Toward an Ecological Definition of an Island: A Northwest European Perspective Author(s): Yrjo Haila Reviewed work(s): Source: Journal of Biogeography,

Vol. 17, No. 6 (Nov., 1990), pp. 561-568 Published by: Wiley-Blackwell Stable URL: http://www.jstor.org/stable/2845140 . Accessed: 20/10/2012 00:44
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Journal of Biogeography (1990) 17, 561-568

Toward an ecological definition of an island: a northwestEuropean perspective

YRJO HAILA

Finland P. Rautatiekatu Department ofZoology,University ofHelsinki, 13, SF-OO100Helsinki,

Abstract.The equilibrium theory of islandbiogeography of MacArthurand Wilson aimed at providing a unifying framework for studyinginsular ecology. However, the theory does not elaboratethe conceptof an 'island' in an I suggest that different ecological framework. types of islands be ecologically definedon the basis of processes relativeto which some ecologically interesting entitieson those 'islands' actually are isolated fromtheirsurroundings. Four main island scales can be discerned,namely, thescales of (1) individuals, (2) populationdynamics,(3) population differentiation, and (4) evolutionary divergence. I illustrate these different island typesand theirinterfaces

Europe. The with examples mainly from northwestem on the equilibriumhypothesisseems relevantprimarily population dynamics scale. A research programme would include two phases: followingsuch differentiation first, assessmentof factorswhich make the environment of specific studied 'insular', and second, construction models for analysing ecological processes in those insular conditions. particular Key words. Islands, island biogeography, isolation, ecological scaling, conservation immigration, extinction, biology,Fennoscandia.

1. INTRODUCTION
Archipelagoes,be they composed of islands at sea or habitat patches on land, present both theoreticaland empiricalchallengesto ecological research.Theoretically the challenge is to understand processes structuring populations and communities in environments that comprisediscretespatial units.In practicesuch an underin enstanding oughtto helppreserving ecological diversity vironments fragmented by humanactivities. The theoryof island biogeographyby MacArthur& Wilson (1967) is the usual starting pointforisland studies. This is no wonder:The theory viewed island communities as dynamic units and offered a unifyingconceptual structure for island ecology, based on such apparently straightforward concepts as immigration and extinction ratesof organisms, assumed to be functions of island area and isolation.After itspresentation thetheory was hailed as a major breakthrough in ecological biogeography(e.g. has become a firm Simberloff, 1974). The theory partof the collective consciousness of ecologists, and has also a centralrole in conservation biology (Soule, 1986). It is difficult to overlookthe theory in insularstudies-islands exist in the ecological science largelyin the lightof interand reinterpretations pretations of the MacArthur-Wilson theory. The legacy of the theory has problematicfeatures, however, as has been pointed out by numerouscritics

(Simberloff,1976; Gilbert, 1980). The theory can be as interpreted in two ways (Haila & Jarvinen, 1982), either a strictexplanatorymodel that predictsthe numberof species in a given taxon on islands, or as a deductive factors schemethatpointsout variouspotentially important view insularcommunities. affecting Accordingto thelatter the theoryis a researchprogramme ratherthan a strict of the explanatoryhypothesis,and the appropriateness theory is assessed indirectly by evaluatingthe fruitfulness of thespecific research problems proposedby thetheory. The narrowinterpretation has dominated the scene. This is explainableby a semioticdimension of scientific theories to the role (Haila, 1986). By a semioticdimensionI refer communication. symbols have in scientific Symbols not only referto externalobjects but also carrywith them a of anothertype,namely,thatof being signs significance ('signifiers'; Barthes, 1967) of more abstractvalues. A value of the MacArthur-Wilson model is thatit particular the relationships formulates between the basic variables, rule. and islandarea,as a simpleanalytic species number have rangedfromhostplants Applicationsof the theory as islands for herbivorous insects (Janzen,1968), or the Galapagos as islands forDarwin's finches(Power, 1975), to pieces of laboratory glass as islands foraquatic diatoms (Patrick, 1975). These are all 'islands'-but of vastly different kinds. This is not a matterof semantics,but a matter of biology:These different environments are insular withrespectto quite different ecological processes.On the 561

