colourbus at the beginning of my PhD and I sort of disappeared to, I really disappeared to Africa for two years, or first two years of my graduate life and worked on Red Colourbus and then came back and roged it up. And that sort of pattern of work was much commoner in the 60s. So people were actually getting the experience of going off to all of the shop and starting their own projects. And I think that actually is quite useful in giving people the confidence that they can do it. Yeah, I think that I agree with everything you just said. I think the biggest barrier to that is just the huge risk that one faces of it not working. And for a grad student, you start out and it doesn't, you spend two years and it doesn't work. I don't know how do you deal with that. And maybe that's unavoidable. I mean, I'm a child of the 60s and I do think that we were more risk, we were more ready to take risks. And that wasn't just us in this area of biology. I think everyone went, the first generation of breaking free from the 50s and the sort of x-war years of when suddenly there was more money around, there were more opportunities around, airfares were much cheaper. But there was the feeling that in the 60s that everything was open and you should look for your opportunities and take them hard. I left school at 17 and went to work in the game department in XAML.
in New England Valley and was working as a range assistant and doing all sorts of things from anti-poting patrols through to building stuff and through to game censuses and that's all. And that gave me a background of working in Africa and knowing what one could do and for how to operate in a in a wild environment where there wasn't much support. The practicality of that and the business saying, well, yes, there are problems, yes, there are some risks. This is how you control them. This is how you do something goes wrong. There's a lot more existing projects one can join now, of course. You look out at the landscape as a prospective graduate student and there's so many people who have been doing what they've been doing for decades and they're going to be able to teach you everything. Yes, at least that's how you're magic, right? And so it's in some ways it feels like the importance of success in graduate school has increased in terms of getting a postdoc in the later job. Like it feels like there's less room for error and at the same time it feels like there's more people who can give that, give you that feeling that you're going to find success because they know what they're doing. And you're doing, you're you're you're you're getting enormous amount by joining long-term studies and doing that but joining doesn't long-term studies like this doesn't necessarily give you the same sort of practical confidence that you can go out into the blue and start something yourself. Totally agree. Yeah. I take my hat off in this kind in the current situation and to people who have started new studies from scratch in whatever is more where I can do it. I do all I can to to help them because I think that's a you know when you see someone doing that with a new study you think yes that that's what more we want more people to do that. Is it harder today? It's different today. It's always been hard. I mean, even in the 60s it wasn't easy raising money. If you look right back at so early studies of people like Timberg and then Black and the states people like Oriens there's a reason Timberg was moving plankons around. Absolutely. Absolutely because he he was getting tiny amounts of money and there's no way that they could afford full-time assistance or postdocs or anything like that. There's no question of that. I think people don't realize in the sense today what a different you know we can raise an RC grant, an NIH grant that funds one or two postdocs plus a couple of students plus perhaps a technician or two you know five salaries or six salaries for three to five years that that would have the the founding fathers of animal behavior would have I'm sure given their IT for that sort of funding. Okay so you want to give one away for free if you're going to start over pick a new a new animal to do a long-term study of what would it be? Well I think the number of one dose in India won't they've not been there a cop to breed a dog. I don't know that. The Kippling's red dog. Okay. The ASEATIC equivalent of wild dogs haven't been studied. Okay. I very much like to see Kuzimance is studied which are the so forest dwelling west African banded Mongoose equivalent. Traditionally there have been few long-term studies on solitary species and few long-term studies on monogamous species but I think that's important. For example I think one of the major discoveries comes from Peter Kaplan's work on on lemurs where he shows that nocturnal lemurs which are solitary quite a number of them look as if they're actually monogamous they you've got a male and a female sharing a common range and just coming together to breed and otherwise foraging and living separately avoiding each other for most of the year for example but breeding repeatedly throughout their lifespan. Well I've talked about the placement of the field of animal behavior relative to science yeah that's to say natural scientists versus social scientists so let's start with your kind of history your your your history and history. Well my history was that I've that to the school level I was never very good at maths and I was quite good with words and so going down the humanities side was the natural thing for me and we specialised much earlier under new doing state in secondary school we either do languages or humanities or we do sciences and you can't mix both we can't easily mix both I did my A levels in history in English, Latin, French and then I did university exams in history to go to chemistry history and the thing about that is that I was this was in the early 60s when economics interpretations of history were dominant and I was really fascinated by the way the difference is in the structure of societies in the control of societies in art in literature in