Who am I?

A/P: Li Daiqin Specialty: Behavioural Ecology; Spiders; Biodiversity of Arthropods Current Research: Animal communication; Sexual selection; UV signalling; Crypsis and mimicry; Spider sociality; Biomaterials - Spider silk and nanomaterials for structural colour production Ultimate Goal: how animals having small brain with few neurons solve the everyday problems that they face within their respective environments. Teaching: LSM2251 and LSM4253 Contact details:

Email: dbslidq@nus.edu.sg Tel: 6516 4372 (Office) Office: S2 01-03

The Straits Times Dec. 25, 2004

My Favourite -- Jumping spider

Predator-prey relationship

Female-male mating

Male-male fighting

Species Interactions (1) Competition

LSM2251 Ecology and The Environment
A/P Li Daiqin

Outline

References Background

Classification of species interactions

Competition Symbiosis Exploitative interactions Predation Parasitism Herbivory

References

Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5th ed. McGraw-Hill. Byers, J. E. 2000. Competition between two estuarine snails: implications for invasions of exotic species. Ecology 81: 1225-1239. Dayan, T. & Simberloff, D. 2005. Ecological and community-wide character displacement: the next generation. Ecology Letters 8: 875-894. Denno, R.F. & Roderick, G.K. 1992. Density-related dispersal in planthoppers: effects of interspecific crowding. Ecology 73: 1323-1334. Tilman, D. & Cowan, M. 1989. Growth of old field herbs on a nitrogen gradient. Functional Ecology 3: 425-438.

Background- Interactions

Ecology is about NUMBERS and CHANGES in Numbers. What can affect/change numbers? How can the numbers be changed? How can we study changes in numbers? Can we predict changes in numbers? How?

Interactions

Classification of interactions

Classification of interactions

Competition

Competition
Intraspecific

Competition

Modes

Competitive

Exclusion and Niches Mathematical and Laboratory Models

Lotka-Volterra

Competition

and Niches

Character Displacement Competition between invasive and native species

Applications

Modes of competition
Interference:

Direct aggressive interaction between individuals.

Intra-specific:

Competition with members of own species.

Inter-specific:

Competition between individuals of two species - reduces fitness of both.

Intra-specific competition among plants

Plant growth rates and weights have been found to increase in low density populations.
Tilman and Cowan (1989) grew the grass Sorghastrum nutans at two different densities: low = 7 per plot, and high = 100 per plot. Found that at low densities S. nutans grew to a large size but not so for those in the high density plots. Competition for resources (nitrogen) was more intense at high population densities (versus little competition at low densities). This leads to mortality among competing plants. Self-Thinning

Population density, soil nitrogen, and the size attained by the grass Sorghastrum nutans (data from Tilman & Cowan 1989)

Molles 2010.

Molles 2010.

Intra-specific competition among planthoppers

Very difficult to show intra-specific competition in herbivorous insects.

Denno and Roderick (1992) demonstrated it within populations of planthopper Prokelesisia marginata. They thought their results would apply to other Homoptera due to a combination of limited resources, rapid population growth and their habit of aggregating.

Population density and planthopper performance (Data from Denno & Roderick 1992)
Molles 2010.

Interference competition among terrestrial isopods

Edward Grosholz (1992) used a field experiment with two treatments: (1) food limitation; and (2) density effects, to demonstrated it in isopods Porcellio scaber.

Supplementing food had no effect on survival. Survival was lower at the higher population density.

Due to cannibalism at the higher density

Molles 2010.

Intra-specific competition: summary

Laboratory

and field studies reveal intra-specific competition. Intra-specific competition for limited resources plays a key role in slowing population growth at higher densities. Interference competition can occur in the absence of obvious resource limitation.

Niches and inter-specific competition

Niche: Environmental factors that influence growth, survival, and reproduction of a species. Hutchinson (1957) defined niche as: n-dimensional hypervolume n equates the number of environmental factors important to survival and reproduction of a species. Gause (1934) came up with the Principle of Competitive Exclusion
Two species with identical niches cannot coexist indefinitely
One will be a better competitor and thus have higher fitness and eventually exclude the other. Thus, coexisting species will have different niches.