562 Yrjo Haila areendemic in the Galapagos thefocusis on speciesthat outsideone or a few archipelago, and exist nowhere thefocus bydiatoms islands. On theglassslidescolonized of sessilereproducing individuals is on local assemblages inthesurroundthat arenotisolated atall from populations in-between. In other is on something ings. casesthefocus by similar can be described Formally the situations and 'extinction'), but such an variables('immigration' invalid. equation may be substantially of the islandecology needsa clarification It seemsthat an ecologically that very concept of 'an island'.I suggest fromthe be soughtby starting meaningful definition is 'an island'actually relative towhich ecological processes an island.
data on island distributions of othergroupsof quantitative and organismsare scarce. The scales forma continuum, theirinterfaces are also interesting and provide important problems forinsular ecology.

2.1. Individualscale On the individualscale a patch of land is an 'island' if some crucialphase of thelifecycle of individual organisms takesplace within its boundaries. Eitherreproobligatorily ductionor survivalmay be important. For instance,most breeding pairs of passerines on the Skokholm island (Williamson,1983), on islands in Lake Malaren (Ahlen & Nilsson, 1982), on the Aland Islands (Haila, Jarvinen & Kuusela, 1983), or on islands in Lake Inari (Haila, 1983b) completetheirreproductive phase on a single island, but somewhere else. The factthattheybreed spend the winter 2. ISLANDS INSPACEANDTIME on real islandsmay be important fortheir but reproduction, their life cycle is influencedby a varietyof additional relatedin ecology,and Space and time are mutually havedifferent characteristic factors. different ecological processes In contrast,large nonpasserines,e.g. predators,may byWiens,1981;Decourt, space-time scales(as discussed include several islands in their foraging territories in Delcourt& Webb, 1983; Birks,1986; O'Neill et al., 1986). northern archipelagoes(Ahlen & Nilsson, 1982; Haila et the space and timescales in In any particular situation together: al., 1983; Haila, 1983b), and no singleisland is an 'island' take placearecoupled which ecological processes forthem. thetimescale.This thespatial scale,thelarger thelarger in thenorth of 'islands', Birdsthatare sedentary a different thedifferentia specifica represent means that 'isolation', and the case. For themwinter survivalis a majorproblem, and they is to be viewedin relation to specific processes ofthose may demand larger areas of high-qualityhabitats for matrix processes. characteristic space-time than for breeding.For such species In Fig. 1 I reproduce thebasicscheme ofWiens(1981), successfulwintering relevant alternative forestfragments may be 'islands' primarily butadding a suggestion as to themost during the in an ecologicalwinter(Helle, 1984; Virkkala,1987). We are still on the 'an island'canbe viewed scalesinwhich In thefollowing I illustrate individualscale, but the processes defining'islands' are the evolutionary perspective. different from thoserelevant forbreeding northern schemewithexamplesmainlyfrom pairs. European If an island is too small for successfulreproduction or birds dominatemy examples because archipelagoes; survivalof individuals of a particular species,thespecies is TIME likelyto be absent.This makes the conceptof 'minimum on the level of individualrearea requirements' important units(Haila, Hanski & Raivio, 1989). Insularity productive Evolutionary and on the individual level may also lead to ecologically interesting consequences in terms of behavioural biogeographic forpreyor forhostsfornest such as searching adaptations dynamics parasitism (Wilcove, 1985). Populations: However,it can hardly be expectedthatinsularity of the environment on the has individual scale direct population demography, or community level consequences,exceptthrough being of dynamics inferior environment (or superior)quality as reproductive forindividual units.Individualslivingon such reproductive islandsare an integral partof theregionalpopulationin the Individual areas. surrounding behaviour, This way of defining 'islands' is plausible forefficiently conditions.A corresponding dispersingbirds in northern SPACE model of island colonizationby land birdscan be derived from an analogue that bird communitieson small nonisolated islands resemblerandomsamples 'drawn' by the island fromthe surrounding avifaunal universe(Preston, 1960; Haila, 1983a). The spatial dynamics of birds in small non-isolated in southern forest breeding fragments (1) (2) (3) (4) Finlandagree withthisview (Y. Haila, I. K. Hanski and S. FIG. 1. A schlcinic time-space scales of ecological of diflerent Raivio, in preparation). define scalesofinsularity. (1) andcriteria that corresponding processes A samplingview seems less plausible for most other Individualscale; (2) Populationscale 1: dynamics; (3) Population groups of organismsfor two main reasons. First,many scale. scale 2: differentiation; (4) Evolutionary

