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Island Biogeography

Islands
Discrete boundaries, and easy to compare & contrast "Oceanic" vs. Continental & "Landbridge" Oceanic Islands: diversity lower but area-effect stronger extinction higher (no rescue-effect) immigration lower (distance to mainland) Examples: Sky Islands Woodlot "Islands" Prairie Potholes

Historical Background

Static Assumption: Island composition fixed


Pattern recognized by Forster & de Candolle Lack, Mayer 1940's; Hutchinson 1950's (islands, diversity)

Equilibrium Theory of Island Biogeography


Munroe, E.G. (1948) dissertation R. McArthur and E. Wilson (1963, 1967) Paradigm: central idea, organizing concept Dynamic Equilibrium (not historic) Diversity Constant but with Turnover (species list)
Munroe, E. G. 1948. "The geographical distribution of butterflies in the West Indies." Ph. D. Dissertation, Cornell University, Ithaca.

Island Patterns - Species-Area Relationship

Power Model S = cAz Semi-logarithmic S = d + k log(A)

On oceanic islands diversity less but area-effect stronger (curve steeper)

Area and Abundance


Oceanic islands are a special case of species-abundance relationship (Preston, 1962) Most Species are rare and only a few species are dominant Lognormal Distribution

Oceanic islands = smaller populations = fewer populous species


Preston 1962. The canonical distribution of commonness and rarity. Part 1 Ecology 43: 185-215. Part 2 Ecology 43: 410-432.

Species-Isolation Relationship
Less "general" than the Species-Area relationship What are the routes of immigration? Exponential S = k1 e-k2(I) Normal S = k1 e-k2(II)

k1 & k2 = constants; S = Species Richness; I = isolation (km)

Krakatau Islands (1883 eruption):


documentation of species turnover Rakata Rakata Sertung Sertung Extinction Immigration Extinction Immigration 1908-1920 2 20 0 28

1920-1933

Is the number of species decreasing, increasing, or stable?

Equilibrium Theory of Island Biogeography


Immigration Rates = Distance Extincton Rates = Area

Equilibrium Theory of Island Biogeography


Strengths and Weaknesses Slopes of Extincton and Immigration rate curves not known Slopes vary from island to island Islands may not be in equilibrium Extincton and Immigration vary among species (continued)

Equilibrium Theory of Island Biogeography


Strengths and Weaknesses (continued) Extincton and Immigration not independent (high immigration rates save species from extinction) "Rescue Effect" Multiple mainlands = multiple immigration routes, rates Assumes no speciations on island (strict sense) 2 islands - area correlated with habitat diversity

Isolation and Speciation


Low Immigration Rates Permit Speciation (vacant niches) Speciation increases diversity

Equilbrium Island Biogeography & Turnover


Turnover on "Landbridge" islands (California Channel Islands)
Island Los Coronados San Nicholas San Clemente Santa Catalina Santa Barbara San Miguel Santa Rosa Santa Cruz Anacapa Area km2 2.6 57 145 194 2.6 36 218 249 2.9 Distance km 13 98 79 32 61 42 44 31 21 Bird Spp. 1917 11 11 28 30 10 11 14 36 15 Bird Spp. 1968 11 11 24 34 6 15 25 37 14 Extinctions 4 6 9 6 7 4 1 6 5 Human Introd. 0 2 1 1 0 0 1 1 0 Immigrations 4 4 4 9 3 8 11 5 4 Turnover %

36 50 25 24 62 46 32 17 31

Diamond, J.M. 1969. Avifaunal equilibria and species turnover rates on the Channel Islands of California. Proc. Natl. Acad. Sci 64: 57-63. Jones, H.L. and Diamond, J.M. 1976. Short-time-base studies of turnover in breeding bird populations on the Channel Islands of California. Condore 73: 526-549.

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California Channel Islands

Porcasi, P., Porcasi, J.F., O'Neill C. Early Holocene coastlines of the California Bight: The Channel Islands first visited by humans. Pacific Coast Archaeological Society Quarterly 35 (2&3): 1-24. http://www.pcas.org/Vol35N23/3523Porcasi.pdf

Effect of Island Size on Turnover rates


Turnover rates lower in larger islands:

Wright, S.J. 1980. Density compensation in island avifaunas. Oecologia 45: 385-389. of turnover of species on islands. Oikos 44:331-340.

Wright, S. J. 1985. How isolation affects rates

Experimental Biogeography
Defaunation of Florida Mangrove Islands rapid increase, overshooting, stabilization "E1" is small isolated island

Simberlof & Wilson 1970. Experimental zoogeography of islands: a two-year record of colonization. Ecology 51: 934-937.

Continental Islands
Rescue Effect: reduced turnover due to replacement

Wright, S.J. 1980. Density compensation in island avifaunas. Oecologia 45: 385-389. of turnover of species on islands. Oikos 44:331-340.

Wright, S. J. 1985. How isolation affects rates

Target Area Effect:


greater immigration rate on larger islands

Buckley, R.C. and S.B. Knedlhans (1986). Beachcomber biogeography: interception of dispersing propagules by islands. Journal of Biogeography, 13: 69-70.

