BIOS 3010: Ecology Lecture 6: Processes: Interspecific competition • Lecture summary: – Definition & examples. – Lotka-Volterra model: Semibalanus balanoides James P. Rowan, http://www.emature.com • Structure. • Competition coefficients. • Predicted outcomes. Chthamalus stellatus Dr. S. Malcolm BIOS 3010: Ecology Alan J. Southward, http://www.marlin.ac.uk/ Lecture 6: slide 1 2. Interspecific Competition: – Like intraspecific competition, competition between species can be defined as: • "Competition is an interaction between individuals, brought about by a shared requirement for a resource in limited supply, and leading to a reduction in the survivorship, growth and/or reproduction of at least some of the competing individuals concerned" – For example, Connell's 2 species of barnacle (asymmetrical interference/contest) in Fig. 8.2 and slides – Gause's Paramecium species of Fig. 8.3 and, – Tilman's diatoms (exploitation/scramble) of Fig. 8.5 – Connell's "the ghost of competition past" = the current product of past evolutionary responses to competition! Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 2 3. Basic outcomes of competition: • These interactions illustrate the two basic outcomes of competition: – 1) coexistence: • if two competing species coexist in a stable environment, then they do so as a result of niche differentiation (of their realized niches) = character displacement (Figs 7.18, 8.23 & 8.25a) – 2) competitive exclusion - the "competitive exclusion principle" or "Gause's principle:” • if there is no niche differentiation, then one competing species will eliminate or exclude the other. • Thus exclusion occurs when the realized niche of the superior competitor completely fills those parts of the inferior competitor's fundamental niche which the habitat provides. • see Fig. 7.4 of exclusion in reed species. Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 3 1 4. The Lotka-Volterra model of interspecific competition: • Based on the logistic equation that describes sigmoidal population growth as a result of intraspecific competition: dN/dt = rN((K-N)/K) – (after Volterra, 1926 & Lotka, 1932) with the inclusion of the competition coefficients α and β we can represent population size changes for the two competing species as: dN1/dt = r1N1((K1-N1-α N2)/K1) – and dN2/dt = r2N2((K2-N2-β N1)/K2) Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 4 5. Competition coefficients: • The competition coefficient α is the effect on species 1 of species 2 (also written as α12): – If α <1 then interspecific competition has less impact than intraspecific competition. – If α >1 then interspecific competition has more impact. • Conversely, β is the effect on species 2 of species 1 (also written as α21): – N1 & N2 are the population sizes of species 1 & 2. – r1 & r2 are intrinsic rates of natural increase for spp. 1 & 2. – K1 & K2 are the carrying capacities for species 1 & 2. Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 5 6. Lotka-Volterra competition model - zero isoclines: – Zero population growth isoclines (dN/dt = 0) are shown in graphs of N2 on the y-axis plotted against N1 on the x-axis in Figs. 8.7 and 8.9. • When this is true for species 1, then r1N1(K1-N1-αN2) = 0 and K1-N1-αN2 = 0 • Therefore N1 = K1-αN2 • When N1 = 0, N2 = K1/α (the result of pure interspecific competition at A in Fig. 8.7a) • When N2 = 0, N1 = K1 (the result of pure intraspecific competition at B in Fig. 8.7a) to give the zero isocline of Fig. 8.7a Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 6 2 7. Four outcomes of the LotkaVolterra competition model: • From Figure 8.9 the 4 outcomes are expected to be: – 1) species 1 wins (competitive exclusion) (8.9a) • species 1 is a stronger interspecific competitor (K1 >K2/β, therefore K1β >K2) even though intraspecific competition within species 1 is stronger than the interspecific effect of species 2 (K1/α > K2, therefore K1 > K2α) • (converse of 1) – 3) either species 1 or species 2 wins (8.9c) • (interspecific competition greater in both species than intraspecific competition - the outcome depends on starting densities) – 4) coexistence (8.9d) • (both species have less competitive effect on the other species than they do on themselves: K1 > K2α, and K2 > K1β - gives a stable equilibrium) Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 7 Figure 8.2: Intertidal distribution of two barnacle species Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 8 Figure 8.3: Intra- and interspecific competition in Paramecium spp. Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 9 3 Figure 8.5: Competition between diatoms & silicate availability Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 10 Figure 7.18 (3rd ed.): Character displacement (mandible variation) in ant communities Data for Veromessor pergandei Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 11 Figure 8.23: Character displacement (gill rakers) in sticklebacks benthic Dr. S. Malcolm limnetic BIOS 3010: Ecology Lecture 6: slide 12 4 Figure 8.25a: Character displacement of freshwater snail shell lengths Hydrobia ulvae Hydrobia ventrosa Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 13 Figure 7.4 (3rd ed.): Asymmetrical competition between cattail species in Michigan Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 14 Figure 8.7: Zero isoclines of the Lotka-Volterra competition equations for species N1 and N2 Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 15 5 Figure 8.9: Lotka-Volterra competition model outcomes Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 16 Barnacles, July 2006, Kintyre, Scotland Dr. S. Malcolm BIOS 3010: Ecology Lecture 6: slide 17 6