ofan island 563 Definition

actively dispersingorganisms,e.g. flyinginsects, move widelyaroundduring thereproductive to periodin contrast territorial birds.Consequently, in any single spatial unita 'rescue effect' (Brown & Kodric-Brown, 1977) may follow. Individuals may invade (and disappear) so frequently that no 'islands' can be discerned on the individual scale. Second, local population dynamics is crucially importantfor species that do not disperse which lead, e.g., to aggregated dispersion efficiently, patterns even within continuous, uniform habitats of (Southwood,1978). The spatialdynamicsof individuals such species may resemble 'sampling' on appropriate spatialand temporal scales, buttheecological relevanceof thisphenomenon assessment. requires separate Quantitativedata on the colonization dynamics of arthropodson islands are extremely scanty, but the thoroughly documented cases of small mangrove (Simberloff, 1969), and salt marsh(Rey, 1985) islands off the Floridan coast as well as sand islets off the Puerto Rican coast (Heathwole & Levins, 1973) overwhelmingly demonstrate the importance of otherthan individual-level processes.Distributional data on carabidbeetles (Niemela, Ranta & Haila, 1985; Niemela,Haila & Halme, 1988), ants (Vepsalainen & Pisarski, 1982) and land snails (Baur & Bengtsson,1987) in the archipelagoesof the Baltic Sea similarly demonstrate thatan individual-level interpretation would be inadequate. We enter anotherscale on which are involved. populations 2.2. Populationscale 1: dynamics Whenislandsbecome larger and moreisolated,theymay of supportpopulationsthat are dynamicallyindependent on other islandsand on themainland. populations Small islands in the Baltic are not islands forbirds on this scale. There is no evidence thatbird populationson singleislandsin theAland archipelago, up to some tensof km2in area, are dynamicallyindependent (Haila et al., 1983). Indeed,thisseems to hold truefora majority of land birdsbreeding on themainislandsof theAland archipelago as well, despite their substantialarea (970 kM2) and isolation(c. 40 km fromthe Swedish and 70 km fromthe Finnishmainland). Both species turnover and population changes documentedin the archipelago since the 1920s parallel trendsobserved on the Finnishmainland (Haila, Jarvinen & Vaisanen,1979). However,fora few sedentary birdspecies the large and most isolated Baltic islands such as Gotland and Aland clearlyare islandson thepopulation level. For instance, the green woodpecker (Picus viridis L.), European nuthatch (Sitta europaea L.) and marshtit (Parus palustrisL.) are abundanton the Swedish mainlandbut absent fromboth Gotland and Aland (Svardson, 1949), although suitable habitats abound on the islands. Furthermore, the greyheaded woodpecker(Picus canus Gmelin),and crestedtit (Parus cristatus L.), have colonizedtheAland Islands since the 1920s (as well as capercaillie(Tetrao urogallusL.) and hazel hen (Tetrastesbonasia (L.)), due to successfulintroductions) (Haila et al., 1979). The thriving Aland populationsof these species are presumably isolated from themainland populations.