Small Island Effect:


no area-diversity effect on small islands too few habitats

Niering, W.A. 1963. Terrestrial ecology of Kapingamarangi Atoll, Caroline Islands. Ecological Monographs 33:131-160.

Sky Islands
Nonequilibrium: no isolation effect, slow extinction, no dispersal S = cAz

Smammal = 1.188 A0.326

Sbird = 2.526 A0.165

Brown, 1978. The theory of insular biogeography and the distribution of boreal birds and mammals. Great Basin Nat. Mem. 2: 209-277.

Great Basin Sky Islands - RELAXATION


Pleistocene Species Supersaturation

Brown, 1978. The theory of insular biogeography and the distribution of boreal birds and mammals. Great Basin Nat. Mem. 2: 209-277.

Sky Island Biogeography


Southwestern Sky Islands In Equilibrium: frequent dispersal

Lomolino, Brown, Davis, 1989. Island biogeography of montane forest mammals in the American Southwest. Ecology 70: 18-194.

Sky Island Biogeography


Southwestern Sky Islands In Equilibrium

Lomolino, Brown, Davis, 1989. Island biogeography of montane forest mammals in the American Southwest. Ecology 70: 18-194.

Sky Island Biogeography


Andean Birds: Holocene colonization

(Nores, 1995)

Freshwater Lakes Biogeography


Speciation vs. Immigration Effect of Antiquity ( !) and Speciation on Diversity

Barbour & Brown, 1974. Fish diversity in lakes American Nat. 108: 473-489

Krakatau Islands Biogeography


Immigration Rates Turnover and Equilibrium

Krakatau Islands Biogeography


Speciation vs. Immigration Volcanic vs. Mature Islands

Thornton 1996. Krakatau: The destruction and reassembly of an island ecosystem. MA, Harvard.

Krakatau Islands Biogeography


Differential Immigration Rates for Plants

Bush Whittaker 1991. Krakatau: Colonization patterns and hierarchies. Journal Biogeography 18: 341-356.

Extinction on Sky Islands


Small generalist herbivores have lower extinction rates. low energy requirements (smaller size, larger popn.) more available energy (lower trophic level)

Brown, J.H. 1971. Mammals on mountaintops: Non-equilibrium insular biogeography American Naturalist 105: 467-478. Brown, J.H. 1978. The theory of insular biogeography and the distribution of boreal birds and mammals. Great Basin Nat. Memoirs 2: 209-277. Patterson, B.D. 1984. Mammalian extinction and biogeography in the southern ROcky MOuntians. p. 247-294 in M.H. Nitecki (ed.) Extinctions Univ. Chicago.

Relaxation and Nested Island Biotas


immigrant pattern: dominated by "best migrators" relict pattern: random extinctions of original biota relaxation model: patterned extinction of original biota

Darlington, 1957. Zoogeography: The Geographical Distriburion of Animals. John Wiley & Sons, NY.

Nested Island Distributions


Malaysian Archipelago ants

Wilson, E.O. 1959. Adaptiave shift and dispersal in a tropical ant fauna. Evolution 13: 122-144.

Single-species Distributions
Metapopulation analysis: minimum area needed to escape extinction Insular Distribution Function: tradeoff of immigration & extincton small islands OK if they're close to the mainland

Lomolino, M.V. 1986. Mammalian community structure on islands: Immigration, extinction and interactive effects. Biol. Journal Linnean Soc. 28: 1-21. Lomolino, M.V. 1998. A species-based, hierarchical model of island biogeography. In E. Wiher and P.A. Keddy (eds.) The search for assembly rules in ecological communities. Cambridge Univ. Press.

Variable Immigration Abilities


Channel Island Distances (& Rates)

Wenner, A.M. and Johnson, D.L. 1980. Land vertebrates on the California Channel Islands: Sweepstakes or bridges? p. 497-530 In: D.M. Power (ed.) The California Islands: Proceedings of a multi-disciplinary symposium Santa Barbara Museum Nat. Hist.

Selective Immigration & Extinction


Biased insular community composition harmonic (balanced) disharmonic (proportions different from mainland)

Establishment of Immigrants
Population-Ecology Life Strategies r-Selected: disturbed ecosystems, broad ecological tolerance, rapid population growth K-Selected: mature ecosystems, stable population

Extinction of Birds in the Hawaiian Islands


Selective extinction of ground-dwelling and carnivorous birds

James, H.F. 1995. Prehistoric extinctions and ecological changes on oceanic islands. p. 87-102. in P.M. Vitousek, L.L. Loope, and H. Anderson. Islands: Biological Diversity and Ecosystem Function. Springer Verlag, NY .

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Nested Island Distributions


Aleutian Island Archipelago Land plant dispersal - "Filter" Dispersal Route

Williamson, M. 1981. Island Populations, Oxford Univ. Press.

