WEBVTT 00:01.840 --> 00:03.360 Prof: Okay, let's get going. 00:03.360 --> 00:07.870 We're into the last segment of the course. 00:07.870 --> 00:09.800 We did evolution, and then we did ecology, 00:09.799 --> 00:11.539 and now we're going to do behavior. 00:11.540 --> 00:15.320 I think the sequence does make sense, 00:15.320 --> 00:18.960 because evolution helps to explain how the things we deal 00:18.963 --> 00:22.843 with in ecology evolved, and it also explains how much 00:22.839 --> 00:25.469 of what we see in behavior evolved. 00:25.470 --> 00:30.470 But I want to say at the outset that the behavioral ecology view 00:30.468 --> 00:35.068 of behavior--which is basically expressed on this slide; 00:35.070 --> 00:38.700 so behaviors evolved--the evolved patterns that we see in 00:38.700 --> 00:42.530 behavior should reflect things that happen frequently to the 00:42.525 --> 00:46.835 organisms in their environment, and the way animals behave 00:46.836 --> 00:51.016 should reflect the consequences of behavior for lifetime 00:51.016 --> 00:52.836 reproductive success. 00:52.840 --> 00:56.910 That is really only part of the biology of behavior. 00:56.910 --> 01:00.510 If you really want to understand it at all levels, 01:00.506 --> 01:03.586 you have to understand how behavior evolved 01:03.591 --> 01:06.431 phylogentically; so you need a comparative view 01:06.433 --> 01:07.033 of behavior. 01:07.030 --> 01:10.630 You need to understand this issue, which is how is it that 01:10.628 --> 01:13.798 behavior is adaptive; is it, or is it a maladaptation? 01:13.799 --> 01:17.749 But then you also need to understand how behavior 01:17.753 --> 01:21.063 develops; that is, if we follow the 01:21.061 --> 01:26.251 organism from zygote to death--you'll see some patterns 01:26.251 --> 01:30.961 of that today--how is it that organisms learn? 01:30.959 --> 01:32.179 How is behavior acquired? 01:32.180 --> 01:34.360 That's a whole field in and of itself. 01:34.360 --> 01:37.310 And then finally we need to understand the mechanistic 01:37.310 --> 01:38.870 underpinnings of behavior. 01:38.870 --> 01:44.620 So in that respect there are a lot of different ways you can go 01:44.617 --> 01:45.357 at it. 01:45.360 --> 01:47.270 You can go at it through neurophysiology; 01:47.269 --> 01:49.209 you can go at it through endocrinology. 01:49.209 --> 01:53.029 There are many different kinds of mechanisms that are involved 01:53.030 --> 01:55.160 in triggering behavior patterns. 01:55.160 --> 01:58.970 So what we're going to concentrate on in this course is 01:58.965 --> 02:02.555 primarily the behavioral ecology approach to it, 02:02.560 --> 02:05.670 which is well exemplified in the book that you've got by 02:05.665 --> 02:06.735 Krebs and Davies. 02:06.739 --> 02:10.379 But these other issues are also very interesting biology, 02:10.378 --> 02:14.718 and I'm just indicating that if you get interested in behavior, 02:14.718 --> 02:16.738 there are lots of ways you can go at it, 02:16.740 --> 02:21.020 and there are entirely different paradigms you can use 02:21.018 --> 02:22.308 to analyze it. 02:22.310 --> 02:25.800 So the five themes that we'll approach--and these are the next 02:25.802 --> 02:28.382 five lectures; so this is a sketch of how the 02:28.378 --> 02:29.318 course finishes. 02:29.318 --> 02:32.328 Today we'll talk about foraging and hunting. 02:32.330 --> 02:35.820 Then next time we'll talk about evolutionary game theory, 02:35.818 --> 02:39.118 which is one of the major analytical frameworks within 02:39.119 --> 02:41.299 which people approach behavior. 02:41.300 --> 02:44.270 We'll have a look at mating systems and parental care, 02:44.274 --> 02:46.804 and they are connected in interesting ways. 02:46.800 --> 02:50.090 We'll take a look at alternative breeding strategies, 02:50.090 --> 02:53.290 which are frequency dependent breeding strategies, 02:53.288 --> 02:57.248 often best analyzed with evolutionary game theory. 02:57.250 --> 03:03.650 And then we'll close with the evolutionary and ecological 03:03.647 --> 03:09.247 analysis of selfishness, altruism and cooperation, 03:09.246 --> 03:12.556 in animals and in humans. 03:12.560 --> 03:15.780 So those are the five themes that I have selected out of 03:15.780 --> 03:18.650 behavioral ecology to emphasize in this course. 03:18.650 --> 03:21.520 It's an introductory course and, you know, 03:21.520 --> 03:24.950 frankly it would be great to give you an entire semester just 03:24.952 --> 03:27.062 on behavior, because it's such an 03:27.056 --> 03:28.236 interesting topic. 03:28.240 --> 03:30.700 But I will signal that we do have other courses in the 03:30.695 --> 03:32.775 department on it, and if you get interested in 03:32.781 --> 03:34.451 them, they might be fun to take. 03:34.449 --> 03:37.649 Okay? 03:37.650 --> 03:41.210 So I'm going to start with foraging by bringing back in 03:41.212 --> 03:44.382 something that you guys presented on that Friday, 03:44.379 --> 03:47.019 which is the marginal value theorem. 03:47.020 --> 03:51.150 And this time I'm going to apply it not to whether you 03:51.150 --> 03:54.090 should fill your plate up, your tray up, 03:54.092 --> 03:56.782 in the dining hall full, if you're going to the far end, 03:56.775 --> 03:59.275 or just with a little bit if you're going to be close to the 03:59.280 --> 04:00.990 counter, but to the issue of how long 04:00.992 --> 04:03.632 you should guard your mate, and indicate that, 04:03.633 --> 04:08.593 in fact, the same intellectual framework applies in both cases. 04:08.590 --> 04:12.410 Then I'll give you an example where we can actually do a 04:12.406 --> 04:16.286 clever experiment to get the foraging organism to tell us 04:16.291 --> 04:18.861 what fitness measure it is using; 04:18.860 --> 04:24.290 and that's often a very satisfying kind of experiment to 04:24.290 --> 04:29.030 do, if you can get the animal, which cannot talk, 04:29.031 --> 04:33.181 to tell you what it thinks it's doing. 04:33.180 --> 04:39.750 Then I'll illustrate how two different birds deal with risk. 04:39.750 --> 04:43.270 Because a small bird, at the end of a cold winter 04:43.266 --> 04:47.656 day, is exposed to extreme risk of dying overnight--and I can 04:47.663 --> 04:50.013 tell you this is quite real. 04:50.009 --> 04:52.269 Over the course of a normal Connecticut winter, 04:52.269 --> 04:55.059 I am often picking up the occasional house sparrow, 04:55.060 --> 04:57.810 or robin, or whatever, which has died next to my house 04:57.807 --> 05:01.987 because we've had a cold night; so that risk is real. 05:01.990 --> 05:06.010 Then I'll discuss a little bit how predators shape crypsis and 05:06.012 --> 05:07.202 conspicuousness. 05:07.199 --> 05:10.649 But then the sort of--the thing that you'll probably remember a 05:10.651 --> 05:14.211 week from now is the part of the lecture that deals with why hunt 05:14.213 --> 05:15.053 in a group? 05:15.050 --> 05:17.710 And at that point I'll show some chimpanzees hunting. 05:17.709 --> 05:20.989 And I want to warn you that is not something where you want to 05:20.985 --> 05:23.985 be bringing- eating the food that you've brought into the 05:23.992 --> 05:26.562 room, or anticipating lunch, 05:26.555 --> 05:30.325 okay, because this is pretty gory stuff. 05:30.329 --> 05:34.439 Okay, marginal value theorem. 05:34.440 --> 05:38.520 The important thing about the marginal value theorem first is 05:38.516 --> 05:41.436 that it's dealing with foraging in space. 