selection, foraging changes in thehabitat Compensatory bird of some sedentary behaviour and even morphology specieson thelargeBalticislandsseemto be connected species. Thisis of potentially competing with theabsence suggestedby abundancedata on large woodpeckers ob1977)anddetailed (Svardson, 1949;Haila & Jarvinen, & tits (Alatalo,Gustafsson servational data on forest Lundberg, 1986). in theAland beetles of carabid Data on thedistribution processesare archipelago suggestthatpopulation-level islands. important for their abundanceson different off the a fewkilometres Population sizes on smallislands derivedfrom fromexpectations coast deviate greatly similarsamples in similarhabitatson the mainland of carabid samples (Niemelaet al., 1985). A comparison three c. 0.5 km2 islands 5-15 kmoffthecoastwith from from themainland revealed and withsamples each other etal., 1988).A few differences (Niemela more pronounced melanarius onthemainland abundant (Pterostichus species werecompletely versicolor (Sturm)) (Ill.) andPterostichus from on theislands. Sevenspecies absent similar habitats among the islands.being had a 'mosaic' distribution butabsent from similar on one or twoof them abundant intheother(s). somespecies (particuFurthermore, habitats in on theislands (Gz.)) werefound larly Calathus fuscipes on the thanon similarhabitats muchhigher numbers a compensatory population-level suggests mainland, which response. on The patterns thatlocal carabid populations indicate thecoastmaybe dynamically off islands a fewkilometres in the boreal populations independent frommainland conditions of theBaltic.Mostof themainland speciesare colonization successis able to colonizeislands, buttheir on Unfortunately, from speciesto species. highly variable to infer the basis of distributional data it is impossible the islandpopulations show real immigrationwhether intime. extinction dynamics thepersistence dynamics control Immigration-extinction in lakesin eastern on smallislands of shrew populations Finland (Hanski, 1986). The abundantand socially common shrew araneus dominant L.), is consistent(Sorex on islands 2 ha,butdemographic than stolarger lypresent of small'populations chasticity maycauselocalextinctions <2 ha. The smaller shrew on islands masked (S. caecutiens L.), are Laxmann),and pygmyshrew (S. minutus shrew to envimore sensitive than thecommon apparently ronmental variation. distributions Theyhavemorepatchy andareat anysingle moment from absent many relatively The immigrationlarge islandswith suitablehabitats. extinction on theseislandsare presumably not dynamics ofmainland populations. independent in archipeladistributions of smallmammals Dynamic connected withpopulation on fluctuations goes are often themainland demonstrated 1986).Thisis amply (Hanski, inpopulation agrestis ofthefield vole(Microtus dynamics (Pokki,1981). The southern Finland (L.)) in Tvarminne, number of islands bv thefield vole varied from inhabited ina 6-year tofifty-two (outofseventyperiod twenty-eight
one islandsincludedin the study).Demographiccharactertheir isticsof thecolonizingsmall mammalsmay influence

