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    Cap 1 Behavioral Ecology Krebs Davies 2006 PDF

    Diunggah oleh

    Tatiana Ferreira

    Hak Cipta:

    © All Rights Reserved
    Unduh sebagai PDF atau baca online dari Scribd
    Tandai sebagai konten tidak pantas
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    Chapter 1 The Evolution of Behavioural Ecology John R. Krebs & Nicholas B. Davies L.1 Observations and questions All natural bistory observations begin with a question. At first our curiosity nay be satislied simply by knowing the species name of the animal we are watching, Then we may want to discover what il is doing and to understand why it is bebaving in a particular way. in 1978, we began the first edition of this book by asking the reader to observe a bind, such as a staring (Sternus vulgaris), searching in Ihe grass for food, The starling walks along. pausing every new and then 1 probe into the ground. Sometimes it succeeds in isding a prey item, such asa beetle larva, and eventually, when it bas collected several prey, ir llies back to the nest to feed its hungry brood. For students of behavioural ecology, @ whole host of questiems came te mind as they observe this behaviour, The first set of questions omrere the way the bird feeds, Why bas it chosen that particular piace t0 foeage? Why is it alone rather than in a flock? What determines its choice ol loragimg path? Does it colle every item of food it encounters or is it selective fet prey type ur size? What influences its decision to stap coliccting lond and By back 1 icod its chicks? Another set of questions emerges when we follow fhe sarting back to the nest. Why has it chasen this sile? Why this brood size? How do the two parent starlings come to an agreement aver how muck work each puts ino offspring care? Why are the chicks begging so nomsly? Ane they each simply signalling Uheir awn degree of hunger or ate they compeung for food? Why such costly begging behaviour? If we observed the stediing ower s longer period of time then we may ask abour what determines how much effori it puls isto reproduction versus its awn maintenance. about the fectors influencing the timing ol its seasonal activities, its choice of maze. end so en. Behavioural ecology provides @ [ramesork for answering Unese kinds of questions. Tt combines ideas from evelusion, ecology and behaviour anil hay emerged [com five schools of thought. developed primarily in the 1960s and carly 1970s. We diseuss them in tact to provide # brick history of the subject and tu show how (he ideas have cvelved in the last 20-30 years, and we point out how this book reflects recent developments, 4 CHAPTER Ff 1.2 Tinbergen’s four questions Hinberven (1963) showed that there are four ways of answering Ihe qucstiat swhy?’ in biology, Retuming to our starling, if we asked why it forged [ea particular way we could answer as folluws. 1 Interms of function, namely how patch choice and prey choice comtribuae 1o the survival of the bird and its offspring 20 Interms of eausaiio, namely the proximate laciors which caused (he Feat to select a foraging site or prey type. These may inchade exes to prey abrutad such as type of soil or vegetation, or Lhe activities of other birds. 3) Interns of developmmur. This answer would be concerned with the role € genctic predispositions and learning in an individual's decision making 40 Intermsol cunwionary kispry, namely how starling bebaviour has cvolst from. its ancestors, This answer might inelude an investigation af how tae starling family has radiated to fill particular ecological niches and the influcnae of competition from other animals on Lae evelution of starling behaviour 2 morphology (e.g. bill size. body size. ‘Tinbergen’s studies on gulls aimed to combine the four kinds of answes and he emphasized the need to study animals in their natural surrounding, namely those where their behaviour had evolved, Ne championed the usc a the fickd ay a natural isberatory Lor observations and controlled experi and showed how ideas can be tested by collecting quantitative data on te haviour pattems ({¢.g. Tinbergen. 1953, 1972). Tinbergen’s legacy is evidersiamr’ curren! field studies of behaviour, many of which use simple experimenw m measure the cests and benefits of traits. A good example is the classic mit manipulation éxpetiment by Andersson and Moller io investigate mate chaie inswidowbirds and swallows discussed by Ryan in Chapter & However, carly sadics in behavioural ccology often focused on Lunct and tended to jenore the other three questions. A caricature af behaviour st in the 1990s & one where researchers imagined theiranimals as litte mact blindly following fixed action patiems in responses to external simul caricature [rom the early days of behavioural ecology and sociabiolagy in she iy animals as scheming 1actiont @ Up The costs and benefits of every conceivable course of action aa always chousing the best one. Current wark is leading to an intermedia position. While we expect selection Lo favour mechanisms that maximize ® individual's fitness, we must. recognize that mechanisms both consirain am serve behavioural outcomes A good example to illustrate this point is Lotem’s recemt studies of bow hosts come to recognize a cuckoo egg in their nest, The cuckoo, Chests coarm is a brood parasite which exploits varions small birds as hosts to raie # ollspring. The female cuckoo lays just ane egg per host nest. The cuckoo st hatches first, whereupon it ejects the host’s eggs aver the side of the nes becoming the sole occupant. Given the cost of parasitism, it is mor sun 1970s is the opposite extreme of regan weigt EVOLUTION OF BEITAVIOURAL ECOLOGY 5 that many hosts have evolved defences such as rejection of add eggs in their nest. Nevertheless, the puzsic is that egg rejection rarely reaches 180% in the host population and, furthermore, Hosts never reject the cuckoo chick. A consideration of mechanisms may help te solve botl: puzzles, Experiments show that the dcte i: hosts _imvolves Jeaming the chatacicristics of their own eggs the first time they breed and then rejecting eggs which differ from this learned set (Lotem etal, 1995). This makes it unlikely that the host population will evulve 100% rejection of cuckoo eggs because hosts which are parasitized during their first breeding attempt will learn the cuckoo egg as part of their own sct_ Nevertheless. the learning rule works quite well and leads hosts lo reject many parasite epns. At the chick stage, however, a learning rule docs tess well than a rule ‘accept any chick in my nest’. ‘This is because (here is a considerable cost of misimprinliag: any host parasitized in its first attempt weuld leacn only the cuckoo chick as and would then subscquently reject its own young in future, unparasitized, brouds (Lorem. 1993). The main message {rom this sLudy is that it is net very feuiliul to discuss the evolution of ‘rejection’ without specifying the mech- ons because these will deiemmine ihe costs and benefits involved. Studies {n. 1975, Wilson predicted the demise of erhology, with mechauisms becom saciobiology, This prediction was fulfilled until recent years, when there bas been a welcome renewed interest in linking mechanism and function. We have marked this change by devoring the first section of this volume to his truithul imerchange. For example. Giraldeay (see Chapter 3) shows how research on foraging behaviour has stimulated new questions abous learning and memory mechanisms, and Sherman et al. isce Chapter 4) point out common leatures of recognition mechanisms of kin, mates and predavors and discuss their functional significance. is OWN, 1.3 Ecology and behaviour Fyen belore Darwin, biologists olien interpreted morphological adaptations in relation to The cnvironment in which Lhe species lived. Darwin's achievement was to show how these could arise witout a Creator. Once the early cthoingists, such as Loxenzand Tinbergen, had demonstrated Laat behaviour patterns were olten as characteristic of a species as its morphological features, attempts were made to correlate diflerences between species in behaviour with differences im ceological factors, such as habitat, food aud predation. a pioneering suudy was that by Cullon (1957), whe was a student of Tinbergen. She Interpreted the reduced anti-predator behaviour of kitliwake gulls, Risse midactyla, compared ta the ground-nesting gulls, in retation to their safer nest sites on step

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