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But and species do not exist in a vacuum… Species interact…

Community A) Five fundamental types of species interactions:

Effect on species A B A B

Predation A B

Mutualism A B

Commensalism A B

Amensalism A B B) Concept of the Niche

1) Best known definition of niche is Hutchinson (e.g., 1957) a) role plays in environment b) role can be determined by measuring all of an organism’s activities and requirements

2) Examples 2-factors 3-factors high

Substratum friability low low high Wave exposure

3) By extension… niche defined as an N-dimensional hyperspace (encompasses all effects and requirements of a species) B) Concept of the Niche

3) Two types of niche a) fundamental: niche space determined by environmental factors and requirements. Manifest in the absence of other . b) realized: niche space determined by combined abiotic and biotic factors. Realized in presence of other organisms

fundamental realized fundamental niche always bigger (or at least as large) - biological interactions can (usually do) limit realized niche C) Competition

Defined: The common use of a resource that is in limited supply.

1) Within and between species a) Intraspecific - among individuals of the same species source of discussed previously b) Interspecific - among individuals of two or more species

2) Two types of competition

a) Interference b) Exploitative C) Competition

2) Two types of competition

a) Interference - direct competition A B i) e.g., aggression ii) e.g., territoriality (fishes, birds, limpets)

b) Exploitation - indirect competition i) Compete through a resource (R) ii) e.g., sessile spp. -- space, filter feeders -- plankton

A B barnacles mussels

R space C) Competition

3) Competitive exclusion principle

The more similar organisms are, the more likely they are to compete.

a) Species occupying the same niche cannot coexist.

b) The greater the niche overlap, the greater the likelihood of competitive exclusion, leading to local extinction of one species.

c) Leads to “resource partitioning” C) Competition

4) Resource partitioning

AB CD E number of individuals

resource gradient* species “packing” ABCD E * e.g., - seed / plankton size -elevation - height on tree / alga resource gradient C) Competition 5) Manifested in patterns a) non-overlapping spatial (or temporal) distribution

AB tidal number height of individuals

resource gradient reef depth

- Implication for relative competitive superiority? - Under what conditions would these patterns be most evident? C) Competition

5) Manifested in patterns a) negative (inverse) relationship in i) gradient in density

A B B A A A B B Abundance B B B sp. A B A A A A

ii) patchy / clumped Abundance sp. B A A B A A A B B A A A A B B B A A A C) Competition

6) Competitive release a) Change in distribution (or some other response such as growth) when separate and together sympatry (together) allopatry – separated in space

absence of mussels tidal height

absence of barns

Could examine observationally or experimentally, which preferred? C) Competition

7) Competitive symmetry a) Relative competitive strength b) superior, inferior (or) dominant, subordinate

A B Symmetrical A = B

A > B A B Asymmetrical A < B A B

How would you assess this?? C) Competition 8) Effects on measured variables a) Individual responses: • Behavioral (feeding rates, distribution) • Physiological (growth rate, reproductive rate) • Morphological (body size, ) Above responses referred to as “trait-mediated” On evolutionary time-scale, manifest as “character displacement”

b) responses: • Abundance (density) • Distribution (zonation) • Demographic rates (population growth) C) Competition 9) Character displacement

When differences among similar species whose distributions overlap geographically are accentuated in regions where the species co-occur, but are minimized or lost where the species' distributions do not overlap.

Reflects the consequences of competition in sympatry, where species co-occur to avoid competitive exclusion.

Example: Darwin’s finch friend’s beaks D) Consumption of one organism (prey) by another (predator), which by definition, occurs between organisms on different trophic levels (vs. competition: within same ) [but murky… cannibalism, “” as forms of competition*] 1) diagrammatically:

Predator A A

Herbivore B B C

Primary producer (plant / alga) C D E F

*(Polis et al 1989 Ann Rev Ecol Syst, Arim & Marquet 2004 Ecology Letters) D) Predation

2) Effects on prey (direct and indirect):

“Direct effects”: direct losses (removal of individuals) - death of individuals - mortality rate of population

“Indirect effects”: influence of predator on variable other than death or mortality • behavioral (feeding rates, foraging distribution) • physiological (growth rate, reproductive rate) • morphological (body size, biomass)

More “trait-mediated responses” vs. other “indirect effects” D) Predation

3) Effects on prey (individual and population): Individual responses: • behavioral (feeding rates, foraging distribution) • physiological (growth rate, reproductive rate) • morphological (body size, biomass) • oh yeah… and you can get completely or partially eaten

Population responses: • abundance, density • distribution ( use) • structure (e.g., size, age, sex ratio, genetic, spatial) • dynamics and persistence (regulation) D) Predation

4) Complex interactions (with other processes)

E.g., competition mediated by predation:

e.g., predator that specializes on barnacles and is restricted to the mid and lower intertidal

With barnacle predators Without barnacle predators

P P P tidal P P height

In absence of predator, barnacle out-competes mussels and expands distribution down into the mid intertidal D) Predation 4) More complex predation interactions:

Effect on species Apparent competition A B C C Where, A and B are prey and C is a common predator. Presence of both prey increases overall predation rates, leading to negative indirect A B effect on one another.