564

YrjoHaila compared with mainland ones can be traced to the dynamics of environmentalchanges in the whole on any single island. thanto conditions archipelagorather For instance, thewaterstrider Gerristhoracicus Schumm.is in southern Finland particularly adapted to the unpredicof rock pool 'archipelagoes' of Baltic table environment skerries(but not to any single rock pool) (Vepsalainen, 1974; Jarvinen& Vepsalainen, 1976; Kaitala, 1987). Another example is provided by carabid beetles. The proportionof brachypterous individuals is high among carabidbeetlesin theAland archipelago comparedwiththe Scandinavian mainland (Lindroth, 1949). This is presumablydue to population processes in the whole rather thanto conditions on singleislands.The archipelago Baltic is a land-uplift area, which produces a continuous as well as in habitatson change in island configurations individual islandsdue to primary succession,whichinduces pronounced faunaldynamics & (Haila et al., 1982; Jarvinen Ranta, 1987). About twenty plant species have taxonomically distinctsubspecies in the outer archipelago of the Baltic, and several species in the taxonomicallyvery complex genera of Hieracium L. and Taraxacum Weber may have undergone considerablegeneticchanges in the area sincethelatestglaciation(Palmgren, 1948). The scales of populationdynamics and populationdifferentiation can usuallybe separated from each other because the respective time-scales differgreatly. In population timeperiodsare measuredby singlererelevant dynamics productivecycles. In population differentiation relevant time periods are longerand highlyvariable fromcase to however. For instance, case. Often the scales interact, continuous immigration-extinction dynamics may bring forth due to founder effect. Such seems to be differentiation the case in shrewpopulationson small islands in eastern Finnish lakes (Hanski,1986). The status of the equilibriumhypothesisis different relativeto population dynamics versuspopulation differentiation. On the population dynamic level immigrationextinction dynamicsare a reasonable concept,which has been documented in several cases, albeit mainly on southern latitudes(Simberloff, 1969; Rey, 1985; Hanski, 1986). However,the relevanceof the conceptis less clear in connection withpopulation differentiation. 2.4. Evolutionary scale The broadestscale in Fig. 1 is constituted by evolutionary processes thatlead to the divergenceof species. The Galapagos and otheroceanic islands are 'islands' on this scale for a greatnumberof taxa. A comparableexample fromnorthern is the freshwater'island' of Lake latitudes Baikal, which hosts numerous endemic species, both vertebrates and invertebrates (Kozhov, 1972). However, endemic formsare scarce on northern islands compared with islands closer to the equator in all taxa (Carlqvist, due to differences in geological 1974). This may be partly history,but there is also real variation in the spatioscale of evolutionary temporal processesin different biogeographicregions.An island needs to be largerand more distant from themainlandin thenorth thanin thetropics to be an 'island' on theevolutionary scale.

to persist on islands,as demonstrated ability experimentally forthebank vole (Clethrionomus glareolus (Schreber))by Ebenhard (1987) in Sweden. For an island to be an island on the populationdynamic need to be fulfilled. scale, two criteria First,the whole life within cycle of the organisms concernedmustbe confined the island. Second, the island populationmustin principle get along for several generations independentlyof populationsin surrounding areas. When the time span of dynamicisolation becomes longer,the island population becomes increasingly also as a genetic unit. independent We enter thenextlevel of insularity. 2.3. Populationscale 2: differentiation Very little is known about island populationsof any organisms in the north in terms of their genetic We do not know whether the populationsof composition. thecarabidCalathusfuscipes,say, are genetically differentiatedon the above-mentioned 0.5 km2 Aland islands from those on the main islands of Aland. The time theyhave been isolatedis fairly of differentibutthepossibility short, ation due to genetic drift,or even to adaptive 'niche variations'(Van Valen, 1965) in theislandenvironments is not excluded. The same guess can be made for the sedentarywoodpecker and tit populations on the large Baltic islandsGotlandand Aland. in the Morphologicalchanges have been demonstrated populationof pygmyshrewon Gotlandcomparedwiththe Swedish mainland. Individuals on Gotland have larger jaws, possiblyas a character displacement connectedwith the absence of common shrew, a larger competitor (Malmquist,1985). Gotlandis an island forthe shrewson thescale of population differentiation. Birds in the boreal zone, being mostlymigratory, are unlikelycandidatesforshowingpopulationdifferentiation on islands. In contrast, further towardthe south a larger of island bird populationsare sedentary, proportion and The wren becomes more important. geneticdifferentiation (Troglodytes troglodytes (L.)) has evolved intoat least six distinctsubspecies in different North-Atlantic archipelaislands goes (Williamson,1981). The large Mediterranean hosttwo endemicbirdspecies,theCorsicannuthatch (Sitta whiteheadi Sharpe) and Cypruswarbler (Sylviamelanothorax Tristram) (Blondel, 1985). Similarly, a variety of small mammalshave differentiated intogenetically distinct forms on theislandsnorth of Britain (Berry,1986). Detailed populationgeneticstudieson invertebrates on northern islandsare scarce. Differentiation, possiblydue to founderprinciple,is highly likely in species that are in their entirely dependent on human transportation dispersal to distantislands, e.g. lumbricidson the Faroe Islands (Enckell et al., 1986; see also Enckell,Bengston& Wiman, 1987). Jarvinen& Ranta -(1987) summarized resultsfrompopulationgeneticstudiesconducted recently on small islandsin thenorthern is Baltic; founder principle important,but some differentiation due to selective pressures has been documented as well (primarilyby Halkka and his coworkers, e.g. Halkka & Halkka, 1974; Halkka & Mikkola,1977). In some cases differentiation of archipelagopopulations