Niches and inter-specific competition

Fundamental niche the ideal hypervolume; the physical conditions under which a species might live, in the absence of interactions with other species. Realized niche includes interactions such as competition that may restrict environments where a species may live. Example: feeding biology as niches of Darwin finches - feeding niches
Availability of suitable food affects the survival and reproduction

The Galpagos Archipelago and Isla Daphne Major home of Darwins finches

Petren et al. 2005. Mol. Ecol. 14: 2943-2957

Feeding niches of Galapagos ground finches

Grant (1986) found differences in beak size among ground finches translates directly into diet. Size of seeds eaten can be estimated by measuring beak depths. Individuals with deepest beaks fed on hardest seeds. After the 1977 drought, the remaining seeds were very hard. Thus, mortality was most heavy in birds with smaller beaks (because they couldnt crack the hard seeds). Molles 2010

Famous Darwins finches the medium ground finch Geospiza fortis

Beak size within species Geospiza fortis

Boag & Grant. 1984. Biol. J. Linn. Soc. 22: 243-287

Relationship between hardness of seeds eaten and beak depth

With the population, individuals with the deepest beaks fed on the hardest seeds, while individual with the shallowest beaks fed on the softest seeds.
Molles 2010.

Seed depletion by the medium ground finch, Geospiza fortis, and average seed hardness (Grand 1986)

As seeds are depleted, finches eat the smallest and softest seeds first, leaving the largest and toughest seeds. Following drought (selection) not only are seeds in short supply, the remaining seeds are also tougher to crack. Mortality fell most heavily on smaller finches with smaller beaks.

Molles 2010.

Selection (drought) for large individuals with large beak size

Boag & Grant. 1984. Biol. J. Linn. Soc. 22: 243-287

Feeding niches of ground finches: summary

Food is the major determinant of survival and reproduction among ground finches, thus beak size gives a good picture of Darwin finches niches. However, the niches of other organisms are determined by entirely different environmental factors.

Mathematical and Laboratory Models

Mathematical

and laboratory models provide a theoretical foundation for studying inter-specific competition in nature Metz (1972) summarized models:
Abstractions and simplifications, not facsimiles of nature. Man-made construct; partly empirical and partly deductive. Used to provide insights into natural phenomena.

Modelling inter-specific competition: Lotka-Volterra competition model

Logistic

model for population growth of a single species:

dN/dt = rmaxN ((K-N) / K)

Population

growth of each species:

dN1 / dt = rmax1N1 ((K1-N1) / K1) dN2 / dt = rmax2N2 ((K2-N2) / K2)

Population growth slows as N increases as the ratio of numbers to carry capacity, either N1/K1 or N2/K2.

Modelling inter-specific competition: Lotka-Volterra competition model

Effect

of inter-specific competition on population growth of each species:

dN1 / dt = rmax1N1 ((K1-N1-12N2) / K1) dN2 / dt = rmax2N2 ((K2-N2- 21N1) / K2)
12: Effect of individual of species 2 on rate of population growth of species 1. 21: Effect of individual of species 1 on rate of population growth of species 2. 12 and 21 are called competition coefficients.

12 > 1, then the competitive effect of an individual of species 2 on the population growth of species 1 is greater than that of an individual of species 1. 12 < 1, then the competitive effect of an individual of species 2 on the population growth of species 1 is less than that of an individual of species 1.

Modelling inter-specific competition: Lotka-Volterra competition model

In

general, LV predicts coexistence of two species when, for both species, interspecific competition is weaker than intraspecific competition. one species is predicted to eventually exclude the other.

Otherwise,

Modelling inter-specific competition: Lotka-Volterra competition model

Predict

population growth for the two species will stop when:

N1=K 1- 12 N2

and

N2 = K 2- 21 N1

Zero Growth Isoclines

Above: Population decreasing Below: Population increasing

Coexistence of two species is only possible when isoclines cross.

