BIOS 6150: Ecology Dr. Stephen Malcolm, Department of Biological Sciences

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1 BIOS 6150: Ecology Dr. Stephen Malcolm, Department of Biological Sciences Week 3: Intraspecific Competition. Lecture summary: Definition. Characteristics. Scramble & contest. Density dependence k-values Discrete & continuous models BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 1

2 2. Competition: Interactive process. Product of combined demand for resources. Leads to competition among individuals, either: intraspecifically or, interspecifically. Results in a negative outcome for all competitors. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 2

3 3. Definition of Competition: 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 (Begon et al., 2006, p. 132) The ultimate effect of competition on an individual is a decreased fitness contribution to the next generation (fewer offspring) compared with what would have happened had there been no competitors. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 3

4 4. Four characteristics of intraspecific competition: (1) Decrease in fitness: The ultimate effect of competition is a decrease in the fitness of all interactants (thus it is a -,- interaction): Often via decreased survivorship or fecundity. Fitness reduction must be measurable to conclude that competition occurred. (2) Limited supply of resources: The resource for which individuals compete must be in limited supply. (3) Reciprocity: Even if the detectable competition is either mostly one-sided, or balanced, it must be reciprocal and have a negative impact on both interactants (symmetrical and asymmetrical). (4) Density dependence: The probability of an individual being adversely affected increases with increasing competitor density (in contrast to density independent effects). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 4

5 5. Extremes of intraspecific competition: Scramble (exploitation) and contest (interference) competition were first described as simplistic extremes by Nicholson (1954) in Australia. Mortality (% or k competition due to competition) or survivorship is plotted against logarithm of initial density to show degree of density dependence. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 5

6 6. Scramble & Contest Competition: Scramble competition: Nicholson described scramble competition for dungflies competing for the limited resources of cow feces. Each member gathers a constant amount of resource at all densities. Thus at high density there is insufficient resource and the whole population dies (slope b = ). Contest competition: Where individuals of the population interfere or contest with each others abilities to harvest resources, some survive. Exact density-dependent compensation is thus described by a mortality slope b = 1. Figs. 5.1 & 5.2 from Begon et al. (2006) to show both kinds and negative effects of competition in single species populations. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 6

7 7. Density dependence: Competition also increases with population density when mortality may increase or survivorship may decrease (Fig. 5.2). The nature of density dependence can also change with increasing density from: density independence, through, undercompensating density dependence, to, exactly compensating density dependence, to, overcompensating density dependence (Figs 5.3, 5.4 & 6.5). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 7

8 8. Density dependence (continued): So intensities of both kinds of intraspecific competition increase with population density and change from density independence to density dependence. Thus density dependent birth and mortality rates lead to the regulation of population size at a stable equilibrium where births = deaths. This is the carrying capacity (K) at the population size sustainable by available resources as shown in Figs. 5.7 & 5.8. Density dependent population regulation generates the sigmoidal or S-shaped curve characteristic of intraspecific competition (see Fig 5.11). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 8

9 9. Density dependent growth: In addition to effects on numbers, competition also negatively influences growth: This in turn influences numbers through reduced per capita reproductive output. Rates of growth and rates of development can be reduced as shown in Figs 6.14 & 5.12: But the total population biomass can remain the same, despite individuals being smaller: The law of constant final yield (exact compensation) Reproductive allocation can also shift with changing resource availability (Figs & 5.15): Within genets, tiller growth was less variable and more regulated than the genets themselves. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 9

10 10. k-values and density dependent mortality: k-values of mortality due to competition can define competition according to the slope b of the relationship (Fig. 5.16) of k competition plotted against the logarithm of initial density (density before the effects of competition). b = 0 density independence. b < 1 undercompensating density dependence. b = 1 (contest) exact density dependent compensation. b > 1 overcompensating density dependence b = (scramble) overcompensating density dependence see Fig. 2-3 from Hassell (1976) of scramble and contest competition. k-mortality is shown in Fig 5.16 & k-fecundity in Fig BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 10

11 11. Discrete breeding season model of intraspecific competition: Using: R net reproductive rate N t population size at time t N t+1 population size at time t+1 In the absence of competition, the model describes population increase simply as: N t+1 = N t R and N t = N o R t This gives the exponential population growth across discrete generations as in Fig BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 11

12 12. Carrying capacity, limited resources and the effect of competition: At high density when the ratio of N t /N t+1 = 1 this is by definition the carrying capacity K. So in the presence of competition, the population rises to K as shown in Fig according to: N t+1 = N t R/1+(aN t ) where a = (R-1)/K so the unrealistic R in the first equation is now replaced by the more realistic R/(1 + an t ) as a and N t increase so does the effect of competition and R is decreased. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 12