Definition of an island 565 The monographof MacArthur & Wilson (1967) deals with evolutionary processes,the dynamicconcept in this connection being the 'taxon cycle' (Wilson, 1961). However, logical ties betweenthe equilibrium hypothesis conclusionsare loose (Haila & Jarvinen, and evolutionary 1982). Furthermore, empirical evidence to support the on islands would be claim thatspeciationand extinction equilibrialprocesses is only suggestiveat best (Faeth & Connor, 1979). It is doubtfulwhetherthe equilibrium hypothesis is actually applicable on the evolutionary scale. of plant populations (Palmgren, 1948) as differentiation occur on the level of the whole archipelagorather thanon thelevel of singleislands(Jarvinen & Ranta,1987).

ANDCONSERVATION 4. ISLANDS
The theory of islandbiography has been applied to conserof habitat vationproblems mainlyin thecontext fragmentation (Soule, 1986). Insular habitatpatches produced by are oftenat the interface betweenthe first fragmentation two island scales. The space isolatinghabitat patchesfrom as each otheris usuallynotequallyhostileto theorganisms wateris formostterrestrial organisms; this is the case for carabid beetles in the taiga (Niemela & Haila, 1986). The to consequence is thatdispersalrates fromone fragment framework anotherincreasewhich makes the equilibrium inadequate.It is necessaryto investigate populationdistributions and habitat requirements on the regional scale (Haila & Hanski,1984; Haila et al., 1989). A crucial conservation challengeconnectedwithhabitat fragmentationis the problem of minimum viable populationsize (Gilpin & Soule, 1986). In the framework of Fig. 1, the problemis mainlyconnectedwith the two largestscales, that is, with situationswhere an endemic species or population is endangeredbecause of habitat destruction. Critical additional effectscan be produced of the population. Lande throughthe spatial structure model of Levins (1969) (1987) applied the metapopulation to territorial organismssuch as birds and concluded that increasingdistance between suitable territories, possibly combinedwitha decreasein their averagesuitability, might drive populations to extinction.In such a case, the is only on the individual 'insularity'of the environment level buta population level consequencefollows. However, the equilibrium hypothesisdoes not give detailed guidelines for elaboratingthe extinction risk of individual species. Even though average extinction in a habitat would be a function of the probability fragment total numberof species presentand fragment area, the the risk of extinctionof specific processes intensifying endangeredspecies (the 'extinctionvortices', Gilpin & Soule, 1986) need studyin each case. Habitatfragmentation is likely to increase extinction risk,but the specific mechanisms must be separately assessed (Soule & Simberloff, 1986).