philosophy and so on were related to changes in the structure of societies so I was always interested in social processes and what the social and physical environment and economic environment was generated was always an at-for-list I went to a Luangua Valley and worked as a ranges assistant in a in a marvellous in an African environment doing really exciting work I came back Cambridge and I faced reading constitutional British history between 1485 and 1830 for the sort of fourth time round I thought no I really don't want to do this and I said would they let me read Zoology and they said absolutely not you've got no science A levels we wouldn't dream of wasting a place in biology on you so I looked around see what I could do and I could do archaeology and anthropology and David Pilbym was there and I could specialize in biological anthropology and I spent my my summers doing expeditions so mostly in archaeology and then I picked up Crook and Gartland 1966 Crook and basic they said well primates belong there's a difference in group size between montane species and forest-reling species and savanna-dwelling species and that's how you should think about it and I I done and there was enough literature leaking through to thinking it was more complex so as my undergraduate project I did a statistical analysis of this and showed that Crook's category weren't very satisfactory and I then went off and worked on on colorbus on red colorbus and black and white colorbus with which were forest-relingly should have been a small group recording we had red colorbus working on in groups of up to 80 and black and white maybe in little family units and that was my PhD I managed to shift to often I done my undergraduate project and done this analysis Crook I persuaded Robert Hine to take me on Robert Hine's supervised Jane Goodall and Diane Fossie Jane Goodall with had no degree at all which in our Cambridge system we were able to do a PhD directly on that and Robert I guess being the sixties he was prepared to he was quite happy to take on people with strange backgrounds actually if you look around they're not quite a high proportion of the early people at that stage and the that half of the sixties early seventies came people like Bob Trivers have I think his background was in economics also ecology it wasn't a traditional biology background David Lack was sort of the main provenant was actually a teacher but a good naturalist and he came into it that way so there were a lot of people that have rather today rather strange backgrounds coming into the field so David Lack set a huge impact on population dynamics and birds and on and on life history and birds and all that tremendous tremendous influence in that area it's a very unfortunate UK that we have this hard division between human sciences and natural sciences and I think the the fact that cultural anthropologists and biological anthropologists have difficulty talking with each other is very unfortunate and very unnecessary I think it's an enormous shame that one can't actually mix around because I think really the sort of area we work on yes easily a but geography it easily but economics it easily buts or social theory and so on and there's a lot that we can learn from those areas and that they can learn from us absolutely yeah so you know from that description it seems obvious but most people would place what we do in natural sciences but it seems clear that the things we're interested in are closer to social sciences than physics yes so hope would be in my head that in that over the next 50 years your generation.
will actually bring together the social sciences and natural sciences. I don't see much sign of it. Yeah, I don't know. I've always found history, psychology, economics, these things all are deeply interesting to me. I desperately feel like I have a lot to learn from them. Social scientists need to realize the sorts of things they're interested in have biological consequences, which in turn have behavioral consequences. I want to kind of close by thinking about how you've seen the field of and all behavior change over time. And you've talked a bit about the great, the funding, kind of things we've talked about a few long term studies. How else do you feel like across your career you've seen the field change? Technology, I did my thesis when I started collecting data. I started collecting on file cards, you used to have a needle through and lifted up and went, "You spent your time punching holes at the edge of cards? Do you know about this?" You didn't know. So you have all the file cards. You don't even know about it. Then if you ever read Cinneseagle, non-parametric statistics with behavioral sounds, wonderful, a Bible on, because you did your statistical test by hand from where it go. You couldn't punch a button and have a man witness you test and did your man, you ranked your numbers in your different categories and you calculated the text. People in my generation, Cinneseagle was their Bible and it was wonderful and we all did that. And I mean even people like Nick Davis would say, "Well, if you can't show it with a man witness you test, it's not this problem up there." So it was kind of a long way from that. The result is, you know, the amazing statistics and the other side is that you just press your button and we could so slightly understand what was happening in there with the U-test. You can't really understand what's happening with that. Well, you're unlikely to understand it. But let's you throw away so much less data. That's right. Like getting to use all your data in well-constructed mixed effects models. Absolutely. What's something that you've thought pretty strongly, how strong feeling about that you ended up changing your mind about? How do we define text or selection? Yes. And how do we define it in a way that it's not a almost inevitably a male characteristic? Here are the huge benefits to female dominance. Is there all with your baboons? Let's take your baboons. And you've got females competing for the dominant position. You've got them competing to breed and