13 13. Density dependence of the model: The k-value for mortality due to competition is thus the difference between log N t R and log N t R/(1+aN t ) and plotting these k values against log 10 N t (Fig. 5.20) shows that the model exactly compensates with a slope b = 1. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 13

14 14. Incorporation of variable density dependence with b: A more realistic model of competition that incorporates a range of competitive regulation was derived by Maynard Smith & Slatkin (1973) in which they simply added the slope b of the k-value plotted against log initial density: N t+1 = N t R/1+(aN t ) b in which b is the slope of mortality (k) against population size (log 10 N t ) and, a substitutes for (R-1)/K as before (see Figs & 6.26 for real data). Also generates realistic ranges of population fluctuations (Fig. 5.22). --- equation 5.18 (p.148) BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 14

15 15. Continuous breeding - the logistic equation: The model above was for discrete time steps described by a difference equation. For continuously breeding populations (birth and death continuous) we need a continuous form of the model using a differential equation. So for exponential population increase the rate of population increase is dn/dt and this speed of change is described in the absence of competition by: dn/dt = rn where r is the intrinsic rate of natural increase which is lnr or lnr o /T So the continuous equivalent to Fig is shown in Fig and this is the differential form of the difference equation N t = N o R t BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 15

16 16. Logistic limitation to a carrying capacity: The differential form of N t+1 = N t R/1+(aN t ) in Fig 5.18 is given by: dn/dt = rn((k - N)/K) This is the famous logistic equation. This shows that exponential increase is decreased to K by the logistic term (K - N)/K BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 16

17 17. Asymmetrical competition: Large vs small Impatiens in a woodland (Fig. 5.26): Small plants did not grow and so the asymmetry increased with time. Root vs shoot competition in morning glory Fig (Weiner expt.): Root competition for nutrients resulted in most biomass reduction, but shoot competition for light generated most size inequality and increase in asymmetry. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 17

18 Figure 5.1: Intraspecific competition among cave beetles for cave cricket eggs (a) scramble or exploitation, (b) contest or interference. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 18

19 Figure 5.2: Survivorship of red deer on the island of Rhum declines with lower birth rate and increased density. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 19

20 Figure 5.3: Density dependent mortality in flour beetles changes from (1) density independence, to (2) undercompensating density dependence, to (3) overcompensating density dependence. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 20

21 Figure 5.4: Exact compensation in trout fry. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 21

22 Figure 6.5 (3 rd ed.): Density dependent mortality in soybeans leading to overcompensation with time. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 22

23 Figure 5.7: Density dependent birth and mortality rates. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 23

24 Figure 5.8: Differences between births and deaths (a), generate recruitment (b), and population increase to a carrying capacity (c). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 24

25 Figure 5.11: Examples of S-shaped population increase for (a) Rhizopertha beetles on wheat, (b) wildebeest after a rinderpest outbreak, and (c) willows after myxomatosis killed rabbit herbivores. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 25

26 Figure 6.14 (3 rd ed.): Effects of density on growth rate and size in (a) Rana tigrina frogs and (b) reindeer. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 26

27 Figure 5.12: Effects of intraspecific competition on growth and final biomass of populations of the limpet Patella cochlear. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 27

28 Figure 6.16 (3 rd ed. see Fig. 5.14, 4 th ed.): Constant final yield of plants sown at a range of densities for (a) subterranean clover, (b, c) the dune annual Vulpia fasciculata. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 28

29 Figure 5.15: Intraspecific competition in rye grass regulates the number of modules (tillers). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 29

30 Figure 5.16: k-values to describe variable density dependent mortality in (a) a dune annual, (b) almond moth, (c) fruit fly, and (d) the moth Plodia interpunctella. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 30

31 Figure 2.3: (Hassell, 1976) BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 31

32 Figure 6.20, 3 rd ed., see Fig. 5.17, 4 th ed.): k-values to describe density dependent reductions in fecundity in (a) limpets, (b) cabbage root fly, (c) grass mirid, and (d) plantain. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 32

33 Figure 5.18: Difference equation model to describe population increase in species with discrete generations BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 33

34 Figure 5.21: Different intensities of intraspecific competition incorporated in equation BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 34

35 Figure 6.26 (3 rd ed.): Equation 5.18 fitted to data for different beetle species in the laboratory (a, b, c & e), and winter moths in the field (d). BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 35

36 Figure 5.22: Range of population fluctuations for (a) values of b and R and (b) population size against time, generated by equation BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 36

37 Figure 5.23: Exponential and sigmoidal models of population increase against time for continuous breeding - the logistic model of population growth. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 37

38 Figure 5.26: Asymmetric competition in the woodland plant Impatiens pallida in SE Pennsylvania. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 38

39 Figure 5.27: Root vs shoot competition in morning glory vines, Ipomoea tricolor. BIOS 6150: Ecology - Dr. S. Malcolm. Week 3: Intraspecific Competition Slide - 39

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