05:41.440 --> 05:44.020 And it's assuming you're starting in one point, 05:44.019 --> 05:46.379 which would normally be your home, your nest, 05:46.379 --> 05:49.019 your refuge, your den, and you are going out 05:49.024 --> 05:52.104 to another point where you are looking for food. 05:52.100 --> 05:53.540 And you have options. 05:53.540 --> 05:55.970 You could either go to this place or you might go to some 05:55.971 --> 05:57.841 other place, when you go out to get food. 05:57.839 --> 06:01.279 So you have to travel to get to that patch of food, 06:01.278 --> 06:02.678 and then you have to search in the patch, 06:02.680 --> 06:05.950 and then once you start getting food in the patch, 06:05.949 --> 06:09.589 you accumulate it--and this is a cumulative curve-- 06:09.588 --> 06:13.638 in a way that expresses diminishing marginal returns. 06:13.639 --> 06:15.639 So the harder- the longer you're in the patch, 06:15.642 --> 06:17.962 the harder you have to look, basically because you've 06:17.958 --> 06:20.138 already eaten some of the stuff in the patch. 06:20.139 --> 06:21.229 Okay? 06:21.230 --> 06:25.180 So there are a number of things to remember about this kind of 06:25.182 --> 06:25.832 diagram. 06:25.829 --> 06:28.159 One is the X axis is time. 06:28.160 --> 06:33.180 Two is it's split up into travel time and search time, 06:33.180 --> 06:36.770 and at the point that you start searching is where you draw your 06:36.771 --> 06:39.351 payoff curve here, because that's the point at 06:39.348 --> 06:42.258 which you're going to draw this cumulative payoff curve. 06:42.259 --> 06:46.289 The vertical axis is some kind of payoff, and it's assumed to 06:46.291 --> 06:48.511 have a relationship to fitness. 06:48.509 --> 06:49.179 Okay? 06:49.180 --> 06:52.910 So it's going to be food, or it could be mates. 06:52.910 --> 06:59.090 And probably the clever thing, the most clever thing about 06:59.088 --> 07:05.378 this--and I well remember when Rick Charnov first drew this 07:05.375 --> 07:07.485 thing; he and I were grad students 07:07.494 --> 07:09.684 together, and he was in his office and he was drawing this 07:09.677 --> 07:10.787 thing on a piece of paper. 07:10.790 --> 07:12.630 So I saw it before it was published. 07:12.629 --> 07:15.599 The clever thing is the nice geometrical solution to the 07:15.603 --> 07:16.743 optimality problem. 07:16.740 --> 07:22.370 And the way to think of that is this: here is the measurement of 07:22.374 --> 07:24.674 time, and the question is at what 07:24.673 --> 07:28.023 point should you stop searching in this patch and go on to 07:28.019 --> 07:28.899 another one? 07:28.899 --> 07:35.739 And if you imagine all the possible lines that you could 07:35.735 --> 07:38.865 draw, that fan out from this axis, 07:38.865 --> 07:43.665 it turns out that the one which is tangent to that curve has the 07:43.672 --> 07:44.972 highest slope. 07:44.970 --> 07:45.810 Okay? 07:45.810 --> 07:47.660 So the slope--I mean, you can see that, 07:47.661 --> 07:49.611 just geometrically from looking at it. 07:49.610 --> 07:52.680 Okay, this is the line which is going to have the highest slope 07:52.678 --> 07:55.448 of all of the possible lines that you could draw that are 07:55.452 --> 07:56.692 within this envelope. 07:56.690 --> 07:59.380 And you can't go above this line. 07:59.379 --> 08:02.629 The reason you can't go above this line basically is that 08:02.629 --> 08:05.299 you're not getting any food above that line. 08:05.300 --> 08:09.370 This line is defining the rate at which you can conceivably 08:09.369 --> 08:12.039 accumulate food, just by the ecological 08:12.035 --> 08:14.135 constraints of that patch. 08:14.139 --> 08:18.899 And so this is the maximal point for the slope. 08:18.899 --> 08:21.879 And then you ask yourself, what is the slope? 08:21.879 --> 08:27.169 Well the slope is ∆y/∆x. 08:27.170 --> 08:28.250 Right? 08:28.250 --> 08:32.050 ∆y is the change that you get, or the amount of payoff you 08:32.049 --> 08:34.949 get per amount of time you spend searching; 08:34.950 --> 08:37.200 so it's the payoff per unit time. 08:37.200 --> 08:39.830 So drawing a line that way, as a tangent, 08:39.827 --> 08:42.257 maximizes the payoff per unit time. 08:42.259 --> 08:43.829 You don't have to write down any equation; 08:43.830 --> 08:45.720 it's just geometry. 08:45.720 --> 08:47.120 Okay? 08:47.120 --> 08:51.650 And that actually is the cool thing about the marginal value 08:51.650 --> 08:52.420 theorem. 08:52.418 --> 08:58.868 Now I want you to imagine you are walking through a field in 08:58.868 --> 09:01.388 the Alps, and there are some cows grazing 09:01.393 --> 09:03.783 in the field, and you look off to the side 09:03.784 --> 09:06.884 and you see two biologists down on the ground, 09:06.879 --> 09:09.979 looking at a cow-pie, and you wonder what the hell is 09:09.982 --> 09:12.142 going on; why do they have their noses 09:12.139 --> 09:13.639 down in a pile of cow dung? 09:13.639 --> 09:17.589 And the answer is they are looking at this guy. 09:17.590 --> 09:19.130 Okay? 09:19.129 --> 09:22.869 And one of the biologists is Geoff Parker and the other one 09:22.873 --> 09:24.693 is me; because I had this happen to me 09:24.692 --> 09:25.702 with Geoff in the Alps. 09:25.700 --> 09:26.900 Okay? 09:26.899 --> 09:31.719 So a cow--let me go back--a cow has come along and it has 09:31.721 --> 09:35.981 dropped a pile of dung here, and it has been rapidly 09:35.984 --> 09:38.554 colonized by a bunch of dung flies, 09:38.548 --> 09:41.748 and because of that, within about a minute, 09:41.750 --> 09:45.670 a male will find a female and they'll go into copulation. 09:45.668 --> 09:49.118 And then the issue is how long should the male stay on that 09:49.115 --> 09:52.555 female before he jumps off and goes to find another one? 09:52.558 --> 09:57.528 You can measure how many of the offspring of that female will 09:57.529 --> 10:00.179 get fertilized if he stays on. 10:00.178 --> 10:03.998 And this is the proportion of eggs that he will fertilize, 10:04.000 --> 10:05.140 if he stays on. 10:05.139 --> 10:06.069 Okay? 10:06.070 --> 10:10.480 So it's starting to look pretty much like a marginal value 10:10.476 --> 10:11.866 theorem problem. 10:11.870 --> 10:15.800 10:15.798 --> 10:20.058 You can set up the analysis as search and guard time, 10:20.057 --> 10:22.347 plus time spent in copula. 10:22.350 --> 10:27.070 So this is his- how long it takes him to search and guard a 10:27.067 --> 10:31.537 female, and this is how long he spends in copula on her, 10:31.543 --> 10:33.743 and this is his payoff. 10:33.740 --> 10:36.900 And now look at what the Y axis is. 10:36.899 --> 10:41.669 It's directly fitness; that's a direct fitness payoff. 10:41.668 --> 10:45.118 So actually this is the purest form of the biological 10:45.123 --> 10:47.983 implementation of marginal value theorem; 10:47.980 --> 10:50.970 finding a mate and fertilizing some eggs. 10:50.970 --> 10:55.250 Now the interesting thing is that he jumps off about ten 10:55.245 --> 10:57.805 minutes earlier than predicted. 10:57.808 --> 11:03.568 Any ideas on why he might jump off ten minutes earlier than 11:03.567 --> 11:04.757 predicted? 