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Community Ecology Niche theory and guilds Click here for supplemental materials for today (PDF) Outline: 1. Definition of niche: response functions vs. resource utilization functions A. Grinnell: emphasis on a species’ "place" (more than just locational) B. Elton: emphasis on a species’ role C. Hutchinson: emphasis on resources used by species i. n-dimensional hypervolume ii. fundamental vs. realized niches iii. ecological release 2. So how can species coexist? Answer: niche partitioning A. Limiting similarity B. Character displacement i. the Hutchinsonian 1.3 size ratio 3. Niche vs. guild vs. functional group Terms/people: niche Elton limiting similarity functional niche 1.3 character size ratio n-dimensional hypervolume guild functional group Hutchinson MacArthur R.B. Root empty niche realized niche niche differentiation character displacement ecological release Grinnell niche Joseph Grinnell (1914) to denote a species’ "place" in the environment (i.e., set of environmental conditions that meet a species’ life-history requirements) Difference between niche and habitat Chas. Elton (1927) emphasized the role of a species in the environment Gene Odum’s distinction between an organism’s “address” (habitat) and its “profession” (niche) Do empty niches exist? G. Evelyn Hutchinson (1957) "n-dimensional hypervolume" 1. fundamental (pre-interactive) niche - potential niche 2. realized (post-interactive) niche - actual niche Example from Orians and Willson (1964) involving Red-winged Blackbirds (Agelaius phoeniceus) and Yellow-headed Blackbirds (Xanthocephalus xanthocephalus) “Ecological release” – mongoose example Ecological niches can thus be defined in terms of: -response functions: how species are distributed on environmental gradients with respect to limitation and optimal performance (a physiological view, prevalent among plant ecologists), i.e., a species’ response to the environment (Whose ideas follow this?) -resource utilization functions: how species use resources (Whose ideas follow this?) So if a niche can only be occupied by one species, but if resources are limited such that competitors must share niche space, how similar in terms of niche can two species be and still coexist? The competitive exclusion principle states that coexistence hinges on niche differentiation (a.k.a. niche partitioning). Consider the words of Gause (1934), when discussing the competitive exclusion principle: "...as a result of competition two species hardly ever occupy similar niches, but displace each other in such a manner that each takes possess of certain kinds of food and modes of life in which it has an advantage over its competitor." This is niche partitioning. But since most organisms are rare and secretive, how do we observe this? We usually do so only indirectly, via morphology (Ricklefs and Travis 1980; see handout): limiting similarity character displacement Hutchinsonian size ratio Robert MacArthur (1958) examined niche overlap in a group of five sympatric warbler species (Bay-breasted, Myrtle, Blackburnian, Cape May, and Black-throated Green) in New Hampshire that were the same size and ate the same arthropod species in the same tree!: if they were so similar, how could they coexist? - found that the birds partitioned the niche space physically Similar but distinct terms: Guild Functional group - References: NOTE: the Proceedings of the National Academy of Sciences vol. 106 suppl. 2 (2009) has a special colloquium devoted to the niche concept. Abrams, P.A. 1983. The theory of limiting similarity. Annual Review of Ecology and Systematics 14:359-376. Alley, T.R. 1982. Competition theory, evolution, and the concept of ecological niche. Acta Biotheoretica 31:165-179. Austin, M.P. 1999. 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Journal of Experimental Marine Biology and Ecology 185:219-233. Hurlbert, S.H. 1978. The measurement of niche overlap and some relatives. Ecology 59:67-77. Hutchinson, G.E. 1957. Concluding remarks. Cold Spring Harbor Symposium on Quantitative Biology 22:415-427. Jaksíc, F.M. 1981. Abuse and misuse of the term "guild" in ecological studies. Oikos 37:397400. Landres, P.B. 1983. Use of the guild concept in environmental impact assessment. Environ. Manage. 7:393-398. Loehle, C. 1998. Height growth rate tradeoffs determine northern and southern range limits for trees. Journal of Biogeography 25:735-742. MacArthur, R.H. 1958. Population ecology of some warblers of northeastern coniferous forests. Ecology 39:599-619. MacMahon, J.A., D.J. Schimpf, D.C. Andersen, K.G. Smith, and R.L. Bayn, Jr. 1981. An organism-centered approach to some community and ecosystem concepts. J. Theoret. Biol. 88:287-307. Murray, M.G., and D. Brown. 1993. 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