this sort of thing. In some cases, in some animals, this can lead through to specific characteristics associated with competition in females. In some cases, weaponry. That's not all that common. That looks to me like a process which is effectively identical to the process of sexual selection in males. With the exception that it is not operating through variation in mating success, it's operating through variation in reproductive success. I don't think it's clear that when Darwin defined these terms, I think if you read sexual selection in the descent of man, Darwin was using sexual selection in terms of a process of intracexual competition to breed. That's right. Not just to mate. Both sexes can be undergoing sexual selection. Exactly. I've wanted to argue that quite a lot of the characteristics of female selection for high testosterone levels to enhance competitive ability should be regarded as a sexual selection trait. And defining sexual selection purely in terms of mating success is that you end up with defining sexual selection in terms of effectively restricted to males. I think it's quite useful to think about sexual selections abroad as characteristic that applies to both males and females. As you look at the current kind of state of anal behavior, what you're excited about for the next, the next 50 years. I mean, it's the thing that the whole string immediately thinks I'm not very excited about. I'm not very excited about the number of papers that do heavy statistical analyses in particular network analysis to generate trivial results. I think referees ought to be much more focusing on whether the paper actually produces a novel result than whether the process is to get there a correct or not. And that's because you feel like social networks are not readily interpretable. Yes. And I feel that in lots of cases, you can use much more direct measures where you can intuitively understand what the confining variables are likely to be. And as you move to network analysis, you've got basically no idea. You don't really know how the taught extent of measures going to screwed up by confining parameters. Let me play devil's advocate for a moment. Selection doesn't care about our ability as researchers to understand exactly what's happening. And so what I see the value of networks as being is the ability to capture complexity that my brain is not capable of rocking very well. And so I do agree when I have to be super careful because when one doesn't know what one's actually measuring, right. And I see this also. And I'm in a neuro biology department when I see people do cluster analyses of behaviors that they don't even know the behaviors are, but you know, they will move in this way and that's a behavior. That that feels odd to me. Yes. Right. And I get a little bit of that feeling from network analysis to, but I also see it as a potential opportunity to overcome our inability to understand complexity. And I can totally agree with that. I'm not against net theory. I'm against. I think a lot of cases when that drug analysis is used in circumstances where you could use a much more direct and much simpler measure. Yes, I do agree with that fact you can find correlations between a boom eigenvectors on on sociality. It doesn't mean say that eigenvectors are necessarily any great benefit rather than comparing centrality and eigenvectors and the different parameters for network analysis rather than comparing their relationships to fitness measures. I'd like to see them being compared to direct measures of time spent within five meters or each other. So direct simple measure trying to understand and maybe it lets you let you start to understand what that complexity is actually capturing. Something I am excited about. Oh, I'm excited. I mean, lots of things are excited about my field, the amazing work on social insects, Laurent Keller's work. Very exciting. I'm pretty excited about the idea that there's a lot more complexity in social species, but I know we should have worked on that myself. I don't know. I'm very excited about maternal effects. Yes. And I think there's a huge area of maternal effects to explore. And I'd love to see that start to encroach more on medicine and feed into the social sciences and all that sort of thing. Specific things I'm excited about. The work that's coming out on Australian cooperative breeders, where you have these whole strange systems where you have multiple breeding females, multiple breeding pairs and then help us move between pairs. So it's a very different system to us once I'm used to. Oh, I'm pretty excited. Although I'm a bit skeptical about the whole idea of being able to look at pilot genetic sequences. I think that's a major. And the genetics makes that. And the genetics makes that is. I think that the problem is that there's probably much more backtracking than we think. So whereas I did think I changed there's an area of general mind. I did think that, you know, if you think about what made in social organization, the start the sort of species and then it evolved into. Gradually increasing forms of sociology and I know tend to think it's probably vast. One of the interesting examples of that. So if you look at the deer species, you very easily think that the most primitive deer species were solitary or monogamous. You know, then you got small groups and then you got progressively bigger groups. But recently they've been able to identify the origins of deer moving into the new world and we know roughly when that happened. And we know roughly where it came from, so the things that moved into the new world were probably something like white tail deer. They were obviously, you know, they were dimorphic. They were medium sized. They had reasonably large antas. They were almost certainly polygons and so on. So they moved into the new world and radiated through the new world. And you know, got these things like who do and Brock it dear. Which are out to this little things living in solitary ranges in South America. And they are derived from a much more social and much more dimorphic. An ancestor.