Patterns Reflecting Interspecific Interactions

Checkerboard Distribution: only one or the other of two competing species occur on each island. Example: Bismark Archipelago flycatchers Pachycephala pectoralis and P. melanura dahli

Patterns Reflecting Interspecific Interactions


Incidence Function: proportion of islands inhabited vs. number of species per island (diffuse competition)

Diamond, J.M. 1975. Assembly of species communities. p. 342-444 In. M.L. Cody and J.M. Diamond (eds.) Ecology and evaluation of Communities. Belknap Press, Cambridge.

Patterns Reflecting Interspecific Interactions


Effect of close-competitors on an island being invaded success correlated with difference from resident species

Moulton, M.P. and Pimm, S.L. 1986. The introduced Hawaiian avifauna: Biogeographical evidence for competition. American Naturalist 121: 669-690.

Patterns Reflecting Interspecific Interactions


Niche shift: atypical habitats occupied on islands where preditor

absent (shrews are predators on voles)

Lomolino, M.V. 1984. Immigrant selection, predatory exclusion and the distributions of Microtus pennsylvanicus and Blarina brevicauda on islands. American Naturalist 123: 468-483.

Patterns Reflecting Interspecific Interactions


Niche shift: Anolis size differentiation when more than species on island

Roughgarden, J. 1974.Niche width: Biogeographic patterns among Anolis lizard populations. American Naturalist 108: 429-442. Roughgarden, J. and Fuentes, E.R. 1977. The environmental determinants of size in solitary population of West Indian Anolis lizards. Oikos 29: 44-51. Foughgarden, J. Heckel, S. and Fuentes, E.R. 1983. Coevolutionary thoery and the biogeography and community structure of Anolis. p. 371-410 IN Huey, R.B. and Pianka, E.R. and Schoener, T.W. (eds.) Lizard Ecology: Studies of a model organism. Harvard Univ. Press.

Patterns Reflecting Interspecific Interactions


Predator Release: species (example spiders) more aboundant where predator absent

Schoener, T.W. and Spiller, D.A. 1987. High population persistence in a system with high turnover. Nature 330: 474-477.

Patterns Reflecting Interspecific Interactions


Density Overcompensation: combined populations on several small islands greater than on a single large island of equal area. Density Compensation: abundance constant despite increasing species (fewer per species) Density Stasis: abundance increases with increasing species biomass

Wright, S.J. 1980. Density compensation on small islands. Oecologia 45: 385-389.

Applications of Island Biogeographic Theory to Nature Preserves


SLOSS Single Large or Several Small

Applications of Island Biogeographic Theory to Nature Preserves

Single Large vs.

Several Small

Circular vs.

"Peninsular"

Clumped vs.

Spread out

Connected vs.

Unconnected

Wilson, E.O. & E.O. Willis 1975. Applied biogeography. pp. 522-534 in M.L. COdy & M.M. Diamond (eds.) Ecology and Evolution of Communities. Cambridge, Belknap Press.

Applications of Island Biogeographic Theory to Nature Preserves


Reserves should be as large as possible Area - Diversity Unique habitats (biotas) should have multiple preserves (nearby) Redundancy Isolation Preserves should be round and continuous Peninsula effect Isolation Priority given to highest endemicity and vulnerability Corridores between preserves should be maintained

Evolutionary Trends on Islands


Flightlessness and reduced dispersal ability Birds absence of predators (no need to fly) limited resources (expensive to fly) Insects Expansion into vacant niches (Orthopteran "wettas" of New Zealand) Precintiveness: dispersal away from hatching area is fatal

Evolutionary Trends on Islands


Land Snails (smaller) Wind-dispersed - smaller is better Plants (bigger, simpler) loss of pappus, heavier, less resistant to saltwater (precintiveness) Become Trees

Evolutionary Trends on Islands


Body Size Change on Islands Dwarfism reduced resource need better shelter in treeless islands Gigantism traits that allowed dispersal (small size a disadvantage) wider range of prey greater energy and water reserves for famines

The Island Rule: "the big get smaller & the small get bigger" Optimal size 250 grams (red squirrel) Predators: canids, felids - dwarfism reduced resources Rodents - gigantism immigrant selection: small animals disperse more easily ecological release: small size not needed to avoid predators

EVOLUTIONARY TRENDS
Size variation squirrel size vs. island area

Heaney, L.R. 1978. Island area and body size of insular mammals: Evidence from the tri-colored squirrel (Callisciurus prevosti) of Southwest Africa. Evolution 32: 29-44.

EVOLUTIONARY TRENDS
Size variation mouse size vs. isolation

Ebenhard, T. 1988. Introduced birds and mammals and their ecological effects. Swedish Wildlife Research 13: 1-107.

EVOLUTIONARY TRENDS
Size variation lizard size vs. number of lizards per island

Soul, M.E. 1966. Trends in insular radiation of a lizard. American Midland Nat. 100: 47-64.

EVOLUTIONARY TRENDS
Taxon Cycles Island speciation ends in extinction 1. Invasion of Most Islands 2. Speciation and Range Restriction 3. Few Relictual Populations 4. Replaced by new Stage 1 Species
Wilson, E.O. 1959. Adaptive shift and dispersal in a tropical island fauna. Evolution 13: 122-144.