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1 FROM Wednesday - end of lecture on comparative life histories: A laboratory evolution experiment - effects of different rates of adult mortality on life history traits in Drosophila (fruit flies) Question: In environments where there is high adult mortality (e.g. high predation) what happens to other aspects of Drosophila life history? Experimenters set up selection lines - in some removed adults 2 x weekly ( High adult mortality treatment ), in others didn t ( Low adult mortality treatment ) kept all other factors constant Kept selection up from Some of their predictions: When adult mortality is high, populations should evolve Higher fecundity early in life Shorter development times Smaller adult size Effect of high adult mortality on amount of early reproduction: Which treatment reproduces earlier? Do they do it from the start of the experiment? High mortality Low mortality Effect of high adult mortality on larval development time: Which treatment has a shorter larval development time? High mortality Low mortality Effect of high adult mortality on body size: Which treatments adults are smaller? High mortality Low mortality Months Months

2 Summary Life history evolution studies show some mechanisms by which animal and plant populations diversify in response to their environment. Some patterns found are general to many organisms. For example, the manipulative selection experiment in Drosophila shows a pattern seen in other descriptive experiments: High adult mortality leads to earlier reproduction, shorter development time, and smaller body size Population regulation and competition I. Population regulation Density-dependent and densityindependent factors II. Intraspecific competition III. Introduction to species interactions Types of interactions The ecological niche IV. Interspecific competition Competitive displacement, exclusion, and dominance A naturalist observed swifts in his village in England... "I am now confirmed in the opinion that we have every year the same number of pairs invariably; at least the result of my inquiry has been exactly the same for a long time past. The number that I constantly find are eight pairs, about half of which reside in the church, and the rest in some of the lowest and meanest thatched cottages. Now as these eight pairs - allowance being made for accidents - breed yearly eight pairs more, what becomes annually of this increase?" White 1778 In 1983, a couple of naturalists returned to the same village Fewer thatched cottages, no swifts in the church, but a total of twelve pairs overall. A net change of 4 pairs of swifts in 205 years... We are likely to focus on dramatic changes in population densities: increases such as locusts, gypsy moths, lemmings, or humans So, what regulates populations? i.e. keeps them within certain limits? decreases such as buffalo, condors, passenger pigeons But most populations are, like the swifts, remarkably constant

3 We can categorize the factors that influence populations as density-dependent factors What s a density-dependent factor? and density-independent factors What s a density-dependent factor? A factor that increases in intensity as density (population size per unit area) increases. Specifically, this means the proportion of the population affected increases with density. A (made-up) example. We want to know if flooding acts as a density-dependent mortality factor of a certain grass growing in a wash. Is flooding a density-dependent mortality factor acting on the grass? From 1989 to 1995 we go out each year after the monsoon, count the number of grass plants that are alive, and the no. dead from flooding. Then we plot our data. No. dead Density (population size)

4 Because we expect a larger number to die as the population size becomes large. If we change the y axis to proportion dead... If a factor is density-dependent it should affect the population in this way... Prop. dead We can see there is no increase in proportional mortality with an increase in density Prop. dead Density (population size) Density (population size) Why do we care about densitydependent factors? Factors which may act in a densitydependent manner? Only density-dependent factors can regulate populations. Other factors can have effects on population dynamics, but don t predictably rein in populations at high densities. Competition - intraspecific (within species) interspecific (between species) Factors which may act in a densitydependent manner? Disease caused by pathogens Contagious diseases are better transmitted at high density Factors which may act in a densitydependent manner? Predation and herbivory predators may be more likely to find abundant prey types predator reproduction greater when abundant food

5 Density-independent mortality factors can be important in a population s dynamics, even if they are not regulating. Example: winter mortality of herons Factors which are generally density -independent? Weather - rain, wind, drought, freezes Fire 5000 No. of nesting pairs = Severe winters Year Experimental demonstration of densitydependent parasitism: Goldenrod gall fly Goldenrod gall fly - an experiment Adult lays eggs in stems of goldenrod Larval fly induces the formation of a gall Fly overwinters in gall All the galls in several fields gathered at the end of a season, put in an unheated shed for the winter In the spring, galls put at one of two densities back into fields Most important mortality agent a parasitic wasp that also spends the winter in the gall High density fields Goldenrod gall fly - an experiment Low density fields Goldenrod gall fly - an experiment Experimenter measured the parasitism of the new season galls in the high and low density fields If parasitism density-dependent, what would pattern would you expect to see? Results: Greater proportion parasitized in high density fields than in low density fields

6 Intraspecific competition increases death rates. Example: Winter moth Proportion surviving II. Intraspecific competition Competition reduces the contribution of the competitors to the next generation through increased death rates or decreased birth rates Numbers of eggs in spring Not a winter moth Intraspecific competition reduces birth rates - example in longhorn cattle III. Species interactions - an introduction Species interactions can be classified according to the net effect on the species involved, positive or negative 1.0 Young per female Density of longhorns Mutualisms are +/+ interactions both partners benefit In commensalism one partner benefits, and the other is unaffected (+/0) Example: Pollinators and flowers more examples to come in next lecture... Example - cattle egrets feed on insects that cattle dislodge. Cattle are unaffected by the egrets.

