l20-Metapopulation Dynamics and Island Biogeography
Introduction and Ecological Observations
Class Observations: Raccoons and Seed Dispersal
The lecture begins with observations from an animal sanctuary webcam featuring raccoons, colloquially known as "trash pandas."
Raccoons are observed eating shelled walnuts. In a natural setting, if a raccoon smashes a hard-shelled nut—causing seeds to shoot off into the forest—this represents a form of seed dispersal known as dyszoochory.
Other animals identified in the footage include an opossum, a tree squirrel, and an armadillo from a rural property.
Metapopulations Context
The lecture covers metapopulations, drawing from textbook Chapters 8 and 19.
Ecology is characterized as a non-linear topic involving feedback loops between population dynamics and life history traits.
Island Biogeography
Definition of Biogeography
Biogeography is the study of relationships between geographic area size, the degree of isolation of that area, and the species diversity (richness) found within it.
Definition of an "Island" in Ecological Terms
An island is any portion of isolated habitat surrounded by a different, potentially hostile environment.
Examples:
Land Islands: Landmasses surrounded by water (e.g., the islands of New Zealand).
Mountaintops: Species evolved for the alpine zone (cool temperatures, thin air) cannot survive at lower elevations. The valleys between peaks act as hostile barriers.
Lakes: Aquatic habitats surrounded by terrestrial environments. For aquatic species, the land is a hostile barrier.
The Species-Area Relationship
Ecologists observe that species richness increases with larger geographic areas.
Continents like Australia exhibit higher species diversity than smaller island systems like New Zealand.
Mechanisms for High Diversity in Large Areas:
Resource Base: Larger areas provide more resources.
Population Stability: They support larger populations, making them less vulnerable to extinction by sheer numbers.
Niche Diversity: There is a greater diversity of ecological niches.
Resilience to Stochastic Events: Environmental perturbations (storms, earthquakes) are less likely to affect the entire landmass. A storm hitting the Chatham Islands will affect a much higher proportion of the land than the same storm hitting Australia.
Trophic Rank and Area
The relationship between richness and area increases with higher trophic rank.
The slope of impact is steeper for predators compared to herbivores, and steeper for herbivores compared to plants.
As area decreases, predators are lost faster than lower trophic levels due to bottom-up food chain control (large islands have more biomass to support multiple levels).
The Simberloff and Wilson Experiment (1969)
Experimental Design
Conducted by E.O. Wilson and Daniel Simberloff in the Florida Keys (Florida Bay).
The study site consisted of mangrove islets (tiny islands) in shallow tidal areas.
Methodology:
The researchers selected islands of varying sizes and distances from the Florida mainland.
They used scaffolding and plastic sheets to enclose entire islets (like fumigating a house).
They sprayed pesticides to kill all arthropod fauna (spiders and other invertebrates).
Results and Conclusions
Species counts increased until they reached an asymptote roughly equal to the original species count prior to fumigation.
While the number of species remained consistent, the species composition (the specific types of creatures) changed.
The experiment proved that species richness is a result of island size and isolation, not simply the age of the island (time allowed for accumulation).
Equilibrium Theory of Island Biogeography
Dynamic Balance
Species richness is a balance between immigration/colonization (arrival of new species) and extinction.
Rate Factors
Immigration Rate: Higher for islands close to the mainland.
Extinction Rate: Higher for small islands due to fewer resources and higher vulnerability to shocks.
The Four Equilibrium States
Small/Far Islands: Lowest species richness; slow colonization, high extinction.
Large/Near Islands: Highest species richness; fast colonization, low extinction.
Small/Near vs. Large/Far: Intermediate richness levels.
The "Target Effect" (Class Experiment)
Small islands are small targets, making them physically harder for wind-dispersed seeds or animals to hit.
Increased distance reduces the likelihood of dispersers surviving the journey across the matrix.
Metapopulation Theory and Landscape Dynamics
Definition of Metapopulation
A subdivided population consisting of a series of discrete subpopulations behaving independently but connected by occasional dispersal (immigration and emigration).
Scale of Dynamics
Patch Scale (Local): Governed by birth () and death () rates. Individuals interact routinely within the patch.
Landscape Scale: Governed by dispersal and colonization. The survival of the metapopulation depends on the recolonization of patches where local populations have gone extinct.
Patch Characteristics
The landscape is viewed as a matrix (non-habitat) containing potentially suitable habitat patches.
Patches vary in size, quality, and isolation.
Necessary Conditions for a Metapopulation
Suitable habitat must be patchy.
All patches must be at risk of extinction.
Recolonization must be possible.
Patch dynamics must be asynchronous (they do not all respond to environmental cues at the same time).
Connectivity and Dispersal
Movement Types
Dispersal can be unidirectional (e.g., wind-blown from Australia to NZ) or bidirectional within a local metapopulation.
Successful colonization requires the ability to move, the tendency to disperse, and survival across the hostile matrix (e.g., snails crossing mowed fields with bird predators).
Types of Connectivity
Structural Connectivity: Based on the physical arrangement of patches (e.g., a riparian corridor or vegetation along a river).
Functional Connectivity: Based on how well the landscape facilitates movement, accounting for the organism's behavioral response to physical structures.
Note: Structural connectivity does not always equal functional connectivity. Human-engineered wildlife overpasses must be placed with biological knowledge to be effective.
Mathematical Framework for Metapopulations
Occupancy Rate
To understand the fraction of patches occupied at any given time (), we use:
Where is the probability of extinction and (or ) is the probability of colonization.
Persistence Requirements
For a metapopulation to persist, the fraction of occupied patches () must be greater than zero (p > 0).
This requires that the probability of extinction () be less than the probability of colonization ().
Case Studies in Metapopulation Dynamics
Sweden Cricket Population
Study of crickets in various-sized patches ranging up to hectares.
Findings: Extinction was common in patches below \text{ hectare}. Larger patches ( to hectares) had only extant (present) populations.
The maximum recorded distance for a cricket to successfully recolonize a patch was .
New York City Camera Traps
A network of camera traps from Downtown Brooklyn to Nassau County studied coyotes and deer.
Found that patch occupancy was related to patch size and the hostility of the urban matrix (concrete vs. suburban greenery).
Source-Sink Dynamics
Source: A patch where population growth is positive; individuals emigrate to other patches.
Sink: A patch with negative population growth. The population only persists due to constant immigration from source patches.
Cat Predation in Dunedin (Otepotee)
House cats were monitored via GPS collars.
For species like fantails () and bellbirds () in urban environments, the catch rate by cats was higher than the estimated total population.
These urban areas are sinks; the birds only remain present because of immigration from nearby native forest sources.
UC Campus Nest Success
In , native species (fantails and silver eyes) had lower nest success on campus compared to literature standards.
Increased predator trapping by the EnviroSoc group has since led to a rise in native bird nest success.
Freshwater Systems: Galaxids and Trout
Introduced trout act as a sink for juvenile galaxids by preying on them.
Galaxid populations are more diverse and include smaller individuals when trout are absent.
Barriers like waterfalls can protect galaxid populations from trout, but this effectively creates habitat fragmentation, preventing the galaxids from moving and functioning as a metapopulation.
Conservation and Reintroduction
Tui on Banks Peninsula
Tui were lost from Banks Peninsula around .
Because Tui are reluctant to fly across large areas of open land (low functional connectivity), human intervention was required.
They were reintroduced in and and are now thriving in areas like Akaroa.
Current research focuses on restoring linear features (native hedgerows) to reconnect the Southern Alps to Banks Peninsula.