11:04.759 --> 11:10.679 11:10.678 --> 11:17.008 It's actually a method of risk minimizing or spreading of risk. 11:17.009 --> 11:22.059 He cannot predict when the next fresh cow-pie is going to hit 11:22.061 --> 11:25.361 the ground, and that's going to be the next 11:25.357 --> 11:28.637 open resource which gets colonized by females, 11:28.639 --> 11:31.819 and he needs some time to find it, because he wants to be the 11:31.823 --> 11:32.783 first one there. 11:32.779 --> 11:35.309 If he can be the first one there, he'll get the best 11:35.312 --> 11:35.762 female. 11:35.759 --> 11:39.399 So he has to jump off this female a little bit earlier than 11:39.403 --> 11:41.793 this simple analysis would predict, 11:41.788 --> 11:47.238 just to hedge against the problem of trying to be first to 11:47.241 --> 11:49.061 the next cow-pie. 11:49.058 --> 11:52.508 This problem is actually almost quantifiable. 11:52.509 --> 11:56.179 Of course you are measuring some rather strange things. 11:56.178 --> 11:58.958 You're sitting there with your stopwatch watching as the 11:58.960 --> 12:01.230 cow-pies arrive on the surface of the pasture, 12:01.234 --> 12:01.744 right? 12:01.740 --> 12:05.010 But it's a completely analyzable problem. 12:05.009 --> 12:08.759 Now, so that's the marginal value theorem applied to mate 12:08.761 --> 12:09.501 guarding. 12:09.500 --> 12:14.300 Now let's look at an experiment that you can do with honeybees. 12:14.298 --> 12:18.758 And this was designed by Paul Schmid-Hempel, 12:18.756 --> 12:22.796 who is a very clever Swiss biologist. 12:22.798 --> 12:27.628 And what Paul did was he built a model in which he could 12:27.634 --> 12:32.914 predict how long bees would spend flying between flowers, 12:32.908 --> 12:35.988 and how many flowers they would visit before they came back to 12:35.985 --> 12:36.535 the hive. 12:36.539 --> 12:39.179 Okay? 12:39.178 --> 12:43.798 And he had two different measures that they might be 12:43.803 --> 12:44.533 using. 12:44.529 --> 12:47.389 So this is the optimal relationship between how long it 12:47.385 --> 12:50.285 is flying between flowers and how many flowers you visit 12:50.294 --> 12:52.204 before you fly back to the hive. 12:52.200 --> 12:57.340 In one model he had calories per minute, which is the usual 12:57.339 --> 12:57.959 rate. 12:57.960 --> 12:58.650 Okay? 12:58.649 --> 13:01.219 In the marginal value theorem, when you maximize that slope, 13:01.220 --> 13:04.350 if you're measuring that payoff curve in calories what you're 13:04.351 --> 13:06.911 measuring then is maximal calories per minute, 13:06.909 --> 13:09.059 in the marginal value theorem. 13:09.058 --> 13:11.548 Paul thought well maybe they have another fitness measure. 13:11.548 --> 13:14.528 They might be using calories gained per calories spent. 13:14.528 --> 13:16.358 So he built the two models, and he had quite different 13:16.360 --> 13:18.840 predictions, and then he manipulated them by 13:18.841 --> 13:22.641 gluing a little wire onto their back and adding a tiny little 13:22.636 --> 13:23.266 weight. 13:23.269 --> 13:26.119 So he made them--you know, he had a series of treatments 13:26.116 --> 13:29.216 where they weren't weighted and then they had a little weight 13:29.221 --> 13:31.191 and then they had a lot of weight. 13:31.190 --> 13:32.230 And this is what they did. 13:32.230 --> 13:34.070 This is the data, okay? 13:34.070 --> 13:38.190 So it really looks like he was able to get them to tell him 13:38.191 --> 13:42.811 which fitness measure they were using while they were foraging. 13:42.808 --> 13:45.498 That's a very, very clever experiment. 13:45.500 --> 13:48.690 And this is the kind of thing that you can do in behavioral 13:48.690 --> 13:49.240 ecology. 13:49.240 --> 13:52.640 It's possible to construct situations in which the 13:52.635 --> 13:56.305 decisions that the animals are making are so precisely 13:56.309 --> 14:00.469 constrained that you can get them to give you an answer. 14:00.470 --> 14:04.440 I am waving over all the details in the mathematical 14:04.438 --> 14:05.138 models. 14:05.139 --> 14:06.289 Okay? 14:06.289 --> 14:09.349 That's graduate student stuff. 14:09.350 --> 14:13.650 But I think the essential point is that you can do an experiment 14:13.653 --> 14:17.823 that will get you to tell you what an animal is actually using 14:17.821 --> 14:19.531 as a fitness measure. 14:19.528 --> 14:24.758 Okay, now two comments on the problem of how to deal with 14:24.764 --> 14:28.044 risk; and this is the small bird in 14:28.035 --> 14:29.475 winter problem. 14:29.480 --> 14:31.570 And this was a nice experiment. 14:31.570 --> 14:35.450 This is a Great Tit, which is a European form of 14:35.448 --> 14:39.988 chickadee, and this is an experiment which is done in an 14:39.990 --> 14:43.870 aviary, and this is a variable environment. 14:43.870 --> 14:44.750 Okay? 14:44.750 --> 14:49.740 So when the experiment starts, the food supply starts becoming 14:49.744 --> 14:53.794 unpredictable in time; and that's just a manipulation 14:53.791 --> 14:57.191 that the experimenter is imposing on the animal. 14:57.190 --> 15:01.540 And as the food starts getting unpredictable, 15:01.535 --> 15:04.495 the bird starts getting fat. 15:04.500 --> 15:06.960 That tells you a couple of interesting things right there. 15:06.960 --> 15:09.880 It says normally the bird doesn't like to be that fat, 15:09.881 --> 15:12.861 but it's going to get fat because it sees that its food 15:12.859 --> 15:15.119 supply is getting very unpredictable. 15:15.120 --> 15:18.800 And then the way that you build one control into this experiment 15:18.797 --> 15:21.947 is then to switch it at this point into a constant food 15:21.947 --> 15:24.917 supply so that the environment just becomes nice and 15:24.924 --> 15:27.264 predictable and the bird relaxes, 15:27.259 --> 15:29.049 shrugs its shoulders and says, "Oh, 15:29.048 --> 15:34.018 I can get away with getting back to a normal weight," 15:34.015 --> 15:36.015 and drops its weight. 15:36.019 --> 15:38.279 So that's one way of dealing with it. 15:38.279 --> 15:41.939 But there's another way of dealing with it, 15:41.937 --> 15:47.417 and that is that if you look at this other relative of the Great 15:47.422 --> 15:53.072 Tit--okay, this is a Marsh Tit; it also looks a bit like a Coal 15:53.070 --> 15:55.590 Tit; it's a little bit smaller, 15:55.586 --> 15:58.716 has a black head-- and you put it in a high 15:58.717 --> 16:02.247 variance environment or a low variance environment, 16:02.250 --> 16:05.500 this one, it doesn't change its weight at all. 16:05.500 --> 16:09.840 It just packs on as much weight as it can, by evening. 16:09.840 --> 16:12.330 And by the way, at that latitude it's getting 16:12.332 --> 16:14.942 dark at 4:00 in the afternoon in the winter. 16:14.940 --> 16:17.370 So it's going up to just about peak weight. 16:17.370 --> 16:22.980 And you can see how much it's losing by the next morning. 16:22.980 --> 16:25.450 What it does though is it stores seeds, 16:25.452 --> 16:28.582 and if you put it in a high variance environment, 16:28.575 --> 16:32.475 it greatly increases the number of seeds that it stores. 16:32.480 --> 16:34.880 So one of these species--they're very closely 16:34.875 --> 16:37.235 related by the way; it's interesting that there 16:37.243 --> 16:39.783 doesn't seem to be much phylogenetic component to this; 16:39.779 --> 16:43.059 one of them decided to pack it on as body mass that it carries 16:43.057 --> 16:45.687 around with itself, and the other one decided that 16:45.690 --> 16:47.410 it was going to store seeds. 