Trophic cascade C Where, A is primary producer, B is an , and C is a predator. B Effect of species on adjacent trophic level has net positive indirect effect on next trophic level. A Trophic cascades

Strong “top-down” effects that produce downward rippling effects through a food chain.

Higher tropic level predators indirectly affect plant biomass via their impacts on herbivore populations.

Strong “bottom-up” effects that produce upward rippling effects through a food chain.

Lower tropic level producers indirectly affect predator biomass via their impacts on herbivore populations. A linear “food chain” Trophic level Relative abundance Predator

Herbivore

Plants

Abiotic (e.g., , , light) Oksanen/Fretwell Model: and Food Chain Length

Predator

Herbivore

Plants

Abiotic resources (e.g., nutrients, water, light) increasing productivity  Oksanen/Fretwell Model A linear food chain

Predator

Carnivores Herbivore Plants Biomass

Plants

Environmental Productivity Oksanen/Fretwell Model: Productivity and Food Chain Length

•Depending on productivity of , food chains can have fewer or more than three trophic levels.

•As primary productivity increases, trophic levels will be sequentially added.

•Food chains that have an odd number of trophic levels should be filled with lush vegetation, because herbivores are kept in check by predators.

•Food chains that have an even number of trophic levels should have low plant abundance because plants are herbivore limited. Estes, J. A. et al. Science 1998. Killer Whale Predation on Sea Otters Linking Oceanic and Nearshore E) /

1) Occurs within or between trophic levels, more often between trophic levels a) mutualisms: e.g., pollinators obligate - required for each others existence - pollinators facultative – not required - cleaner fish and parasitized hosts b) : e.g., facilitation

mutualism Abundance A = B A B sp. A (symmetrical)

Abundance commensalism A < B A B sp. B (asymmetrical) How would you assess this?? F) Community metrics (w/ focus on diversity)

1) : number of species in a community

2) Species composition: identity of species that constitute a community 3) : species richness and relative abundance

Shannon-Weiner index of diversity:

H' = -Σ pi (ln pi)

Where pi is the proportion of individuals in the community that are species i F) Community metrics

4) Illustration of diversity

100 100 100 H'= 0.87 H'= 1.39 H'= 1.10 75 75 75 No. of 50 50 50 indiv.s 25 25 25 0 0 0 ABCD ABCD ABCD

Evenness: measure of the relative similarity of species abundance in a community

E= H'/(ln S) where, S is species richness G) Spatial scales of species diversity

1) Alpha (α): within habitat diversity

2) Beta (β): between habitat diversity

3) Gamma (γ): the total species diversity in a landscape

Gamma diversity is the product of alpha and beta diversity: γ = α * β H) Components of diversity

Biodiversity is, broadly speaking, the variety of life. It exists at all hierarchical levels, including genes, populations, species, functional groups, or even or ecosystems. Functional group is a collection of species with similar in a community. Can be of widely different taxonomic groups. Examples: primary producers, , herbivores, Multiple components of diversity within a community: i. Diversity of species within trophic levels ii. Diversity (number) of functional groups iii. Diversity of species within functional groups Stachowitz et al 2007 Ann Rev Ecol Syst. I) Mechanisms of diversity – community stability

Ways by which a more diverse community can enhance its stability (i.e. persistence in the face of perturbations)

Generally, multiple weak interactors create greater community stability than few strong interactors.

Consider trophic cascades versus complex food webs and the consequence of losing a single species

A A Impact of losing

species “B” ? B versus B C

C D E F I) Mechanisms of diversity – community stability

Complementarity refers to greater performance of a species in mixture than expected from its performance in monoculture. Examples: (i) facilitation, (ii) differential resource use among multiple species enhances community productivity and stability (plants and nutrients, predators on prey control)

Functional redundancy is when two or more species fulfill similar ecological functions in a community (e.g., trophic guilds such as planktivores, detritivores) that differ in their vulnerability to perturbations. Redundancy can contribute to community stability by compensating for relative vulnerability and loss of species. I) Mechanisms of diversity – community stability

Identity and Composition effects recognize the important effects of particular species within a community and the variation among species or particular combinations of species in their influence on an process or property.

Examples: presence of a , keystone predator, etc.

Sampling effects reflects the likelihood of including one of these important species as diversity or richness increases. J) Metacommunities

Metapopulations create metacommunities

A set of interacting communities linked by the dispersal of multiple, potentially interacting species.

Variation in rates of movement of species between communities influence species composition, diversity and community functions (e.g., planktivory, herbivory, detritivory, predation) and ecosystem functions (e.g., productivity, cycling)