Evolution Behavioural Ecology is the scientific study of behavioural phenomena in the light of ecological and evolutionary theory 1. A refresher in evolution by natural selection 2. What is behaviour in the light of evolutionary theory 3. How did behavioural ecology develop as a discipline? 4. There is evolution in action all around us PSYCH 3T03 Lecture 02 Animal Behavior – Chapter 1 Common misunderstandings in evolution 1. Evolution is not random – mutation is random, selection is not 2. There is no ‘aim’ to evolution – whatever survives survives 3. What survives is not necessarily what is ‘best’ for long term survival of the species 4. No species is ‘more evolved’ than any other – same ~3.7 billion years of evolution 5. Evolution is not necessarily slow – big changes can occur in few generations 6. Not all evolutionary change is adaptive – neutral processes such as drift Three things that behavioural ecology is not 1. Behavioural Ecology is not behavioural genetics 2. Behavioural Ecology is not behavioural determinism 3. Behavioural Ecology is not primarily about humans * This course will not be about humans Three things that behavioural ecology is 1. Behavioural Ecology is Adaptationist We will be looking at adaptive traits 2. Behavioural Ecology is Selectionist Natural selection is the force driving adaptation 3. Behavioural Ecology is Comparative We will be comparing across different taxa and sexes Adaptation The evolutionary process by which organisms fit better into their environments or habitats. A physical or behavioual trait maintained and evolved by natural selection that enhances the probability of survival and reproduction Adaptation- Bird Flight Wings – large chest muscles Bones – hollow, lightweight Aerodynamic – body structure Feathers – help birds fly Powerful heart and respiratory system – extract more oxygen and power flight Why study Behavioural Ecology? 1. Behavioural Ecology benefits us We use animals and plants in many different ways Protect our crops from pests, improve the welfare of pets/livestock, find food etc. 2. Behavioural Ecology benefits the natural world Know what animals need and when/where they need it Monitor how ecosystems are changing due to habitat and climate change 3. Behavioural Ecology is interesting >5000 mammals, >30,000 fishes, >40,000 spiders, >400,000 beetles, etc. Who studies Behavioural Ecology? Evolutionary Biologists Population and Community Ecologists Zoologists Psychologists Physiologists Geneticists Neuroscientists Endocrinologists Anthropologists Conservation Biologists Wildlife Resource Managers Animal Breeders And where do they publish? Nature Science Proceedings of the Royal Society Animal Behaviour Behavioral Ecology Behavioural Ecology & Sociobiology Hormones & Behavior Ethology Behavior Ecotoxicology Neuroscience Experimental Physiology And many more…. How did the field develop? Behavioural Ecology unites two fields: 1. Ethology The study of behaviour under natural conditions 2. Evolutionary Biology The study of how life develops through natural selection Ethology – studying behaviour Niko Tinbergen Konrad Lorenz Karl von Frisch Nobel Prize in 1973 The great insight of the classical ethologists was that they realized that just as each animal species has its characteristic anatomy, physiology, biochemistry so too each species has its characteristic behaviour. Behavioural traits can be analyzed in the same way as physical traits, including the study of their evolution. A curious naturalist Obliged to the public Valued communication Science is a social effort Richard Dawkins Marian Stamp Dawkins Iain Douglas Hamilton Desmond Morris How did the field develop? Behavioural Ecology unites two fields: 1. Ethology The study of behaviour under natural conditions 2. Evolutionary Biology The study of how life develops through natural selection Evolutionary Biology Charles Darwin On the Origin of Species 1859 Alfred Russell Wallace Natural selection occurs because 1. The is variation within populations Individuals differ in traits that affect their chance of survival / reproduction 2. Variation is heritable Successful traits can be passed on to the next generation 3. Reproductive success can differ within a population Some individuals will have more offspring that others Gregory 2009. Evo Edu Outreach. 2: 156-175. Remember…. when differential survival and reproduction are related to individual differences and these individual differences are heritable, evolution can ensue. We say evolution "can" ensue, not that it will ensue, because even if all 3 conditions are fulfilled, selection is often stabilizing: if a trait's is already optimal (because of past selection), current selection tends to prevent it from evolving ("drifting") away from that optimum. Evolution in action Anyone know of any examples of contemporary examples of evolution? Share them in the chat Pollution adapted peppered moths SARS-COV-2 variants Running cane toads in Australia Antibiotic resistant ‘superbugs’ Anolis lizards adapted to urban substrates Shrinking fish because of over fishing Tuskless elephants Darker owls in Finland due to climate change Evolution in action – Darwin’s Finches This is the adaptive radiation that most influenced Darwin : By standard morphological criteria of taxonomy, these 11 species are one another’s closest relatives (now confirmed by molecular methods), but they are much more diverse in bill morphology, diet and behaviour than is typical within a genus of songbirds. Arriving first at the newly formed volcanic islands of the Galapagos, they “radiated” to fill unoccupied ecological niches that other songbird genera occupy in South America and elsewhere. The Grants in The Galapagos 50 km Rapid evolution of beak morphology in the medium ground finch (Geospiza fortis) G. fortis has been studied on Isla Daphne for > 50 years by Rosemary & Peter Grant Unusual aspects of this study 1. Long term study. 