3. INTERFACES
is It seems thatthe domain of the equilibrium hypothesis scale. restricted mainlyto the second,population dynamics On this scale, empirical evidence on the realism of immigration-extinction dynamics has been obtained in insular species assemblages. In contrast, as regards individual scale islands the equilibrium hypothesis is irrelevant, and forthetwo broadestscales it is an empirical to decide whether matter immigration-extinction dynamics in natural At present are real and relevant conditions. direct positiveevidenceis lacking. However,the scales grade fromone to another. Various combinations of differentecological processes may influence and communities populations on islandsand, as a consequence, the definitionof an 'island' relative to ecological processesis often ontologically ambiguous.This implies thatthe equilibrium hypothesis may be a fruitful of insularsituations; starting pointforstudiesin a variety thisconclusionagreeswiththeview of,e.g., Brown(1986) and Lomolino (1986) on the relevance of the theoryfor studiesof mammalson islands. A particularly interface is between'islands' on important the individualand populationdynamicsscales. Single cow are 'islands' fordungbeetles(Hanski, 1980) and droppings so are carcasses for carrionflies (Hanski, 1976) on the individual scale. The developmentcycle of individual beetles or fliesis confined to single resourcepatches,but a 'population'onlyfora partof one singlepatchessupport generation. However, distancesof the patches fromeach affect other beetlesand flieson thepopulation level. greatly When distancesbetweenthe patches increase it becomes moreand moredifficult fordispersing individuals to finda new suitable patch, and a population level response follows:thepopulation goes extinct (Hanski,1982). Population size of colonizing species in source area variable (mainlandor nearbylarger islands) is an important in islandecology as has been demonstrated for particularly mammals (Lomolino, 1986; Peltonen et al., 1989). fromlarge thanfromsmall Propagulesare more frequent source populations, which increases the persistenceof island populations,and decreases the likelihood of their geneticdifferentiation. The land uplift archipelagoes of the northern Baltic exhibitmixedpatterns as well. Vegetation of the islands is characterized successionalchange,and both by directional, than plants and animals have far higher immigration extinction rateson singleislands.Population such responses

A RESEARCH 5. CONCLUSIONS: PROGRAMME FOR ISLAND BIOGEOGRAPHY

A particular patchof land is an 'island' in a different sense fordifferent in the species. This causes seriousrestrictions domainof validity of equilibrium islandbiogeography. It is usually not ecologically realisticto include all different on an island in a single species presentsimultaneously variable'species number'. Immigration-extinction dynamics are often real with to individual on islands,buttheir respect populations extrapolation to the community level may be invalid. This is because all species in any given taxon do not necessarily

566 Yrjo Haila participate in theimmigration-extinction dynamics. As a consequence, thedynamics maynotbe reflected on the community levelatall. Carabidbeetleson small Baltic islandsserve as an ACKNOWLEDGMENT
Ilkka Hanski, Olli Jarvinen and Kari Vepsalainen made fruitful comments on an earlydraft of thepaper. REFERENCES
Ahl6n,I. & Nilsson, S.G. (1982) Samband mellanfagelfauna och biotopareal pa oar med naturskog i Mdlarenoch Hjalmaren.Var Fagelvarld,41, 161-184. Alatalo, R.V., Gustafsson, L. & Lundberg, A. (1986) Interspecific competition and nichechangesin tits(Parus spp.): evaluationof nonexperimental data.Amer.Natur.127, 819-834. Barthes,R. (1968) Elementsof semiology.Hill and Wang, New York. Baur, B. & Bengtsson, J. (1987) Colonizing abilityin land snails on Baltic uplift achipelagos.J. Biogeogr. 14, 329-341. Berry,R.J. (1986) Genetics of insularpopulationsof mammals, withparticular reference to differentiation and founder effects in British smallmammals. Biol. J. Linn.Soc. 28, 205-130. Birks,H.J.B. (1986) Late-Quaternary biotic changes in terrestrial lacustrine environments, withparticular reference to north-west Europe. Handbook of Holocene palaeoecology and palaeohydrology (ed. by B. E. Berglund), pp. 3-65. JohnWiley & Sons, New York. Blondel, J. (1985) Habitat selection in island versus mainland birds. Habitat selection in birds (ed. by M. L. Cody), pp. 477-516. Academic Press,London. Brown, J.H. (1986) Two decades of interaction between the MacArthur-Wilson model and the complexitiesof mammalian distributions. Biol. J.Linn.Soc. 28, 231-251. Brown,J.H. & Kodric-Brown, A. (1977) Turnover ratesin insular effect of immigration on extinction. biogeography: Ecology,58, 445-449. Carlquist,S. (1974) Island biology. Columbia University Press, New York. Delcourt,R.H., Delcourt,P.A. & Webb, T. (1983) Dynamicplant of vegetational ecology: thespectrum change in space and time. Quat. Sci. Rev. 1, 153-175. Dyke, C. (1988) The evolutionary A dynamics ofcomplexsystems: OxfordUniversity study ofbio-social complexity. Press. Ebenhard, T. (1987) An experimental test of the island colonization survival model: bank vole (Clethrionomys glareolus) populations with different demographicparameter values.J. Biogeogr. 14, 213-223. Enckell, P.H., Bengtson,S.A., Douwes, P., Niklasson,M., Stille, B. & Wiman, B. (1986) The dispersal pattern of an anthropochorous species: Genetic variationin populationsof Lumbricus terrestris L. (Lumbricidae) in the Faroe Islands. Hereditas,104, 253-261. Enckell, P.H., Bengtson,S.-A. & Wiman, B. (1987) Serf and waif colonization: distribution and dispersal of invertebrate species in Faroe Island settlementareas. J. Biogeogr. 14, 89-104. and relaxing Faeth, S.H. & Connor,E.F. (1979) Supersaturated island faunas: a critique of the species-age relationship. J. Biogeogr.6, 311-316. Gilbert, F.S. (1980) The equilibrium theory of island factor fiction? J. Biogeogr.7, 209-235. biogeography: Gilpin,M.E. & Soul6, M.E. (1986) Minimumviable populations: processes of species extinction.Conservation biology. The science of scarcity and diversity (ed. by M. E. Soul6), pp. 19-34. Sinauer,Sunderland, Mass. islands: a sampling Haila, Y. (1983a) Land birds on northern forinsular metaphor colonization. Oikos,41, 334-351.