16:47.408 --> 16:49.688 It may have something to do with the risks of predation. 16:49.690 --> 16:52.470 Big, fat birds don't get away from predators quite as easily 16:52.469 --> 16:53.929 as nice slender little birds. 16:53.929 --> 16:57.809 16:57.808 --> 17:02.328 Now what are some of the consequences of predatory 17:02.326 --> 17:04.536 behavior for the prey? 17:04.538 --> 17:09.528 Well one of them is so-called aposematic coloration; 17:09.528 --> 17:14.078 and that is that if you're carrying around something that 17:14.075 --> 17:18.535 is going to poison your predator, you want your predator 17:18.539 --> 17:19.919 to know that. 17:19.920 --> 17:23.660 You don't want--you know, if you put a bunch of ham 17:23.663 --> 17:26.513 sandwiches out in Saybrook Commons, 17:26.509 --> 17:29.419 and you've got cyanide in five of them, 17:29.420 --> 17:32.400 and it's in the interest of those five cyanide-laced ham 17:32.395 --> 17:36.055 sandwiches not to be eaten, then you want a big warning 17:36.063 --> 17:38.723 label on it that says, "Do not eat." 17:38.720 --> 17:39.460 Right? 17:39.460 --> 17:42.670 Well that's what these are, these warning colorations. 17:42.670 --> 17:47.560 If you go out and you pick up a warningly colored millipede, 17:47.555 --> 17:52.105 and you shake it in your hand, it will smell like bitter 17:52.108 --> 17:53.018 almond. 17:53.019 --> 17:55.169 And it's perfectly safe, by the way. 17:55.170 --> 17:57.750 There's not enough cyanide in it to get hurt by sniffing it. 17:57.750 --> 18:00.280 So you're perfectly safe picking up and shaking a 18:00.275 --> 18:02.515 millipede, except you may feel a little 18:02.518 --> 18:05.908 moral compromise at the act of shaking an invertebrate nervous 18:05.913 --> 18:06.473 system. 18:06.470 --> 18:07.460 Okay? 18:07.460 --> 18:09.580 But they will emit cyanide all over your hand, 18:09.580 --> 18:11.420 and it does smell like bitter almond. 18:11.420 --> 18:15.740 And of course the monarch butterfly caterpillar, 18:15.740 --> 18:20.060 which gets its cardiac glycosides from milkweed, 18:20.059 --> 18:24.839 will cause tachycardia in the birds that eat it. 18:24.838 --> 18:29.098 Tachycardia is a condition where your heart jumps say from 18:29.104 --> 18:32.184 a pulse rate of 80, in a human--let's say if you've 18:32.181 --> 18:35.041 been exercising a little bit you might have your pulse at 80 or 18:35.044 --> 18:37.884 100 or something like that; with tachycardia you go up to 18:37.882 --> 18:40.542 250 or 300 and you pass out, because your heart starts 18:40.536 --> 18:42.536 fluttering and it can't pump anymore. 18:42.538 --> 18:46.088 So that's what eating that will do to a Blue Jay; 18:46.089 --> 18:49.199 it will have cardiac arrest. 18:49.200 --> 18:52.770 So it would take a few of them to do that to you. 18:52.769 --> 18:54.629 So I suggest that if you want to try this one, 18:54.626 --> 18:55.696 don't eat more than one. 18:55.700 --> 18:56.940 Okay? 18:56.940 --> 18:59.060 If you really want to get into it, you're probably getting 18:59.060 --> 19:00.960 threatened if you eat maybe five or six of them, 19:00.960 --> 19:02.820 or make a milkshake out of them or something like that. 19:02.819 --> 19:06.539 19:06.538 --> 19:09.358 One can do experiments with this kind of thing as well. 19:09.358 --> 19:14.518 This is a situation in which chicks, just regular domestic 19:14.517 --> 19:18.587 chicks, were given different colored baits. 19:18.589 --> 19:19.399 Okay? 19:19.400 --> 19:23.480 And in both of these situations they were given seeds that had 19:23.481 --> 19:27.101 been stained green or blue, and they had been soaked in 19:27.096 --> 19:29.586 quinine; and chicks do not like seeds 19:29.585 --> 19:31.265 that are soaked in quinine. 19:31.269 --> 19:34.059 So they were both distasteful. 19:34.058 --> 19:37.568 The only difference here is that in one case the green and 19:37.567 --> 19:41.077 blue seeds are on a green background, and here they are on 19:41.076 --> 19:42.426 a blue background. 19:42.430 --> 19:46.140 And what you can see is that the ones that match the 19:46.140 --> 19:48.760 background continue to get eaten, 19:48.759 --> 19:51.449 and the ones that stand out from the background start 19:51.449 --> 19:52.379 getting avoided. 19:52.380 --> 19:53.450 Okay? 19:53.450 --> 19:56.200 So there's a bit of learning; oh, I don't like to taste these 19:56.201 --> 19:56.641 things. 19:56.640 --> 19:59.800 But then they avoid the ones that they see most easily. 19:59.798 --> 20:01.798 And that's what's going on over here. 20:01.798 --> 20:03.948 It's this process that has produced these colors. 20:03.950 --> 20:08.000 20:08.000 --> 20:09.490 Of course it can go in the other direction. 20:09.490 --> 20:13.400 If you, in fact, are not distasteful, 20:13.400 --> 20:17.670 and you want to avoid being eaten, then often natural 20:17.667 --> 20:22.177 selection will change your morphology in such a way that 20:22.182 --> 20:25.962 it's rather difficult to see what you are. 20:25.960 --> 20:27.150 Okay? 20:27.150 --> 20:30.160 The head is here; that's the end of the right 20:30.161 --> 20:32.491 wing, that's the end of the left wing; 20:32.490 --> 20:35.250 there's a nice wing vein running down the middle. 20:35.250 --> 20:39.720 Some of these things are just remarkably precise in resembling 20:39.721 --> 20:41.921 a dead leaf or other things. 20:41.920 --> 20:44.690 I think some of the ones that I like the most are the praying 20:44.693 --> 20:46.593 mantises that look like flower petals, 20:46.588 --> 20:49.388 and sit on flowers, and grab things when they come 20:49.393 --> 20:53.483 in; they're very nasty. 20:53.480 --> 20:54.970 Here's another one that's very nasty; 20:54.970 --> 20:56.230 same kind of thing, okay. 20:56.230 --> 20:57.740 So this is aggressive mimicry. 20:57.740 --> 21:03.160 These are the light signals which are given out by different 21:03.163 --> 21:09.603 species of so-called fireflies-- in fact, these are beetles--and 21:09.596 --> 21:15.556 they have a light organ, and you can see that there's a 21:15.555 --> 21:19.055 species specific signal pattern. 21:19.058 --> 21:21.878 Now normally you would think, okay fine, they're just 21:21.875 --> 21:23.495 dividing up the frequencies. 21:23.500 --> 21:27.600 They don't need to have an FCC to regulate which frequency they 21:27.602 --> 21:28.002 use. 21:28.000 --> 21:29.770 They're flying through the night. 21:29.769 --> 21:31.929 They see a signal over there and they can say, 21:31.930 --> 21:33.840 "Oh, that is another one of my species, 21:33.838 --> 21:38.188 I'll go check it out and potentially mate with it." 21:38.190 --> 21:42.120 So you might think, oh, that's all just normal mate 21:42.115 --> 21:42.975 behavior. 21:42.980 --> 21:47.530 However, these are males that are flickering, 21:47.532 --> 21:52.192 and then they get a response from a female. 21:52.190 --> 21:52.870 Okay? 21:52.868 --> 21:55.868 So this would be male-male, female, male-male, 21:55.874 --> 21:57.414 female kind of thing. 21:57.410 --> 22:01.200 And so there are some that mimic the light signals of 22:01.204 --> 22:03.894 another species; some females that are sitting 22:03.890 --> 22:05.450 there saying--they're faking it. 22:05.450 --> 22:06.340 Okay? 22:06.338 --> 22:08.638 A male blinks, the female can see, 22:08.636 --> 22:12.666 oh, that's not a male of my species, therefore I can safely 22:12.673 --> 22:13.513 eat him. 22:13.509 --> 22:15.929 So she goes blink, with the signal of the other 22:15.925 --> 22:18.285 species, he flies in, and she chews him up. 22:18.289 --> 22:21.519 In so doing she gets two things. 