2. Every individual was marked and tracked. So LRS was known and the traits beneficial for fitness could be assessed. 3. Natural environmental perturbations in the form of droughts were “naturalistic experiments” or manipulations allowed the Grants to observe natural selection in a relatively short time period. Excellent popular account of their work: J Weiner (1994) The beak of the finch. NY: Knopf. Beak size/depth Birds with large beaks eat large hard seeds Birds with medium beaks eat both small soft and large hard seeds Birds with small beaks eat small soft seeds 1976 was an unusually dry year on Isla Daphne; 1977 was worse. Over < 2 years, finch food (seed) abundance plummeted, and the finch population crashed. Not only were seeds rarer, but species composition of the seed bank changed. Before the drought the most abundant were small, soft Portulaca seeds. After the drought, most of what remained were large hard Tribulus seeds… which not all G. fortis were capable of cracking open. Result : Directional selection on beak size in G. fortis as a result of differential survival of the drought N of birds 751 90 1976 population 1978 survivors 1976 Number of individuals 9.4 mm 10.2 mm 5 7 9 11 Beak depth (mm) 1978 13 Average beak depth 9.4 mm 10.2 mm Recall : When differential survival and reproduction are nonrandomly related to individual differences and these individual differences are heritable, evolution can ensue. Survival (hence reproduction) was obviously selective with respect to beak dimensions. But is beak size demonstrably heritable? Yes, it is All the necessary conditions for evolution were met, and evolution of bill size indeed occurred: ensuing generations of Daphne Island G. spiza had bigger bills than the 1976 pre-drought population. Boag & Grant 1981. Science Review Evolution Is a change in the genetic composition of a population over successive generations. Driven by natural selection acting on the genetic variation in reproductive success among individuals (resulting from a heritable trait). Results in new species. Review Evolution-long term process Natural Selection 1976 9.4 mm Number of individuals is a process by which traits become more or less common in a population because these inherited traits lead to differential survival and reproduction of individuals possessing them. 1978 10.2 mm 5 7 9 11 Beak depth (mm) 13 Review Adaptation is a change in a state or a trait that increases the reproductive success of individuals that have it. Important to Remember 1. Evolution is not Natural Selection Evolution is gene frequency change - Evolution occurs across generations. Individuals do not evolve. 2. The major causes of evolution are mutation, genetic drift and natural selection. 3. Natural selection is differential reproduction Differential reproductive success is called FITNESS 4. Natural selection is the only evolutionary force predictably leading to adaptation (or the fit between an organism and its environment). Important to understand the following terms: Evolution Natural Selection Fitness Adaptation Evolution in Action: The Story Continues Grant PR & Grant BR (2006) Evolution of character displacement in Darwin’s finches. Science 313 : 224-226. Result : Directional selection on beak size in G. fortis as a result of differential survival of the drought N of birds 751 90 1976 population 1978 survivors 1976 Number of individuals 9.4 mm 10.2 mm 5 7 9 11 Beak depth (mm) 1978 13 Average beak depth 9.4 mm 10.2 mm Review MEDIUM GROUND FINCH WITH A SMALL BEAK 1982 NEW COMPETITOR:LARGE GROUND FINCH MEDIUM GROUND FINCH WITH A BIG BEAK Review Grant PR & Grant BR (2006) Evolution of character displacement in Darwin’s finches. Science 313: 224-226. Figure 2. Year-by-year variation in Geospiza fortis beak size on Daphne Major. The 1977 and 2004 “selection events” were both droughts that led to seed bank depletion and crashing finch populations due to starvation. But the selective effects on G. fortis beak dimensions were opposite in direction. The Grants argue that the critical difference was the presence of G. magnirostris in 2004 cf. absence in 1977. “Character displacement” Divergence in traits that is caused by competition between species (interspecific competition). CD means that the differences are exaggerated in sympatry, presumably as a result of a recent selective history of interspecific competition. CD caused bigger differences in beak dimensions between the medium ground finch (G. fortis) and the large ground finch (G. magnirostris) G. fortis- MGF G. magnirostris - LGF After 2005 40 20 % Individuals 8 In Each Size 40 class 20 8 10 12 1 4 Before 2004 10 12 Beak size (mm) 1 4 Review “Character release” In the absence of competition from the small ground finch, (G. fulginosa), the medium ground finch (G. fortis), on Daphne had smaller beaks than is typical on other islands. Greater overlap in beak size when each species found alone (allopatric). In the past, the medium ground finch was released from competition and had unusually small beaks. Any Questions? PSYCH 3T03 Lecture 02