And one of the interesting things that is in the old world, solitary species, with the don't have antlers, they have canines instead. So things like Montreal and Chinese water, have waking, the males have waking, great canine, they fight by biting. In the new world, solitary species, they don't have canines at all. Well, if they have canines, they're normal, they don't have done morphic, sexes, they don't morphic canines. Very interesting. So totally different. I'm excited by people starting to work on social dynamics. My group start and are looking at the life history of groups, as well as the life history of individuals. And we finally have, we're getting to the point where we have the power to do that. Yes. And I'd like to see people using the same sort of approach on relationships. I want to know more, not just about how the factors that affect whether individuals acquire dominance, but the factors affect the persistence of dominance. And I'd like to understand the loss of dominance. We're missing a pup. We're missing two pup. Oh, there they are. What excites you? What excites me is senescence and differences between chronological senescence and biological senescence. Increasingly, I think a lot about terminal effects, how much life does an animal have left and how does that shape its behavior rather than how old is it. And I think that part of the power of really long term studies is you know an animal's die. And so you can look backwards instead of saying how old is it alive, you can say how long is it too long? What have you previously? And that's something that I'm excited about. At this kind of stage in my career, I'm thinking about things that I'm, that I think I'm good at and things that I know I'm not good at. So what's something that you see as a weakness of yours and how do you manage, how do you overcome that scientifically? Well, I'm weak on technology and I'm aware of that and I overcome that by having people working with me who are not and I'm quite happy to leave it to them. I'm quite good at delegating and possibly I delegate too much. I'm not, I don't know, where do you, there are masses of people around for a much more perfectionistic than I think is good for them. I think I'm probably, possibly at the other extreme that as far as day to go, I'm principally interested in whether I think they're adequate, rather than whether I think they're perfect. And Dave Gainer is fantastic because he did, he really absolutely wants to understand every, every step of the way. I'm prepared to take things from granted that I don't understand because I know I'm unlikely ever to understand them. So I never really, I'll never be able to understand the animal model fully. I can see what it does, but I don't follow the mathematics of it. As someone like to his feet and the whole, once I just have accepted it. And that's definitely, that's obviously weakness. But by working with people who are able to under, we have confidence are using it expertly, that overcomes it. Well, it may do, but it means, if it was judge people's reliability, yes, they can have problems. Right. I guess that's kind of the way I'm thinking about it is, you know, when I obviously can't do everything within a lab group or within a project. And how do you know when you can trust other people who be doing things to the level of excellence you would expect at yourself? Difficult. And you make mistakes on that. But you make mistakes all, I mean, I'm very struck that it's serious, it's difficult to pick who are good, going to be good PhD students. And if I look back over my career, I've really had no concept of who we're going to be stars and who we're not going to be stars. And I often got that wrong. I think one of the benefits I was saying to, is that on the whole, we live our lives working with people who will inform recent or pleasant interesting. And that's one actually one of the major benefits of the academic life. Well, Tim Clinton-Brock, thanks so much for taking this time and being so giving with both your hospitality and your thoughts and your time. You're very welcome. Very welcome. It's a pleasure talking about. It's a pleasure talking about a lot of the process that's involved in doing an academic life. I've seen something as a general alarm. So they've seen, can we see it? Pale chanting girls walk. Yeah, pale chanting girls walk would take pups. So that for a group like this with pups with small pups with the group, that is a real risk. So they're bunching and defensively and they may well head back to the barrow. I just think that's where they're going at the moment. But this is, look, look here, they've got the tales happens for war, dark salams, go. So they're all bunching defensively. And this is very much a costume with the tales up like this. The Animal Behavior Podcast is created by a talented team of animal behavior researchers. We have three excellent content editors, Nico Hensley, an NSF postdoctoral fellow, studying the evolution of neurocensory systems and their impact on animal communication and speciation at Cornell University, Camilla Cheny, who studies tool use, object play, and animal innovation in non-human primates, and Logan James, a postdoctoral fellow at the Smithsonian Tropical Research Institute, studying acoustic communication in frogs and birds. Our communications director is Casey Patmore, a PhD student at the University of Edinburgh, studying the behavior of varying beetles. You can follow us on Twitter @AnimalBehavePod or check out our website at animalbehaveyourpod.com. Our education team makes lesson plans and classroom materials that you can incorporate into your undergraduate classes. You can find those materials on our website. The education team is Emily McClain, an assistant professor of biology at Oxford College at Emory University. Georgia Lambert, a PhD candidate, studying parental cooperation in varying beetles at the University of Edinburgh, and Smile Chaudry, a recent master of research graduate in biological sciences from the University of Exeter, who worked on camouflage and escape responses in green shore crabs. Our sound director is Brian Level, a PhD candidate studying the evolution of acoustic signals in Hemenon Bernal's lab at Purdue University. This season, I'll be recording my side of most conversations in the Cornell Broadcast Studios with engineering support from Bert Odom Reed. Our art is all produced by animal behavior researchers. Our logo was designed by Adelaide Johan Montaille. Our theme music is by Sally Street and transitions are by Andreg Gensolsch. I direct and host the show along with my co-host Amy Strauss. 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