Island Biogeography 2

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    Island Biogeography

    IslandsDiscrete boundaries, and easy to compare & contrast

    "Oceanic" vs. Continental & "Landbridge"Oceanic Islands:

    diversity lower but area-effect strongerextinction higher (no rescue-effect)immigration lower (distance to mainland)

    Examples:Sky IslandsWoodlot "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) dissertationR. McArthur and E. Wilson (1963, 1967)Paradigm: central idea, organizing concept

    Dynamic Equilibrium (not historic)Diversity Constantbut with Turnover (species list)

    Munroe, E. G. 1948. "The geographical distribution of butterflies in the West Indies." Ph. D. Diss ertation, Cornell University, Ithaca.

    Island Patterns - Species-Area Relationship

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    Power Model S = cAz

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

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

    Area and Abundance

    Oceanic islands are a special case of species-abundancerelationship (Preston, 1962)

    Most Species are rare and only a few species are dominantLognormal 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 relationshipWhat arethe routes of immigration?

    Exponential S = k1

    e-k2(I)

    Normal S = k1 e-k2(II)

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

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    Krakatau Islands (1883 eruption):

    documentation of species turnover

    RakataExtinction

    RakataImmigration

    SertungExtinction

    SertungImmigration

    1908-1920

    2 20 0 28

    1920-1933

    5 4 2 7

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

    Equilibrium Theory of Island Biogeography

    Immigration Rates = DistanceExtincton Rates = Area

    Equilibrium Theory of Island Biogeography

    Strengths and Weaknesses

    Slopes of Extincton and Immigration rate curves not knownSlopes vary from island to islandIslands may not be in equilibriumExtincton and Immigration vary among species(continued)

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    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, ratesAssumes 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)

    IslandArea

    km2

    Distance

    km

    Bird Spp.

    1917

    Bird Spp.

    1968Extinctions

    Human

    Introd.Immigrations

    Turnover

    %

    LosCoronados

    2.6 13 11 11 4 0 4 36

    San Nicholas 57 98 11 11 6 2 4 50

    San

    Clemente145 79 28 24 9 1 4 25

    SantaCatalina

    194 32 30 34 6 1 9 24

    Santa

    Barbara2.6 61 10 6 7 0 3 62

    San Miguel 36 42 11 15 4 0 8 46

    Santa Rosa 218 44 14 25 1 1 11 32

    Santa Cruz 249 31 36 37 6 1 5 17

    Anacapa 2.9 21 15 14 5 0 4 31Diamond, 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. PacificCoast 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. Wright, S. J. 1985. How isolation affects ratesof turnover of species on is lands. Oikos 44:331-340.

    Experimental Biogeography

    Defaunation of Florida Mangrove Islandsrapid increase, overshooting, stabilization"E1" is small isolated island

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

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    Continental Islands

    Rescue Effect: reduced turnover due to replacement

    Wright, S.J. 1980. Density compensation in island avifaunas. Oecologia 45: 385-389. Wright, S. J. 1985. How isolation affects ratesof turnover of species on is lands. Oikos 44:331-340.

    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 ofBiogeography, 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.

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    Sky Islands

    Nonequilibrium: no isolation effect, slow extinction, no dispersal

    S = cAz

    Smammal = 1.188 A0.326 Sbird = 2.526 A

    0.165

    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

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    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)

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    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 RatesTurnover and Equilibrium

    Krakatau Islands Biogeography

    Speciation vs. ImmigrationVolcanicvs.Mature Islands

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

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    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 ROc ky 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 biotarelaxation model: patterned extinction of original biota

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

    Nested Island DistributionsMalaysian 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 escapeextinction

    Insular Distribution Function: tradeoff of immigration & extinctonsmall islands OK if they're close to the mainland

    Lomolino, M.V. 1986. Mammalian comm unity structure on islands: Immigration, extinction and interactive effects. Biol. Journal LinneanSoc. 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.