shows a regular increasewithisland area, but this is strongly connected with habitatheterogeneity of the islands.A number of species are good colonizers of islands andseemto be virtually always present on islands close to the shore,some of thempresumably being specialistsof shore-linehabitats.Consequently, the speciesthatpossiblyhave a dynamic balancebetween immigration and extinction in theareaare confounded in thetotalspecieslists;their influence on thetotalspecies number oftheislands is negligible. The sameis true of carabid beetles in terrestrial habitat in southern fragments Finland.Distributional data on forest in smallforest carabids patches surrounded by agricultural and suburban in thevicinity habitats of Helsinki (Halme, 1987) suggest thata few forest species have in establishing difficulties viable populations in the patches. their However, community level contribution is confounded by generalist speciesand speciesfavoured by human influence. The equilibrium modeldoes notprovide the unifying scheme forinsular conceptual studies it purports to be. I that theconstruction suggest ofa new,more realistic view of insular ecology than the one providedby the MacArthur-Wilson framework include twomain steps: First, outinwhich finding particular sensethe ecological islandsstudiedare 'islands' for the organisms one is in. interested Second,developing modelsthatreflect the particular areimportant inthesystem that dynamics studied. Different types of questionsneed to be asked in - or,to putit differently, different situations theobserved needto be related patterns to different 'contrast spaces' which defineexcluded alternatives (Dyke, 1988). In smaller island scalesan appropriate 'contrast space' would be basedon variables colonization influencing propensity ofindividual species & (e.g.Simberloff, 1981;Vepsalainen Pisarski, 1982;Haila et al ., 1983),and in broader island scalesonvariables their influencing long-term persistence. Whatever thedetailed of theensuing structure models, is nottheonlyoption idiosyncracy that follows from the oftheMacArthur-Wilson framework. Island disintegration studiesdiversify, but lower-level and generalizations models can be developed coverspecified that andecologirealistic subsets ofall possible kinds ofarchipelagoes. cally Generalism andrealism can be combined in various ways inconstructing models ecological (Levins, 1966). Williamson(1989) evaluated the legacy of the 'true buttrivial'. equilibrium theory by thestatement My conclusion is different. The domainof validity of the is more narrow thanoriginally equilibrium hypothesis - in many situations inthehypothesis is simply supposed whenthescale of islands matches appropriate. However, of the hypothesis and relevant implicitassumptions
immigration-extinction dynamics can be demonstrated, the hypothesis is highly non-trivial.

example (Niemela et al., 1988). The numberof species

of an iAland 567 Definition

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