7 In amensalism one partner is harmed, and the other is unaffected (-/0) In predation and parasitism one partner benefits, and the other is harmed (+/-) Examples - Large animals may negatively influence plants near water holes. Larger herbivores may incidentally consume smaller herbivores like insects, etc. Predation In competition, both partners are harmed (-/-) Parasitism Exploitative vs. interference competition Exploitative competition: competition through exploitation of a common resource. Winners of competition appropriate more of the resource. For example, gila woodpeckers and starlings compete for nest cavities Exploitative competition: Example Competition between trees for light * * C. Exploitative vs. interference competition Seedlings Slight disadvantage of some seedlings (e.g. late germination, slower growth rate) becomes exaggerated as lose access to light. Interference competition: Competitors directly interfere with each other. The victor is the one that wins the contest. Mature trees

8 Interference competition. Example: Young larvae of parasitic wasps are often equipped with large mandibles used to kill competitors in the same insect host. Interactions with other species may affect a species ecological niche. What s a niche? The ecological niche Many definitions in the literature A simple one: The niche is the sum of the ecological requirements of a particular species. E.g., habitat requirements, food requirements, climatic requirements, etc. Two kinds of niches The fundamental niche - that which a species could use in the absence of predation or competition Two kinds of niches The fundamental niche - that which a species could use in the absence of predation or competition. The realized niche - what the species uses when predators and competitors are present Two kinds of niches Which is likely to be bigger? The fundamental niche or the realized niche?

9 Niches in the presence of interspecific competition - a classic example from the intertidal region on the coast of Scotland Interspecific competition in barnacles Intertidal region area between high and low tides sometimes under water, sometimes dry vs. Barnacles (crustaceans, Arthropoda) start life as free-swimming larval forms. They then settle on rocks and grow in size by molting, but do not move. Researcher found two species: Results of manipulation Balanus - covered by water most of the time Effects on Balanus - the one covered by water most of the time Chthamalus - exposed most of the time Distribution didn t change when Chthamalus removed - couldn t colonize area ordinarily occupied by Chthamalus because too dry From different areas he removed all adults of one or other species, and continued to remove their settling larvae. Results of manipulation So what does this have to do with niches? Effects on Chthamalus - the one exposed most of the time When Balanus removed, Chthamalus colonized the entire region. Ordinarily Balanus would out-compete Chthamalus in the lower regions by growing over them.

10 So what does this have to do with niches? So what does this have to do with niches? Chthamalus Balanus The niche occupied by Balanus is unaffected by Chthamalus The niche occupied by Chthamalus is reduced in the presence of Balanus The barnacle study provides an example of competitive displacement: one species niche was contracted in the presence of the other. What do you think would have happened to Chthamalus (the exposed species) if both species had the same fundamental niche? What do you think would have happened to Chthamalus (the exposed species) if both species had the same fundamental niche? The principle of competitive exclusion: when two species occupy the same fundamental niche, they cannot coexist. Chthamalus would have become extinct wherever Balanus was.

11 Competitive exclusion: An example from the Sonoran desert Graduate student at the U of A looked at competition for seeds among seed feeders 1) Competition among rodents Put fences around areas with holes that excluded kangaroo rats, allowed smaller rodents to pass Competitive exclusion: An example from the Sonoran desert In plots where Kangaroo rat the kangaroo rats were excluded: Copyright (c) Grolier Interactive Inc. There were more species of small rodents, and they were more abundant So, kangaroo rats competitively exclude other rodents Competitive dominance. Another example of competition for seeds in the desert. Competitive dominance. Perhaps most commonly, competition leads simply to the greater abundance of one species than the other. 2) Are ants competing with rodents for seeds? Harvester ant with large seed 2) Are ants competing with rodents for seeds? Removed ants from some plots, rodents from others, kept some as control plots. Rodents Ants Control removed removed plots 2) Are ants competing with rodents for seeds? Yes. Both kinds of seed predators are more abundant when the other kind is removed. No. ant Colonies No. of Rodents

12 What species dominates in competition may depend on the environment. An example with flour beetles. Experimenter put 2 species of beetles together at 29 C - A and B No. beetles Days B A At 29 At 29 C, A wins Experimenter put the same 2 species of beetles together at 32 C - A and B No. beetles Days B At 32 C, B wins A Why the reversal in outcome? A grows faster than B at 29 C, and B grows faster than A at 32 C

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