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    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 ecologicaltolerance, rapid population growth

    K-Selected: mature ecosystems, stable population

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    Extinction of Birds in the Hawaiian Islands

    Selective extinction of ground-dwelling and carnivorous birds

    James, H.F. 1995. Prehistoric extinctions and ecological c hanges 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 ArchipelagoLand plant dispersal - "Filter" Dispersal Route

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

    Patterns Reflecting Interspecific Interactions

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    Checkerboard Distribution: only one or the other of twocompeting species occur on each island.Example: Bismark Archipelago flycatchers Pachycephalapectoralisand P. melanura dahli

    Patterns Reflecting Interspecific Interactions

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

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

    Patterns Reflecting Interspecific Interactions

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

    Moulton, M.P. and Pimm, S.L. 1986. The introduced Hawaiian avifauna: Biogeographical evidence for c ompetition. American Naturalist121: 669-690.

    Patterns Reflecting Interspecific Interactions

    Niche shift: atypical habitats occupied on islands where preditor

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    absent (shrews are predators on voles)

    Lomolino, M.V. 1984. Immigrant selection, predatory exclusion and the distributions of Microtus pennsylvanicusandBlarina brevicaudaon islands. American Naturalist 123: 468-483.

    Patterns Reflecting Interspecific Interactions

    Niche shift: Anolissize differentiation when more than specieson 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 s ize in solitary population of West IndianAnolis lizards.Oikos 29: 44-51. Foughgarden, J. Heckel, S. and Fuentes, E.R. 1983. Coevolutionary thoery and the biogeography and comm unitystructure of Anolis. p. 371-410 IN Huey, R.B. and Pianka, E.R. and Schoener, T.W. (eds.) Lizard Ec ology: Studies of a model organism. HarvardUniv. Press.

    Patterns Reflecting Interspecific Interactions

    Predator Release: species (example spiders) more aboundantwhere predator absent

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

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    Patterns Reflecting Interspecific Interactions

    Density Overcompensation: combined populations onseveral small islands greater than on a single large island ofequal area.Density Compensation: abundance constant despiteincreasing species (fewer per species)Density Stasis: abundance increases with increasing

    species biomass

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

    Applications of Island Biogeographic Theory toNature Preserves

    SLOSSSingle Large or Several Small

    Applications of Island Biogeographic Theory toNature Preserves

    Single Large vs. Several Small

    Circular vs. "Peninsular"

    Clumped vs. Spread out

    Connected vs. Unconnected

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

    Applications of Island Biogeographic Theory toNature Preserves

    Reserves should be as large as possibleArea - Diversity

    Unique habitats (biotas) should have multiple preserves(nearby)

    RedundancyIsolation

    Preserves should be round and continuousPeninsula effectIsolation

    Priority given to highest endemicity and vulnerabilityCorridores between preserves should be maintained

    Evolutionary Trends on Islands

    Flightlessness and reduced dispersal ability

    Birdsabsence of predators (no need to fly)

    limited resources (expensive to fly)InsectsExpansion into vacant niches

    (Orthopteran "wettas" of New Zealand)Precintiveness: dispersal away from hatching area isfatal

    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

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    Evolutionary Trends on Islands

    Body Size Change on Islands

    Dwarfismreduced resource needbetter shelter in treeless islands

    Gigantismtraits that allowed dispersal (small size a disadvantage)wider range of preygreater 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 resourcesRodents - gigantism

    immigrant selection: small animals disperse moreeasilyecological release: small size not needed to avoidpredators

    EVOLUTIONARY TRENDS

    Size variation

    squirrel size vs. island area

    Heaney, L.R. 1978. Island area and body size of insular mammals: Evidence from the t ri-colored squirrel (Callisciurus prevosti) of

    Southwest Africa. Evolution 32: 29-44.

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    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 extinctionInvasion of Most Islands1.Speciation and Range Restriction2.Few Relictual Populations3.Replaced by new Stage 1 Species4.

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

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