22:21.519 --> 22:27.179 She gets calories out of him, but in some cases she also is 22:27.181 --> 22:32.651 absorbing a defensive chemical that will protect her from 22:32.646 --> 22:35.376 birds, bats and spiders. 22:35.380 --> 22:37.400 So she gets a double dose. 22:37.400 --> 22:41.320 She gets both calories and she gets defense from doing this. 22:41.318 --> 22:45.038 This kind of aggressive mimicry is reasonably widespread, 22:45.036 --> 22:48.816 and it has been evolved convergently a number of times. 22:48.818 --> 22:52.298 The one that always got me as a kid is the saber-tooth blenny. 22:52.298 --> 22:55.658 You know about cleaning wrasses that come into the mouths of big 22:55.660 --> 22:58.700 fish and clean the parasites off of them and so forth, 22:58.700 --> 23:02.360 and the big fish have evolved, because this is a beneficial 23:02.361 --> 23:03.981 thing, to kind of relax. 23:03.980 --> 23:06.540 And so you will see giant groupers and barracudas opening 23:06.539 --> 23:09.239 their mouths and letting these little fish swim through them 23:09.238 --> 23:10.608 and clean off their teeth. 23:10.608 --> 23:14.188 Well, there is another fish called the saber-tooth blenny 23:14.194 --> 23:17.914 that mimics both the color and the approach behavior of the 23:17.909 --> 23:20.399 cleaning wrasse; which is, by the way, 23:20.396 --> 23:21.486 a sigmoidal dance. 23:21.490 --> 23:22.670 It goes through the water like this. 23:22.670 --> 23:25.260 And so you see one of these things coming up and you kind of 23:25.261 --> 23:27.041 relax and say, "Oh, it's just a cleaning 23:27.039 --> 23:28.219 wrasse, it's going to be fine." 23:28.220 --> 23:29.880 And it comes up, and if it's a human, 23:29.880 --> 23:31.510 it takes a chunk out of your thigh, 23:31.509 --> 23:35.129 and if it's a fish that has its mouth open, 23:35.130 --> 23:37.530 it rips out a chunk of gill and goes running off. 23:37.529 --> 23:41.699 So this is the sort of thing that led Darwin to think that 23:41.703 --> 23:46.103 evolution just fills the world up with things that are taking 23:46.096 --> 23:49.316 advantage of every possible opportunity; 23:49.318 --> 23:54.098 and aggressive mimicry is a good example of this. 23:54.098 --> 24:01.758 One of the interesting issues with predation and parasitism 24:01.758 --> 24:07.038 and mimicry has to do with the cuckoo. 24:07.038 --> 24:10.098 And I want to mention it because it really is an 24:10.103 --> 24:12.193 interesting series of puzzles. 24:12.190 --> 24:13.300 Okay? 24:13.299 --> 24:14.759 So here's a cuckoo. 24:14.759 --> 24:17.449 Cuckoos, by the way, feed on caterpillar larvae, 24:17.445 --> 24:20.525 and so they tend to disappear from places where lots of 24:20.532 --> 24:22.022 insecticides are used. 24:22.019 --> 24:24.049 So they're kind of a canary in a coalmine. 24:24.048 --> 24:27.228 If there have been cuckoos on the landscape and you can't hear 24:27.233 --> 24:29.393 them anymore, it means that intensive 24:29.392 --> 24:32.502 agricultural practice has probably wiped out the entire 24:32.497 --> 24:35.877 large fauna of caterpillars; so they like to eat 24:35.880 --> 24:37.060 caterpillars. 24:37.058 --> 24:40.398 And what they do is they go around and they find a nest, 24:40.398 --> 24:43.798 like a robin's nest here, and they lay their own egg into 24:43.799 --> 24:44.589 the nest. 24:44.588 --> 24:48.458 And they have their whole developmental program set up in 24:48.458 --> 24:52.808 such a way that their baby will hatch earlier than the babies of 24:52.808 --> 24:54.258 the host species. 24:54.259 --> 24:57.459 And its behavior is set up in such a way that the first thing 24:57.464 --> 24:59.814 that it does is-- you know, it's just a tiny 24:59.807 --> 25:02.237 little baby bird, and it's just hatched out, 25:02.244 --> 25:05.014 but it has enough muscular coordination and enough 25:05.008 --> 25:08.278 behavioral complexity to take the other eggs and shove them 25:08.277 --> 25:11.707 out of the nest onto the ground, so it's the only one left. 25:11.710 --> 25:13.790 And then it sits there. 25:13.788 --> 25:17.078 And it's got a very effective feeding behavior. 25:17.078 --> 25:19.158 It opens its mouth, it gives all of the 25:19.163 --> 25:21.473 morphological and behavioral cues that say, 25:21.465 --> 25:23.765 "Feed me, feed me, feed me." 25:23.769 --> 25:26.859 And the parents work really hard, the parents of the other 25:26.856 --> 25:29.616 species work really hard to come in and feed it, 25:29.618 --> 25:31.368 and you get a baby cuckoo out of the nest, 25:31.368 --> 25:33.428 instead of a warbler or whatever. 25:33.430 --> 25:35.110 Okay? 25:35.108 --> 25:39.618 Well why don't the hosts throw out the cuckoo egg? 25:39.619 --> 25:40.579 There they are. 25:40.579 --> 25:41.379 They're good parents. 25:41.380 --> 25:43.220 They come back to the nest. 25:43.220 --> 25:46.190 There's an egg in it that they haven't laid. 25:46.190 --> 25:48.800 Sometimes it looks quite a bit like their egg, 25:48.797 --> 25:51.377 sometimes it doesn't; it all depends on which 25:51.380 --> 25:54.530 particular host species it is and how close that particular 25:54.534 --> 25:57.584 race of cuckoo is to matching the host species color. 25:57.578 --> 26:00.858 Well there are really, I think, two reasons, 26:00.862 --> 26:04.452 but they may not be quantitatively sufficient to 26:04.450 --> 26:06.740 explain everything we see. 26:06.740 --> 26:09.660 One is the source-sink distinction. 26:09.660 --> 26:13.080 The hosts can't adapt as fast as the cuckoo because the 26:13.075 --> 26:16.735 parasitized nests are sinks for the host, and unparasitized 26:16.742 --> 26:19.022 nests are sources for the hosts. 26:19.019 --> 26:22.299 Most of the sources are coming out of nests that have not had 26:22.304 --> 26:23.294 cuckoos in them. 26:23.288 --> 26:25.618 If a cuckoo has gotten into the nest, it's wiped out that 26:25.615 --> 26:26.235 reproduction. 26:26.240 --> 26:28.660 Okay? 26:28.660 --> 26:31.700 So the adaptation is to the source, which is to the 26:31.700 --> 26:34.680 condition without cuckoos, and not to the sink. 26:34.680 --> 26:37.740 But there's another issue, and that is if you're just 26:37.743 --> 26:41.223 starting to evolve the behavior of throwing eggs out of your 26:41.220 --> 26:43.160 nest, and you're not very good at it 26:43.160 --> 26:45.180 yet, you can make a serious mistake 26:45.176 --> 26:47.096 by killing one of your own kids. 26:47.098 --> 26:51.268 So there's kind of a threshold there that you have to get over. 26:51.269 --> 26:54.429 You have to actually--this is a behavior where you actually have 26:54.430 --> 26:57.390 to be accurate and pretty good at it, before it pays off. 26:57.390 --> 27:00.980 Before you get good at it, you are indulging in some very 27:00.977 --> 27:02.127 costly behavior. 27:02.130 --> 27:02.880 Okay? 27:02.880 --> 27:04.570 So that's another reason. 27:04.568 --> 27:08.478 Another reason that we don't see the hosts throwing the 27:08.482 --> 27:12.622 cuckoo egg out is that the cuckoos may be moving on to new 27:12.615 --> 27:13.335 hosts. 