Modelling inter-specific competition: Lotka-Volterra competition model

Molles 2010.

Paramecia lab experiments

Gause (1934) used laboratory experiments to test the major predictions of LV competition model. He demonstrated resource limitation with freshwater, ciliated protozans Paramecium caudatum and Paramecium aurelia in presence of two different concentrations of the bacterium Bacillus pyocyaneus.

When grown alone, carrying capacity determined by intra-specific competition. When grown together, P. caudatum quickly declined. Reduced resource supplies increased competition.

Paramecia lab experiments

Would one of the two species drive the other to extinction if grown together in microcosms where they were forced to compete with each other for a limited food supply. When grow alone, both species did well, the carrying capacity is determined by intra-specific competition. When grown together, P. aurelia survived, while the population of P. caudatum quickly declined. Competition exclusion results from competition for limited food supply. Molles 2010.

Flour beetle experiments

Tribolium

products.

beetles infest stored grain

Park (1954) studied inter-specific competition between T. confusum and T. castaneum under varied environmental conditions.

When grown separately in hot-wet environments, both species did well. When grown together in hotwet environments, Tribolium castaneum usually excludes T. confusum. In contrast, when grown together in cool-dry conditions, T. confusum usually excludes T. castaneum. How can we interpret the results of these laboratory experiments in terms of the effects of competition on these species niches?

Competition and niches

Competition

can have short-term ecological effects on the niches of species by restricting them to their realized niches.

But if competitive interactions are strong and pervasive enough, they may produce an evolutionary response in the competitor population.

Changes fundamental niche (fuller range of environments in the absence of inter-specific competition).

Niches and competition among plants

Tansley (1917) suggested inter-specific competition restricts realized niche of each of two species of bedstraw (Galium spp.) to a narrower range of soil types.
One species is usually found on acidic soils, the other on limestone soils. When grown together, both showed clear competitive dominance on their own soil types. But both species can grow well on either soil type if they are not planted together.

Figure 13_17

Niche overlap and competition between barnacles

Connell (1961) discovered interspecific competition in barnacles. Balanus plays a role in determining lower limit of Chthamalus within intertidal zone. Chthamalus will grow in the middle intertidal zone if Balanus are removed. Balanus crowd out Chthamalus. Balanus are not found in the upper intertidal as it cannot withstand the high levels of desiccation.

Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5th ed. McGraw-Hill.

Character displacement
Because degree of competition is assumed to depend upon degree of niche overlap, inter-specific competition has been predicted to lead to directional selection for reduced niche overlap.

Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5th ed. McGraw-Hill.

Character displacement
Taper

and Case: Necessary criteria:

Morphological differences between sympatric species are statistically greater than differences between allopatric populations. Differences between sympatric and allopatric populations have genetic basis. Differences between sympatric and allopatric populations evolved in place, and are not derived from different founder groups already differing in the character.

Character displacement
Variation in the character (e.g. beak size) must have a known effect on use of resources (e.g., seed sizes). Must be demonstrated competition for the resource (e.g., food) and competition must be directly correlated with character similarity (e.g., overlap in beak size). Differences in character cannot be explained by differences in resources available to each of the populations.

Applications: competition between native and invasive species

Species invasion is one of the most significant contemporary environmental problems. What are the mechanisms by which allow invasive species to invade communities of native species? Superior competition ability. Byers (2000) used field experiments to explore the ecological relationships of native (Cerithidea californica) and an invasive species of mud snail (Batillaria attramentaria) in North America.

Density effects of native and invasive snails on the abundance of diatoms and growth rate resource competition

Both reduce diatom density, but no differences in the effect on diatom abundance

Invasive snails grow faster than native snails at all densities, showing potential competition between native and invasive species

Competition: summary
Three types of competition. Laboratory and field studies reveal intra-specific competition. Two species with identical niches cannot coexist indefinitely, thus coexisting species will have different niches. Mathematical and laboratory models provide a theoretical foundation for studying inter-specific competition Competition can have significant ecological and evolutionary effects on the niches of species.