27:13.338 --> 27:16.488 So it may be that the cuckoos for say a hundred years 27:16.494 --> 27:18.774 parasitized robins, and the robins may 27:18.770 --> 27:21.020 slowly--slowly, because of these reasons-- 27:21.019 --> 27:23.719 start to evolve a response to cuckoos, 27:23.720 --> 27:26.380 at which point the cuckoos just switch over and start 27:26.381 --> 27:27.611 parasitizing warblers. 27:27.608 --> 27:30.028 And they do that for awhile, and they just keep moving 27:30.028 --> 27:32.628 around among the different species in their landscape, 27:32.630 --> 27:35.630 so that they're always able to stay ahead, 27:35.630 --> 27:41.240 because their evolution is a little bit faster than that of 27:41.239 --> 27:42.399 the hosts. 27:42.400 --> 27:45.040 So that process is hard to observe. 27:45.038 --> 27:48.098 And the egg mimicry isn't very precise. 27:48.098 --> 27:51.728 You can see here, this isn't a very good match. 27:51.730 --> 27:53.420 This is one kind of cuckoo egg. 27:53.420 --> 27:55.070 There are some that are a bit closer to robins. 27:55.068 --> 27:57.078 This might be appropriate for another kind of species. 27:57.078 --> 28:01.608 The egg mimicry isn't very precise, and it is still kind of 28:01.605 --> 28:06.125 puzzling why the hosts don't throw out more cuckoo eggs. 28:06.130 --> 28:08.270 It's an interesting problem. 28:08.269 --> 28:13.269 Now I'd like to talk about hunting in a group. 28:13.269 --> 28:19.329 And this is a situation that is interesting, 28:19.328 --> 28:25.508 both because we can quantify the benefits of foraging styles, 28:25.509 --> 28:29.489 and we can see whether or not animals are actually doing what 28:29.486 --> 28:31.736 is quantitatively best for them. 28:31.740 --> 28:35.500 But it also addresses the whole issue of why animals should 28:35.501 --> 28:36.671 exist in groups. 28:36.670 --> 28:40.340 And since we are a group living and hunting primate, 28:40.338 --> 28:44.438 this is a very interesting thing for us to contemplate. 28:44.440 --> 28:49.450 Now when an individual joins a group, it's making a pretty 28:49.448 --> 28:51.468 fundamental decision. 28:51.470 --> 28:55.760 It's basically deciding that the payoff it's going to get, 28:55.759 --> 28:57.639 from the coordination of group hunting, 28:57.640 --> 29:00.790 is going to more than compensate for the fact it's 29:00.791 --> 29:04.381 going to have to share the food; unless it's an extremely 29:04.375 --> 29:07.565 confident dominant type, it's going to have to share the 29:07.574 --> 29:08.044 food. 29:08.039 --> 29:09.349 Okay? 29:09.348 --> 29:13.118 So what you see in wolves, coyotes, 29:13.118 --> 29:16.518 African hunting dogs and hyenas is that all of these things will 29:16.517 --> 29:19.807 hunt alone for little things and they'll hunt together for big 29:19.806 --> 29:20.396 things. 29:20.400 --> 29:24.340 So go up to Ellesmere Island in the Arctic, pop yourself down 29:24.342 --> 29:27.562 onto a wolf study site, and go out and look at the 29:27.560 --> 29:28.810 wolves hunting. 29:28.808 --> 29:32.068 And if they are hunting for voles and mice, 29:32.068 --> 29:34.858 which are about this big, you'll see individual wolves 29:34.859 --> 29:38.499 moving about the landscape, trapping them with their paws 29:38.503 --> 29:40.073 and munching them up. 29:40.068 --> 29:43.688 However, if they decide to tackle something like a muskox, 29:43.690 --> 29:47.110 they will form up into a pack and go into coordinated 29:47.109 --> 29:49.349 behavior, with a division of labor and 29:49.348 --> 29:52.488 assigned roles, to bring down the muskox. 29:52.490 --> 29:54.200 And the muskox, of course, have a 29:54.195 --> 29:56.745 counter-adaptation, which is a group adaptation, 29:56.748 --> 29:58.518 and they form a protective circle, 29:58.519 --> 30:00.369 and they all face outward towards the wolves, 30:00.368 --> 30:02.698 and they defend themselves with their horns. 30:02.700 --> 30:05.820 Similarly for coyotes, African hunting dogs and 30:05.818 --> 30:06.428 hyenas. 30:06.430 --> 30:08.880 With African hunting dogs, they will be going after 30:08.876 --> 30:11.516 usually small rodents or birds or things like that, 30:11.519 --> 30:14.589 on the ground, but they're actually capable of 30:14.585 --> 30:16.215 bringing down a zebra. 30:16.220 --> 30:17.940 This is a dog which is this big. 30:17.940 --> 30:18.400 You know? 30:18.400 --> 30:21.300 A zebra is a horse, which is this big. 30:21.298 --> 30:26.458 And five or six African hunting dogs, each of which weighs say 30:26.459 --> 30:30.689 about 75 pounds maybe, at max--50 to 75 pounds--can 30:30.688 --> 30:33.478 bring down a 500 pound zebra. 30:33.480 --> 30:38.180 So the interesting thing is that they switch facultatively 30:38.182 --> 30:43.132 from solitary hunting to group hunting, as group size- as the 30:43.132 --> 30:45.362 size of prey increases. 30:45.358 --> 30:50.098 Now in chimpanzees--I'm going to show you a bit of the gory 30:50.097 --> 30:53.037 detail of chimpanzees in a minute. 30:53.038 --> 30:55.258 But before I do that I just want to show you some of the 30:55.262 --> 30:56.762 kinds of things that these capture. 30:56.759 --> 31:01.229 So we have wolves and coyotes and hyenas and African hunting 31:01.232 --> 31:04.302 dogs, and these are pictures where 31:04.299 --> 31:09.219 they have done a sophisticated, coordinated group hunt to bring 31:09.217 --> 31:10.907 down a big piece of meat. 31:10.910 --> 31:22.560 31:22.558 --> 31:24.378 Now let's see what chimpanzees do. 31:24.380 --> 31:31.880 31:31.880 --> 32:14.840 <> 32:14.839 --> 32:15.959 Prof: That's Brutus. 32:15.960 --> 32:20.380 32:20.380 --> 32:24.490 Brutus was born in 1952, August. 32:24.490 --> 36:02.730 <> 36:02.730 --> 36:04.730 Prof: That's an important point. 36:04.730 --> 36:13.440 36:13.440 --> 36:17.920 They use seasoning, a little spice from the weeds. 36:17.920 --> 37:11.050 <> 37:11.050 --> 37:15.010 Prof: So that's in Christophe Boesch's study site, 37:15.005 --> 37:18.605 which is in Tai National Park in the Ivory Coast. 37:18.610 --> 37:27.700 Those pictures were taken in about 1990,1991. 37:27.699 --> 37:31.439 I had been there in February of 1989, and I had been out on a 37:31.436 --> 37:33.176 hunt with that same group. 37:33.179 --> 37:36.569 I had had lunch, which was by the way fruit, 37:36.572 --> 37:37.522 with them. 37:37.518 --> 37:40.148 I ate the same stuff they did, but I was eating fruit, 37:40.148 --> 37:41.238 not colobus monkeys. 37:41.239 --> 37:47.959 Those guys are all dead now, the chimps. 37:47.960 --> 37:50.530 They died of Ebola and they died of poaching. 37:50.530 --> 37:54.750 However, a smaller group has replaced them. 37:54.750 --> 37:58.460 Of all of the chimps that were in that group in 1990, 37:58.463 --> 38:01.753 there is only one female who is still alive. 38:01.750 --> 38:04.600 But the group is back to about oh fifteen or twenty; 38:04.599 --> 38:07.319 at that point there were sixty in it. 38:07.320 --> 38:13.130 So let's look at an analysis of why they choose to hunt in a 38:13.130 --> 38:13.920 group. 38:13.920 --> 38:15.600 Okay? 38:15.599 --> 38:18.769 So the increase must more than balance the cost: 38:18.766 --> 38:19.436 sharing. 38:19.440 --> 38:24.020 They do it during the rainy season. 38:24.018 --> 38:26.208 During the dry season they crack nuts. 38:26.210 --> 38:29.800 So these chimps actually have a culture where they teach their 38:29.802 --> 38:33.162 offspring how to crack nuts with a hammer and an anvil. 38:33.159 --> 38:36.539 This is true west of a certain river in the Ivory Coast and 38:36.536 --> 38:38.746 it's not true in the rest of Africa. 38:38.750 --> 38:43.320 And you can see that they hunt a lot more frequently in 38:43.318 --> 38:45.348 September and October. 38:45.349 --> 38:48.929 And I was actually there and saw the hunt in February; 38:48.929 --> 38:51.429 so they weren't so frequent in February, I was lucky to see 38:51.431 --> 38:51.691 one. 38:51.690 --> 38:53.970 But these chimps hunt a lot. 38:53.969 --> 38:57.879 And if you look at the hunting success as a function of the 38:57.880 --> 39:02.620 numbers in the hunting party-- you can see on the top here a 39:02.617 --> 39:07.857 comparison of solitary and group hunts in the Ivory Coast in 39:07.864 --> 39:09.914 Gombe and at Mahale. 39:09.909 --> 39:14.379 So Gombe is Jane Goodall's study site in Tanzania, 39:14.380 --> 39:19.400 Tai is in the Ivory Coast, and Mahaleis a Japanese study 39:19.398 --> 39:20.218 site. 39:20.219 --> 39:23.279 And you can see that they hunt quite a bit in Tai, 39:23.278 --> 39:26.088 and they do a lot more group hunting there. 39:26.090 --> 39:29.390 And if you look at the impact of group size on capture 39:29.393 --> 39:31.053 success, you can see that the more 39:31.047 --> 39:32.557 chimps that are hunting in the group, 39:32.559 --> 39:36.349 the more likely it is they are to make a kill, 39:36.349 --> 39:41.409 the longer the hunt lasts, and the greater the degree of 39:41.407 --> 39:44.347 collaboration during the hunt. 39:44.349 --> 39:48.839 So this is team behavior, where individuals have roles, 39:48.840 --> 39:51.340 and they learn to play a team role, 39:51.340 --> 39:54.600 and they learn to do what is good for the team so that the 39:54.601 --> 39:56.491 team will have greater success. 39:56.489 --> 40:01.249 If we look at success as a function of group size, 40:01.250 --> 40:02.630 you can see that the net--and by the way, 40:02.630 --> 40:06.880 this is now measured in net benefit in calories, 40:06.880 --> 40:11.100 and in order to calculate that, you have to be able to estimate 40:11.097 --> 40:14.427 how many calories a chimpanzee is putting out, 40:14.429 --> 40:16.949 if it's running along the ground or climbing a tree or 40:16.952 --> 40:18.002 something like that. 40:18.000 --> 40:20.370 And so this is taken from exercise physiology, 40:20.371 --> 40:23.061 the estimates are taken from exercise physiology. 40:23.059 --> 40:26.469 And then you can figure out how many calories are there in a 40:26.471 --> 40:28.381 colobus monkey of a given size. 40:28.380 --> 40:32.970 And it turns out that the right number to have--if you are a 40:32.965 --> 40:37.935 hunter you do better up until you get to a group size of five; 40:37.940 --> 40:40.940 after that you have to share with too many others and the 40:40.943 --> 40:42.233 payoff isn't so great. 40:42.230 --> 40:45.560 If you are a bystander, it's pretty much the same, 40:45.559 --> 40:47.189 the curve mimics that. 40:47.190 --> 40:49.790 And if you are a latecomer who's coming in, 40:49.789 --> 40:53.319 then normally you don't get too much of what is caught. 40:53.320 --> 40:58.600 So in this particular hunt, Brutus led the hunt and he made 40:58.603 --> 40:59.883 the capture. 40:59.880 --> 41:03.030 So that was the chimp that you saw going up as the blocker; 41:03.030 --> 41:04.330 that was Brutus. 41:04.329 --> 41:06.579 Brutus, by the way, was the oldest male in the 41:06.583 --> 41:09.543 group, and he also had a couple of strategic innovations. 41:09.539 --> 41:12.579 Brutus had figured out that in competition with neighboring 41:12.576 --> 41:14.326 groups-- and chimps do have wars with 41:14.327 --> 41:17.067 neighboring groups-- if you were in a state of being 41:17.070 --> 41:20.710 the weaker group and you were being confronted by a big one, 41:20.710 --> 41:23.380 what you would do is you would go over and you would display at 41:23.380 --> 41:23.940 the border. 41:23.940 --> 41:26.190 They would come in, and then you would quiet down-- 41:26.190 --> 41:28.530 and he got them all to quiet down--and the pack of males 41:28.527 --> 41:31.077 would run around the back and steal a female from the back of 41:31.079 --> 41:31.929 the other group. 41:31.929 --> 41:34.759 And this caused such chaos and disarray that they would 41:34.755 --> 41:37.785 normally be able to overpower a larger neighboring group by 41:37.791 --> 41:39.521 using this sneaking behavior. 41:39.518 --> 41:42.228 Just think of how much strategic thinking it requires 41:42.226 --> 41:44.096 to figure something like that out. 41:44.099 --> 41:46.309 And that was Brutus; he was a smart guy. 41:46.309 --> 41:47.779 And there he is right there. 41:47.780 --> 41:53.280 So Macho was contending with Brutus to be an alpha male-- 41:53.280 --> 41:56.280 there are two or three different ways you can define 41:56.275 --> 41:59.985 group hierarchy in chimps-- and Macho was trying to be 41:59.987 --> 42:01.067 group leader. 42:01.070 --> 42:03.540 And he was actually a very collaborative hunter. 42:03.539 --> 42:06.749 It's one way to sort of weasel your way into power is to 42:06.751 --> 42:07.921 collaborate a lot. 42:07.920 --> 42:09.990 And Rousseau never made it. 42:09.989 --> 42:13.259 And Rousseau actually got attacked by a leopard once, 42:13.255 --> 42:16.705 who managed to rip off half of his scrotum, and Rousseau 42:16.708 --> 42:22.128 survived that; pretty amazing. 42:22.130 --> 42:25.730 So this is the function--it's sort of a hill-shaped or 42:25.728 --> 42:29.058 hump-shaped function of success versus group size, 42:29.056 --> 42:32.246 with the best group size being around five. 42:32.250 --> 42:37.140 And then if you look at who shares in the capture, 42:37.135 --> 42:42.515 what you see is a breakdown where there is considerable 42:42.518 --> 42:45.508 sharing which is going on. 42:45.510 --> 42:48.710 And you can see that bystanders, who are often 42:48.708 --> 42:51.758 females, are eating meat often after a 42:51.759 --> 42:56.789 capture for nearly half an hour, and the captor usually gets 42:56.793 --> 42:57.853 most of it. 42:57.849 --> 43:00.919 But it's very interesting how much gets shared. 43:00.920 --> 43:05.880 And if you look at the kind of hunt that was involved, 43:05.880 --> 43:08.660 you can see interestingly, if you classify the hunts by 43:08.657 --> 43:11.127 kind of a primitive, not terribly well-coordinated 43:11.128 --> 43:13.118 hunt, which is a half-anticipation, 43:13.121 --> 43:15.271 through a single-anticipation hunt, 43:15.268 --> 43:18.278 where one of the chimps is actually successfully 43:18.278 --> 43:21.988 anticipating where the colobus are going and manages to get 43:21.992 --> 43:25.162 over and block them off, to a doubled-anticipation hunt, 43:25.159 --> 43:26.639 which is even more sophisticated, 43:26.639 --> 43:30.899 you can see that there is a reward in terms of amount of 43:30.900 --> 43:35.550 meat eaten for participating in a more sophisticated hunt. 43:35.550 --> 43:36.720 Okay? 43:36.719 --> 43:41.099 43:41.099 --> 43:43.379 They pay a tax, and they are willing to pay a 43:43.378 --> 43:44.878 tax to belong to the group. 43:44.880 --> 43:47.760 The tax that they pay is by sharing what they capture. 43:47.760 --> 43:50.770 43:50.768 --> 43:55.588 So there is an anticipation of being willing to give up 43:55.592 --> 43:58.632 personal gain for group benefit. 43:58.630 --> 44:03.060 And there's another thing going on here, and that is that quite 44:03.056 --> 44:06.766 a bit of the reward of bystanders is trading food for 44:06.768 --> 44:07.338 sex. 44:07.340 --> 44:12.890 So both taxes and prostitution appear to be present in 44:12.885 --> 44:14.345 chimpanzees. 44:14.349 --> 44:15.929 Now what about learning? 44:15.929 --> 44:19.549 Well this is the frequency of ambushes that are used by 44:19.550 --> 44:21.430 hunters of different ages. 44:21.429 --> 44:24.439 Ambushing is, you know, a moderately 44:24.440 --> 44:26.420 sophisticated tactic. 44:26.420 --> 44:29.000 It's not quite as sophisticated as a big group hunt, 44:29.001 --> 44:31.731 but it's an indication that, you know, young chimps are 44:31.733 --> 44:33.003 learning how to hunt. 44:33.000 --> 44:34.810 And this is how old they are. 44:34.809 --> 44:38.109 And the interesting thing about this is--well of course they're 44:38.106 --> 44:40.276 weaned at five; so they aren't really going to 44:40.284 --> 44:42.924 start participating until they get there, and they don't do too 44:42.918 --> 44:44.828 much hunting before they become teenagers. 44:44.829 --> 44:47.199 But then let's suppose they start playing here, 44:47.204 --> 44:49.274 they start going out and hunting here. 44:49.268 --> 44:52.498 It takes them until they're thirty-five-years-old before 44:52.496 --> 44:54.196 they really hit their peak. 44:54.199 --> 44:57.339 It takes them about twenty years to learn how to be an 44:57.342 --> 44:58.472 effective hunter. 44:58.469 --> 45:00.289 That's pretty amazing. 45:00.289 --> 45:03.389 45:03.389 --> 45:05.909 And if you look, if you break this down by 45:05.911 --> 45:08.681 half-anticipations and full-anticipations, 45:08.679 --> 45:12.889 what you see is the frequency with which full-anticipations 45:12.894 --> 45:17.694 are practiced in hunts is lower, and it takes a long time before 45:17.692 --> 45:21.182 they get up to the level of half-anticipations. 45:21.179 --> 45:24.549 So these are categories of sophistication in hunting, 45:24.552 --> 45:27.992 and it takes a long time to get more sophisticated. 45:27.989 --> 45:34.049 So to sum up on the behaviors that are involved in foraging 45:34.052 --> 45:40.432 and hunting, I've really given you today two paradigms for how 45:40.429 --> 45:42.729 to think about it. 45:42.730 --> 45:45.100 One of them is the marginal value theorem, 45:45.099 --> 45:48.669 which tells you how you should decide when to stop hunting in a 45:48.670 --> 45:51.380 certain place, or when to stop copulating in a 45:51.378 --> 45:53.848 certain place, and go off to find either food 45:53.847 --> 45:56.707 or mates in another place; and that works in a spatial 45:56.710 --> 46:00.070 situation where you have to move some distance before you get to 46:00.074 --> 46:00.774 the patch. 46:00.768 --> 46:04.368 And the other paradigm that I've given you is simply the 46:04.367 --> 46:08.287 cost-benefit analysis of how much caloric reward do I get out 46:08.293 --> 46:11.633 of hunting in a group versus hunting by myself? 46:11.630 --> 46:15.600 And you can see that if you do that cost-benefit analysis, 46:15.599 --> 46:20.969 it turns out that organisms don't do it perfectly, 46:20.969 --> 46:24.959 but they do get better and better at approximating the 46:24.960 --> 46:26.240 optimal payoff. 46:26.239 --> 46:30.799 So that tells us that hunting is basically microeconomics. 46:30.800 --> 46:33.400 It's a very short-term, kind of selfish behavior, 46:33.400 --> 46:37.140 where the group hunting behavior, which looks like it 46:37.137 --> 46:42.217 might not be so selfish, in terms of caloric reward is 46:42.224 --> 46:45.664 quite selfish, and it looks like it's 46:45.659 --> 46:47.129 long-term selfish. 46:47.130 --> 46:52.670 So the sharing is stabilizing relationships that are paying 46:52.666 --> 46:58.676 off in terms of future hunts and future sexual opportunities. 46:58.679 --> 47:02.219 We've seen that predation is shaping both the behavior and 47:02.224 --> 47:03.784 the morphology of prey. 47:03.780 --> 47:06.270 We saw that with both conspicuous coloration, 47:06.268 --> 47:08.868 aposematic coloration, which is advertising the 47:08.871 --> 47:12.291 presence of poisons; and we saw it with crypsis, 47:12.291 --> 47:16.491 which is cryptic coloration, which is hiding and looking 47:16.492 --> 47:18.252 like something else. 47:18.250 --> 47:21.810 We saw that aggressive mimicry illustrates an important 47:21.809 --> 47:25.429 evolutionary trend, which is that every available 47:25.431 --> 47:30.001 opportunity will eventually be seized by some species evolving 47:29.996 --> 47:31.416 into that niche. 47:31.420 --> 47:34.990 And we saw it with fireflies, and then I told you about 47:34.994 --> 47:37.844 cleaning wrasses in saber-tooth blennies. 47:37.840 --> 47:41.600 And then finally I showed you the cooperative hunting, 47:41.599 --> 47:43.939 complex example in chimpanzees. 47:43.940 --> 47:48.430 It requires strategic thinking; it requires a kind of teamwork 47:48.434 --> 47:53.184 that really is only attained in fairly complex organisms. 47:53.179 --> 47:55.639 I'll mention one other, which is deeply cool, 47:55.635 --> 47:58.365 and that is the bubble-nets of humpback whales. 47:58.369 --> 48:01.579 Who has heard of a humpback whale bubble-net? 48:01.579 --> 48:03.099 A few. 48:03.099 --> 48:06.999 Humpback whales hunt in family groups in Alaska and down near 48:07.001 --> 48:10.711 Antarctica, and they hunt for fish that are in schools. 48:10.710 --> 48:11.890 They hunt herring. 48:11.889 --> 48:15.569 And a mama humpback will go down below a group of herring, 48:15.570 --> 48:19.890 and she'll start up in a spiral, just at the right rate, 48:19.889 --> 48:22.909 so that the bubbles that she's blowing out of her nose will all 48:22.913 --> 48:25.593 rise up in a curtain to the surface around the school of 48:25.594 --> 48:28.464 herring at the same time; so a curtain of bubbles, 48:28.463 --> 48:31.553 that looks like a net, is going up around the school 48:31.552 --> 48:32.402 of herring. 48:32.400 --> 48:36.150 And at the same time she gives a few little signals with her 48:36.150 --> 48:39.380 complex calling behavior, and the whole family of whales 48:39.378 --> 48:41.408 will dive down, and they'll come up as a group 48:41.414 --> 48:43.564 and they'll open their mouths underneath the school of 48:43.563 --> 48:45.333 herring, and all of the mouths will 48:45.333 --> 48:47.313 emerge from the water at the same time. 48:47.309 --> 48:50.509 They can take out an entire school of herring with one bite 48:50.512 --> 48:51.012 apiece. 48:51.010 --> 48:55.350 They're taking in maybe two or three tons of fish when they do 48:55.351 --> 48:56.841 this; like five or six whales doing 48:56.840 --> 48:58.240 it all at once, they're right next to each 48:58.242 --> 48:58.552 other. 48:58.550 --> 49:05.010 So there are very interesting examples where other species, 49:05.010 --> 49:09.040 that appear to have complex cognition and good signaling 49:09.039 --> 49:11.239 capacity, learn really rather 49:11.244 --> 49:12.924 sophisticated behavior. 49:12.920 --> 49:17.530 And I think the chimpanzee hunt is one of the most interesting. 49:17.530 --> 49:19.790 I'm sorry it's so bloody, and I do hope you enjoy your 49:19.789 --> 49:20.129 lunch. 49:20.130 --> 49:43.000